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SPECIAL PUBLICATIONS
THE MUSEUM

TEXAS TECH UNIVERSITY

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Biology of Bats of the New World Family
Phyllostomatidae. Part III

Edited by

Robert J. Baker , J. Knox Jones, Jr., and Dilford C. Carter

January 1979

TEXAS TECH UNIVERSITY

Cecil Mackey, President

Regents. — Robert L. Pfluger (Chairman), J. Fred Bucy, Jr., Clint Formby, Roy K. Furr,
A. J. Kemp, Jr., James L. Snyder, Lee Stafford, Judson F. Williams, and Don R. Workman.

Academic Publications Policy Committee. — J. Knox Jones, Jr. (Chairperson), Dilford C.
Carter (Executive Director and Managing Editor), Robert J. Baker, David K. Davies, Harold
E. Dregne, Leslie C. Drew, Charles S. Hardwick, Ray C. Janeway, Walter R. McDonald,
George F. Meenaghan, Charles W. Sargent, and J. Dalton Tarwater.

The Museum

Special Publications No. 16
441 pp.

12 January 1979

$20.00

Special Publications of The Museum are numbered separately and published on an irregular
basis under the auspices of the Dean of the Graduate School and Director of Academic Pub¬
lications, and in cooperation with the International Center for Arid and Semi-Arid Land
Studies. Copies may be obtained on an exchange basis from, or purchased through, the Ex¬
change Librarian, Texas Tech University, Lubbock, Texas 79409.

ISSN 0149-1768
ISBN 0-89672-068-3

Texas Tech Press, Lubbock, Texas

1979

SPECIAL PUBLICATIONS
THE MUSEUM
TEXAS TECH UNIVERSITY

Biology of Bats of the New World Family
Phyllostomatidae. Part III

Edited by

Robert J. Baker, J. Knox Jones, Jr., and Dilford C. Carter

No. 16

January 1979

TEXAS TECH UNIVERSITY

Cecil Mackey, President

Regents. — Robert L. Pfluger (Chairman), J. Fred Bucy, Jr., Clint Formby, Roy K. Furr,
A. J. Kemp, Jr., James L. Snyder, Lee Stafford, Judson F. Williams, and Don R. Workman.

The Museum

Special Publications No. 16
441 pp.

12 January 1979
$20.00

ISSN 0149-1768
ISBN 0-89672-068-3

Texas Tech Press, Lubbock, Texas

1979

CONTENTS

Introduction . 5

Systematic and Distributional Notes . 7

J. Knox Jones, Jr., and Dilford C. Carter, The Museum, Texas Tech
University, Lubbock, 79409.

Morphometrics . 13

Pierre Swanepoel and Hugh H. Genoways, Kaffrarian Museum, King

William’s Town, 5600, Republic of South Africa; Carnegie Museum of
Natural History, 4400 Forbes Avenue, Pittsburgh, Pennsylvania
15213.

Karyology . 107

Robert J. Baker, Department of Biological Sciences and The Museum,
Texas Tech University, Lubbock, 79409.

Biochemical Genetics . . 157

Donald O. Straney, Michael H. Smith, Ira F. Greenbaum, and Robert J.
Baker, Museum of Vertebrate Zoology, University of California, Berkeley,
94720; Savannah River Ecology Laboratory, Aiken, South Carolina
29801; Department of Biology, Texas A&M University, College
Station, 77843; Department of Biological Sciences and The Museum,

Texas Tech University, Lubbock, 79409.

Sperm Morphology . 177

G. Lawrence Forman and Hugh H. Genoways, Department of Biology,
Rockford College, Rockford, Illinois 61101; Carnegie Museum of
Natural History, 4400 Forbes Avenue, Pittsburgh, Pennsylvania 15213.

Alimentary Tract . 205

G. Lawrence Forman, Carleton J. Phillips, and C. Stanley Rouk, De¬
partment of Biology, Rockford College, Rockford, Illinois 61101; De¬
partment of Biology, Hofstra University, Hempstead, New York
1 1550; Barton County Community College, Great Bend, Kansas 67530.

Morphometric Analysis of Chiropteran Wings . 229

James Dale Smith and Andrew Starrett, Department of Biological
Sciences, California State University, Fullerton, 92634; Department of
Biological Sciences, California State University, Northridge, 91330.

Reproductive Patterns . 317

Don E. Wilson , U.S. Fish and Wildlife Service, National Fish and
Wildlife Laboratory, National Museum of Natural History,

Washington, D.C. 20560.

Embryology . 379

William J. Bleier, Department of Zoology, North Dakota State
University, Fargo, 58102.

Ontogeny and Maternal Care . 387

D. G. Kleiman and T. M. Davis, National Zoological Park,

Smithsonian Institution, Washington, D.C. 20008.

General Physiology . 403

John M. Burns, Department of Biological Sciences, Texas Tech
University, Lubbock, 79409.

Population and Community Ecology . 409

Stephen R. Humphrey and Frank J. Bonaccorso, The Florida State
Museum, University of Florida, Gainesville, 3261 1; University
College, European Division, University of Maryland, im Bosseldorn
30, 6900 Heidelberg, German Federal Republic.

INTRODUCTION

Because of their adaptive diversity and, in many instances, unique morphologi¬
cal attributes, bats of the family Phyllostomatidae long have fascinated biologists.
Known only from the New World, most species of phyllostomatids are limited
distributionally to tropical environments, but some representatives occur as far
north as the southwestern United States and others southward to the northern parts
of Argentina and Chile; some species also are distributed on the Bahamas and
islands of the Greater and Lesser Antilles. With the advent in recent years of
improved methods of collecting bats, a tremendous wealth of information on
phyllostomatids has accumulated, and it is the purpose of this three-part pub¬
lication, which contains a total of 27 individual chapters, to bring these data
together in order to assess what now is known about the family and to provide a
departure point for future studies.

Owing to the large number of contributions, all of which were solicited by us
from persons we felt to be knowledgeable of the subject matter, and the fact that
several contributions are necessarily lengthy, the decision was made to group
chapters into three volumes, each separately numbered as a Special Publication of
The Museum at Texas Tech University. In order to establish a workable approach
by which reference could be made consistently to taxa throughout the series, an
annotated checklist by Jones and Carter (published in the first part of the trilogy)
was circulated to all authors. Each was asked to follow the nomenclature and
systematic arrangement in the checklist or, alternatively, to document departures
therefrom. This system, it is hoped, will allow readers to relate information from
one chapter to another and from one volume to the next without the handicap of
conflicting names for the same organism.

Manuscripts first were requested from contributors in 1973 and most had
been received by the end of 1974. Part I of the series was published in 1976 and
Part II in 1977. As editorial work progressed, some authors provided up-dated
information and all authors had the opportunity to insert limited materials at
the time they received galley proofs. Therefore, content is as current as reasonably
could be anticipated for a project of this kind. Organization and editorial style
follow that established for the Special Publications of The Museum at Texas
Tech University. Otherwise, authors were allowed broad latitude concerning
material to be included in their chapters. Accordingly, and for obvious other
reasons, some chapters overlap others in content.

Even though some redundancy has resulted, we thought it best to have a section
on the cited literature with each contribution. Citations to manuscripts in Part
III are carried in text as “this volume.”

For the convenience of readers who may not have seen Part I of the series
(Spec. Publ. Mus., Texas Tech Univ., 10:1-218, 1976), the titles, authors, and
pagination of its contents are as follows: Introduction (Baker, Jones, and Carter),
p. 5; Annotated checklist, with keys to subfamilies and genera (Jones and Carter),

SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

pp. 7-38; Zoogeography (Koopman), pp. 39-47; Chiropteran evolution (Smith),
pp. 49-69; Collecting techniques (Tuttle), pp. 71-88; Care in captivity (Green-
hall), pp. 89-131; Economics and conservation (C. Jones), pp. 133-145; Brain
anatomy (McDaniel), pp. 147-200; and Lactation and milk (Jenness and
Studier), pp. 201-218.

Following a two-page introduction by the editors, Part II (Spec. Publ. Mus.,
Texas Tech Univ., 13:1-364, 1977) includes: Endoparasites (Ubelaker,

Specian, and Duszynski), pp. 7-56; Ectoparasites (Webb and Loomis), pp.
57-1 19; Oral biology (Phillips, Grimes, and Forman), pp. 121-246; Echolocation
and communication (Gould), pp. 247-279; Thermoregulation (McManus),
pp. 281-292; Feeding habits (Gardner), pp. 293-350; and Movements and
behavior (Fenton and Kunz), pp. 351-364.

February 1978

Robert J. Baker
J. Knox Jones, Jr.
Dilford C. Carter

SYSTEMATIC AND DISTRIBUTIONAL NOTES

J. Knox Jones, Jr., and Dilford C. Carter

Since completion of the manuscript for an annotated checklist of phyllostomatid
bats, which appeared in the first part of this trilogy (Jones and Carter, 1976),
several publications have come to our attention that alter the systematic arrange¬
ment originally presented or extend the known distribution of included species.
These papers are summarized here for the convenience of those who may not have
all the recent literature available to them and also in order to make the three-
volume set on the biology of the Phyllostomatidae more useful as a source of
references. Some of this new information also is incorporated in an annotated
checklist of the bats of Mexico and Central America by Jones et al. (1977).

Systematics

In a recent appraisal of the taxonomy and zoogeography of Macrotus water-
housii in the West Indies, Buden (1975) reached the conclusion that only two
subspecies should be recognized there: waterhousii (jamaicensis a synonym)
on Jamaica, Hispaniola, and Puerto Rico, and in the southern Bahamas; minor
( compressus a synonym) on Cuba, Grand Cayman, and in the northern Bahamas.
Anderson and Nelson (1965) had recognized four subspecies in the Antillean
segment of the distribution of M. waterhousii.

Greenbaum et al. (1975) convincingly argued, on the basis of karyotypes,
that Mesophylla is generically distinct from Ectophylla, a conclusion earlier
reached on the basis of morphologic comparisons by Starrett and Casebeer
(1968).

We earlier listed the subgenus Xenoctenes to include Micronycteris hirsuta.
Davis (1976) provided evidence for abandoning Xenoctenes as valid and
returned M. hirsuta to the nominate subgenus.

Distributional records listed for Peru by Gardner (1976) were taken into
account in preparation of our checklist, but the publication arrived too late to
insert remarks relating to systematics. Among these, Gardner suggested that all
species of small Tonatia ( hrasiliensis , venezuelae, and minuta ) probably are
conspecific and that Lichonycteris degener may be synonymous with L. obscura.
He also questioned the report of Lonchophylla concava from Peru.

Buden (1976) studied the genus Erophylla systematically and reduced
the then-recognized two species, including a total of six subspecies, to two sub¬
species of a single species, E. sezekorni, as follows: sezekorni ( mariguanensis ,
planifrons, and syops synonyms) from the Bahamas, Cuba, Jamaica, and the
Cayman Islands; bombifrons ( santacristobalensis a synonym) from Hispaniola
and Puerto Rico.

Buden (1977) also reviewed morphological variation in Brachyphylla and
concluded that all extant populations should be referred to the one species B.

SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

cavernarum. Subspecies recognized by Buden were: cavernarum (Puerto Rico,
Virgin Islands, Lesser Antilles south to St. Vincent); minor (Barbados); nana
(Cuba and Grand Cayman); and pumila (Hispaniola and the Caicos Islands in
the southern Bahamas). Verona (1974) earlier arranged all named taxa of
Brachyphylla as subspecies of the single species cavernarum, but gave no reasons
for having done so.

In a paper on activity patterns of bats taken near Iquitos, Peru, Davis and
Dixon (1976) used the names “Artibeus planirostris ” and “ Artibeus fuliginosus ,”
evidently based at least in part on information contained in the unpublished
doctoral dissertation of Donald R. Patten. They also listed Artibeus pumilio
as a distinct species; we referred to pumilio as a subspecies of A. cinereus.
Similarly, Smith and Genoways (1974) used the name combination “ Artibeus
planirostris trinitatis ” in reference to a population on Margarita Island,
Venezuela. They cited Patten’s unpublished dissertation as the basis for recognition
of specific status for planirostris (which we listed as a subspecies of jamaicensis).
We have read Patten’s dissertation and do not believe he intended to apply the
specific name planirostris to jamaicensis- like bats from the Caribbean coastal
area of northern South America and adjacent islands; nevertheless, we deplore
the use of manuscript names and strongly suggest that such information not be
incorporated into the published literature without appropriate documentation.

Handley (1976) provided a valuable annotated checklist of Venezuelan
bats in which there are several departures from the systematic scheme we
employed. Unfortunately, none of these departures is documented with evidence
or other explanation; rather, it is indicated that the author will describe new
taxa and discuss nomenclatural changes in another paper that was “in press”
but which, to our knowledge, has not yet appeared.

Finally, Jones (1978) described a new subspecies of the Artibeus jamaicensis
complex from the Antillean island of St. Vincent ( schwartzi ), and Davis and
Carter (1978) named as new Tonatia evotis, which occupies a distribution from
Chiapas southeastward in the Caribbean versant of Central America to
Honduras within the range earlier ascribed to T. silvicola (note change in
spelling). They also described a new subspecies of the latter (T. s. centralis )
from Honduras, Nicaragua, and Costa Rica, and a second new subspecies
( T. s. occidentalis ) from western Ecuador and Pern, while restricting the dis¬
tribution of the nominate subspecies to the region from Panama into South
America as far as Amazonian Brazil, Bolivia, and Peru.

[Koopman’s (1978) important contribution on systematics and zoogeography
of Peruvian bats was received after our report was in galley proof. It contains
accounts for 71 species of phyllostomatids. Among the important systematic
comments are the following: Mimon koepckeae was regarded as a subspecies of
M. crenulatum ; Choeroniscus inca was synonomized with C. minor, Vampyrops
nigellus was placed as a subspecies of V. lineatus\ Enchisthenes was reduced to
subgeneric status under Artibeus, as has been done by several other authors;
Artibeus glaucus and A. watsoni were regarded as conspecific with A.

BIOLOGY OF THE PHYLLOSTOMATIDAE

cinereus, but A. anderseni was recognized as a distinct species; Diaemus was
considered congeneric with Desmodus. Additionally, Koopman recognized and
defined the species Artibeus fraterculus, A. fuliginosus, and A. planirostris as
distinct from A. jamaicensis — we listed fraterculus and planirostris as subspecies
of A. jamaicensis , and fuliginosus represents the “underscribed species” men¬
tioned in the same account. ]

[After this paper was in paged proof, we became aware of a review of the
genus Lonchorina by Hernandez-Camacho and Cadena-G. (Caldesia, 13:1 99-25 1 ,
1978), which included description of a new species, Lonchorhina marinkellei
(p. 229), with type locality at Durania, near Mitu, Colombia.]

Faunistics

Starrett (1976) and LaVal (1977) recorded species of bats, including
phyllostomatids, new to the fauna of Costa Rica. The latter paper contains the
first reported specimen of Micronycteris daviesi from North America under the
generic (instead of subgeneric) designation Barticonycteris. Koopman (1975)
summarized the bat fauna of the Virgin Islands and its zoogeographic relation¬
ships. In a report on bats from southern Haiti, Klingener et al. (1978) recorded
the first whole specimens of Phyllonycteris poeyi obtusa, previously known only
from skeletal remains.

Greenbaum and Jones (1978) reported new records of phyllostomatids
from several Middle American countries and Carter and Jones (1978) recorded
several new species for the Mexican state of Hidalgo, including the northeastern-
most record of Chiroderma villosum. Furthermore, Baker and Genoways
(1978) summarized in a useful way the zoogeography of Antillean bats, and
Baker et al. (1978) reported on bats from the island of Guadeloupe.

In our checklist, we indicated that Vampyrops dorsalis was known from Costa
Rica eastward into South America. Our inclusion of Costa Rica within the
known distribution of this bat evidently was in error as we now can find no
published accounts of this species to the north of Panama. Regarding new
distributional records, Belize and Costa Rica can be added to the countries
previously listed as within the known distribution of Phylloderma stenops,
Michoacan included within the known distribution of Musonycteris harrisoni,
and Oaxaca added to that of Uroderma magnirostrum. Also, Centurio senex
now is known on the mainland of South America from Venezuela.

Readers should be aware of the Mammalian Species series, published by the
American Society of Mammalogists, in which useful summaries of the biology
of individual species of mammals are published. More than 100 accounts thus
far have been distributed or are in press, of which eight of those previously
published deal with phyllostomatids: Ardops nichollsi (Jones and Genoways,
1973), Hylonycteris underwoodi (Jones and Homan, 1974), Macrophyllum
macrophyllum (Harrison, 1975), Macrotus waterhousii (Anderson, 1969),
Monophyllus redmani (Homan and Jones, 1975a), M. plethodon (Homan and
Jones, 19756), Stenoderma rufum (Genoways and Baker, 1972), and Sturnira

SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

thomasi (Jones and Genoways, 1975). Also of interest is a catalogue of type
specimens of bats in European museums that was compiled by Carter and
Dolan (1978). In this work, evidence was presented to establish the correct
spelling of Vampyrodes caraccioli (spelled caraccioloi in our checklist).

Literature Cited

Anderson, S. 1969. Macrotus waterhousii. Mammalian Species, 1: 1-4.

Anderson, S., and C. E. Nelson. 1965. A systematic revision of Macrotus (Chiroptera).
Amer. Mus. Novit., 2212:1-37.

Baker, R. J., and H. H. Genoways. 1978. Zoogeography of Antillean bats. Pp. 53-97,
in Zoogeography in the Caribbean (F. B. Gill, ed.), Spec. Publ. Acad. Nat. Sci.
Philadelphia, 1 3 : iii + 1-128.

Baker, R. J„ H. H. Genoways, and J. C. Patton. 1978. Bats of Guadeloupe. Occas.
Papers Mus., Texas Tech Univ., 50:1-16.

Buden, D. W.. 1975. A taxonomic and zoogeographic appraisal of the big-eared bat

( Macrotus waterhousii Gray) in the West Indies. J. Mamm., 56:758-769.

- . 1976. A review of the bats of the endemic West Indian genus Erophylla. Proc.

Biol. Soc. Washington, 89:1-15.

- . 1977. First records of bats of the genus Brachyphylla from the Caicos Islands,

with notes on geographic variation. J. Mamm., 58:221-225.

Carter, D. C., and P. G. Dolan. 1978. Catalogue of type specimens of Neotropical bats
in selected European museums. Spec. Publ. Mus., Texas Tech Univ., 15:1-136.
Carter, D. C., and J. K. Jones, Jr. 1978. Bats from the Mexican state of Hidalgo. Occas.
Papers Mus., Texas Tech Univ., 54:1-12.

Davis, W. B. 1976. Notes on the bats Saccopteryx canescens Thomas and Micronycteris
hirsuta (Peters). J. Mamm., 57:604-607.

Davis, W. B., and D. C. Carter. 1978. A review of the round-eared bats of the Tonatia
silvicola complex, with descriptions of three new taxa. Occas. Papers Mus.,
Texas Tech Univ., 53:1-12.

Davis, W. B., and J. R. Dixon. 1976. Activity of bats in a small village clearing near
Iquitcs, Peru. J. Mamm., 57:747-749.

Gardner, A. L. 1976. The distributional status of some Peruvian mammals. Occas.

Papers Mus. Zool., Louisiana State Univ., 48:1-18.

Genoways, H. H., and R. J. Baker. 1972. Stenoderma rufum. Mammalian Species,
18:1-4.

Greenbaum, I. F., and J. K. Jones, Jr. 1978. New records of bats from El Salvador,
Honduras, and Nicaragua. Occas. Papers Mus., Texas Tech Univ., 55:1-7.
Greenbaum, I. F., R. J. Baker, and D. E. Wilson. 1975. Evolutionary implications of
the karyotypes of the stenodermine genera Ardops, Ariteus, Phyllops, and
Ectophylla. Bull. S. California Acad. Sci., 74:156-159.

Handley, C. O., Jr. 1976. Mammals of the Smithsonian Venezuelan project. Sci.

Bull. Brigham Young Univ., Biol. Ser., 20(5): (4) + 1-89 + (2).

Harrison, D. L. 1975. Macrophyllum macrophyllum. Mammalian Species, 62:1-3.
Homan, J. A., and J. K. Jones, Jr. 1975«. Monophyllus redmani. Mammalian Species,
57:1-3.

- . 19756. Monophyllus plethodon. Mammalian Species, 58:1-2.

Jones, J. K., Jr. 1978. A new bat of the genus Artibeus from the Lesser Antillean island
of St. Vincent. Occas. Papers Mus., Texas Tech Univ., 51:1-6.

Jones, J. K., Jr., and D. C. Carter. 1976. Annotated checklist, with keys to subfamilies
and genera. Pp. 7-38, in Biology of bats of the New World family Phyllostomatidae.
Part I (R. J. Baker, J. K. Jones, Jr., and D. C. Carter, eds.). Spec. Publ. Mus., Texas
Tech Univ., 10:1-218.

BIOLOGY OF THE PHYLLOSTOMATIDAE

Jones, J. K., Jr., and H. H. Genoways. 1973. Ardops nichollsi. Mammalian Species,
24:1-2.

- . 1975. Sturnira thomasi. Mammalian Species, 68: 1-2.

Jones, J. K., Jr., and J. A. Homan. 1974. Hylonycteris underwoodi. Mammalian
Species, 32:1-2.

Jones, J. K., Jr., P. Swanepoel, and D. C. Carter. 1977. Annotated checklist of the bats
of Mexico and Central America. Occas. Papers Mus., Texas Tech Univ., 47:1-35.
Klingener, D., H. H. Genoways, and R. J. Baker. 1978. Bats from southern Haiti.
Ann. Carnegie Mus., 47:81-99.

Koopman, K. F. 1975. Bats of the Virgin Islands in relation to those of the Greater and
Lesser Antilles. Amer. Mus. Novit., 2581:1-7.

- . 1978. Zoogeography of Peruvian bats with special emphasis on the role of the

Andes. Amer. Mus. Novit., 2651:1-33.

LaVal, R. K. 1977. Notes on some Costa Rican bats. Brenesia (Museo Nacional de
Costa Rica), 10/1 1:77-83.

Smith, J. D., and H. H. Genoways. 1974. Bats of Margarita Island, Venezuela, with
zoogeographic comments. Bull. S. California Acad. Sci., 73:64-79.

Starrett, A. 1976. Comments on bats newly recorded from Costa Rica. Contrib.
Sci., Los Angeles Co. Mus. Nat. Hist., 277:1-5.

Starrett, A., and R. S. Casebeer. 1968. Records of bats from Costa Rica. Contrib.

Sci., Los Angeles Co. Mus. Nat. Hist., 148:1-21.

Varona, L. S. 1974. Catalogo de los mamfferos vivientes y extinguidos de las Antillas.
Acad. Cien. Cuba, viii + 139 pp.

MORPHOMETRICS

Pierre Swanepoel and Hugh H. Genoways

In this paper, we have attempted to cite all relevant literature in which mensural
data pertaining to phyllostomatid bats has appeared. We are not so naive as to
believe this goal was reached, but we do believe most pertinent publications
are listed, including all major works relating to each species. This information
serves as a summary of what currently is known concerning morphometries of
phyllostomatids and hopefully provides a basis for future morphometric studies
of members of the family.

Early descriptive accounts of phyllostomatids were based mostly on material
preserved in fluid and generally lacked mensural data; most measurements that
were included were of external dimensions only. In the late 1800s and 1900s,
cranial measurements began to appear in the literature as did the first systematic
reviews of phyllostomatid groups, notably those dealing with Micronycteris
(Andersen, 1906a), Carollia (Hahn, 1907), Uroderma and Artibeus (Andersen,
1908), and Glossophaga (Miller, 1913 b). Through the years, systematic studies
have become more and more sophisticated, involving substantial mensural
data and complex methods of analysis, culminating in multivariate analyses such
as those of Davis and Baker (1974), Baker et al. (1972a), and Power and
Tamsitt (1973).

In the following accounts, papers in which measurements have appeared
are listed for each species. Additionally, when appropriate information is
available in the published record one or more of the following kinds of variation
are discussed: age, individual, secondary sexual, and geographic. Accounts are
included for all species listed by Jones and Carter (1976). Within each subfamily,
genera and species are listed alphabetically. A standard set of measurements
for specimens of all species of phyllostomatids is given in Appendix 1. One
external (length of forearm) and seven cranial measurements (greatest length
of skull, condylobasal length, zygomatic breadth, postorbital constriction,
breadth of braincase, length of maxillary toothrow, breadth across upper molars)
were taken with dial calipers from each specimen. Four males and four females
were measured for each species except in those instances when fewer specimens
were available to us.

Acknowledgments

We are especially grateful to Rina Swanepoel for aiding us in innumerable
ways including typing early drafts of the manuscript, arranging citations, and
reading proof. We also thank Catherine H. Carter and Margaret Popovich
for their help in checking proof and Flora Gibson for clerical assistance.

We acknowledge the following curators for allowing us to measure specimens
in their care: Karl F. Koopman, American Museum of Natural History (AMNH);

SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

Albert Schwartz, private collection (AS); John Edwards Hill, British Museum
(Natural History) (BMNH); Robert Goodwin, Colgate University (COLU);
Jerry R. Choate, Fort Hays State University (FHKS); Robert S. Hoffmann,
University of Kansas (KU); Lan A. Lester, Natural History Museum of Los
Angeles County (LACM); George H. Lowery, Jr., Louisiana State University
(LSU); Randolph L. Peterson, Royal Ontario Museum (ROM); David J.
Schmidly, Texas A&M University (TCWC); Robert J. Baker, Texas Tech
University (TTU); Charles O. Handley, Jr., National Museum of Natural
History (USNM).

Subfamily Phyllostomatinae
Chrotopterus auritus (Peters, 1857)

Measurements of Chrotopterus auritus have been recorded as follows: Peters (1857),
external measurements of the holotype of Chrotopterus auritus; Dobson (1878a),
external measurements of one specimen; Elliot (1904), and Goodwin (1942a), external
and cranial measurements of one specimen; Elliot (1917), external measurements of one
specimen; Anthony (1920), external and cranial measurements of holotype of C. colombianus
(sex unknown) from Colombia; Lima (1926), external measurements of a male from
Brazil; Cunha Vieira (1942), external measurements of four males and a female and cranial
measurements of a male and female from Brazil; Goodwin (1946), external and cranial
measurements of a male from Brazil; Hall and Kelson (1959), cranial measurements of a
male and female from Veracruz; Burt and Stirton (1961), external and cranial measurements
of a specimen from El Salvador; Villa-R. (1967), external and cranial measurements of a
male from Mexico; Rick (1968), forearm and cranial measurements of three males and a
female from Costa Rica; Goodwin (1969), forearm and cranial measurements of two males
(one subadult) from Chiapas; Villa-R. and Villa Cornejo (1969), external and cranial
measurements of a male and two females from Argentina; Taddei (1975a), external
measurements of six specimens and cranial measurements of seven specimens (mean,
se, range) of males and females combined from Brazil.

Individual variation. — Coefficients of variation for external (N= 6, males and females
combined) and cranial measurements (N— 7, males and females combined) of specimens
from Brazil ranged from 1.89 to 5.37 in external measurements and from 0.84 to 4.08
in cranial measurements (Taddei, 1975a).

Lonchorhina aurita Tomes, 1863

Measurements for Lonchorhina aurita have been recorded as follows: Tomes (1863),
external and cranial measurements of the holotype of L. aurita; Peters (18666), external
measurements of one specimen; Dobson (1878a), external measurements of the holotype
from Trinidad; Elliot (1904), external and cranial measurements of one specimen; Miller
(1912), external and cranial measurements of a male and female from Panama; Anthony
(1923), external and cranial measurements of the male holotype of L. aurita occidentalis
from Ecuador, and forearm measurements of three specimens and cranial measurements
of one specimen of L. aurita aurita from Venezuela; Cunha Vieira (1942), external
measurements of three males and cranial measurements of two males from Brazil; Goodwin
(1942a), external measurements of a specimen from Honduras; Goodwin (1946), external
and cranial measurements of a male and female from Panama; Goodwin (1953), external
and cranial measurements of the holotype of L. a. occidentalis as given by Anthony (1923);
Felten (1956a), external measurements of two males and cranial measurements of a male
from El Salvador; Hall and Kelson (1959), external and cranial measurements of a male
and female from Panama; Burt and Stirton (1961), external measurements of two males

BIOLOGY OF THE PHYLLOSTOMATIDAE

and cranial measurements of a male from El Salvador; Goodwin and Greenhall (1961),
forearm and cranial measurements of two females and a juvenile male from Trinidad;
Pirlot (1967), external measurements of one specimen; Villa-R. (1967), external
measurements of 22 and cranial measurements of 21 males and females combined (mean,
sd, and range) from Mexico; Goodwin (1969), forearm and cranial measurements of four
females from Oaxaca; Tuttle (1970), external measurements of a male and two females
from Peru; Linares and Ojasti (1971), external and cranial measurements of 26 specimens
from Trinidad and Venezuela.

Lonchorhina orinocensis Linares and Ojasti, 1971

Linares and Ojasti (1971) gave external and cranial measurements (mean, sd, range)
of five specimens from Venezuela, including the female holotype.

Macrophyllum macrophyllum (Schinz, 1821)

Measurements for Macrophyllum macrophyllum have been recorded as follows: Dobson
(1878a), external measurements of a specimen from Brazil; Cunha Vieira (1942) external
measurements of two females and cranial measurements of one female from Brazil;
Goodwin (1946), external and cranial measurements of a male from Guyana; Felton
(1956a), external measurements (mean, range) of five males and cranial measurements of
three males from El Salvador; Hall and Kelson (1959), external and cranial measurements
of a male from Guyana; Hill and Bown (1963), external and cranial measurements of a
male and female from Ecuador; Davis et al. (1964), external measurements of two males
from Nicaragua; Hill (1964), forearm and cranial measurements of a male from Guyana;
Starrett and Casebeer (1968), forearm and cranial measurements of a male and female
from Costa Rica; Harrison and Pendleton (1974), external and cranial measurements
of nine males and three females from El Salvador; Harrison (1975), forearm and cranial
measurements (range) for the species; Taddei (1975a), external and cranial measurements
(mean, sd, range) of eight males from Brazil.

Individual variation. — Taddei (1975a) gave coefficients of variation for external (0.48-
8.03) and cranial measurements (0.27-3.51) for eight males from Brazil.

Macrotus cal ifornicus Baird, 1858

Measurements of Macrotus californicus have been recorded as follows: Baird (1858),
external measurements of a specimen from California in the original description of M.
californicus; H. Allen (1864), external measurements of eight specimens; H. Allen (1894a,
18946), mean external and cranial measurements of four individuals and external measure¬
ments of another eight specimens; Elliot (1901), external measurements of one specimen;
Elliot (1904), external and cranial measurements of one specimen; Rehn (1904), external
measurements (mean, range) of five topotypes (Imperial Company, California), and
cranial measurements (mean, range) of six specimens; Stephens (1906), external measure¬
ments of one specimen; Grinnell (1918), external and cranial measurements of 18 females
from California; Hall (1946), external measurements of two males and mean and range
of nine females and cranial measurements of a male and a female from Nevada; Anderson
and Nelson (1965), external and cranial measurements (mean, sd, range) of four samples from
throughout the geographic range of the species; Villa-R. (1967), external measurements
(mean, se, range) of five males and eight females and cranial measurements (mean, se,
range) of five males and four females from Mexico; Anderson (1972), external measure¬
ments of a large sample and cranial measurements of one individual from Chihuahua.

Secondary sexual variation. — Anderson and Nelson (1965) reported no secondary
sexual dimorphism in 28 males and 30 females from California.

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Geographic variation. — According to Anderson and Nelson (1965), there is a
geographic uniformity in characters of populations from the southern end of Baja
California north to California, Nevada, and Arizona and then southward through Sonora.
Consequently, they recognized no geographic races within the area that is now considered
to constitute the distribution of M. californicus.

Macrotus waterhousii Gray, 1843

Measurements of Macrotus waterhousii have been recorded as follows: Saussure
(1860c), external measurements of one specimen; Gundlach (1872, 1877), external

measurements of a Cuban specimen; Dobson (1876), external measurements of the
holotype of M. hocourtianus from Guatemala; Dobson (1878a), external measurements
of two specimens; H. Allen (1890a), external measurements of one specimen in the original
description of M. vv. bulleri from Jalisco; H. Allen (1894a), external measurements of one
specimen probably from Jalisco; J. A. Allen (1904), external measurements (mean) of
seven specimens from Tehuantepec, Oaxaca, compared to those of one specimen from
Yautepec, Morelos; Elliot (1904), external and cranial measurements of four specimens;
Rehn (1904), external and cranial measurements of the various subspecies (revision of
the genus); Elliot (1905), range of external and cranial measurements of the different
subspecies; Shamel (1931), external and cranial measurements of the male holotype of
M. w. herberfolium from Providencialis Island and the measurement range of five
specimens ( = M. vv. waterhousii) Hispaniola; Martinez and Villa-R. (1938), external
measurements of three specimens and cranial measurements of two from Morelos;
Martinez and Villa-R. (1940), external and cranial measurements (mean, sd) of samples
of males and females from the Guerrero; Goodwin (1942a), external and cranial measure¬
ments of one specimen; Anderson and Nelson (1965), external and cranial measure¬
ments (mean, sd, range) of 12 samples from throughout the geographic range of the species;
Choate and Birney (1968), cranial measurements of subfossil specimens from Puerto
Rico; Anderson (1969), external measurements for the genus as the two species are
treated conspecifically under M. waterhousii-, Goodwin (1969), forearm and cranial
measurements of three males and two females from Oaxaca; Alvarez and Ramirez-
Pulido (1972), external and cranial measurements (mean, range) of 11 specimens from
Tamaulipas and San Luis Potosi; Silva-Taboada (1974), measurements of fossil
mandibles from Cuba; Buden (19756), external and cranial measurements (mean, sd,
range) of large samples from northern Bahamas, southern Bahamas, Cuba, Hispaniola,
Jamaica, and means of smaller samples from Isle of Pines, Grand Cayman, and Navassa
for sexes combined.

Individual variation. — In specimens from Guerrero, coefficients of variation
(CV) for external measurements varied in males from 1.93 to 11.16 and in females from
1.67 to 8.09; for cranial measurements, in males from 1.36 to 3.08 and in females from
0.65 to 3.90 (Martinez and Villa-R., 1940).

According to Anderson and Nelson (1965), length of skull proved to be the least
variable character, and then in order of increasing variability were the breadth of brain-
case, length of bulla, interorbital breadth, and breadth at canines. External measure¬
ments were generally more variable than cranial measurements. The coefficient of
variation for total length, however, was usually no greater than that of the more variable
cranial measurements.

Buden (19756) showed in West Indian specimens that cranial (except breadth at
canines) and forearm measurements were the least variable measurements, whereas tail
length generally showed extremely high CVs. Forearm and cranial CV values, other than
that of breadth at canines, ranged from 1.03 to 3.58; values for breadth at canines varied
from 2.78 to 4.63. The coefficient of variation values observed in tail length ranged
from 6.19 to 9.13.

BIOLOGY OF THE PHYLLOSTOMATIDAE

Geographic variation. — Anderson and Nelson (1965) noted an increase in size from
northwest to southeast through the range of Macrotus waterhousii. This held true for all
measurements except length of bulla, which increased in size from southeast to northwest.
Specimens from eastern Cuba were larger than those from the western end of the island.
However, samples from different parts of western Cuba and the Isle of Pines did not differ
significantly in size (Anderson and Nelson, 1965). Geographic variation was found within
Hispaniolan samples — those from Haiti averaged larger than those from the Dominican
Republic. Populations on Hispaniola were larger in size than those on Cuba and the
southern Bahamas. Specimens from several northern Bahaman islands were not significantly
different in size but averaged larger than those from Cuba (Anderson and Nelson, 1965)
and smaller than those from the southern Bahamas and Hispaniola. Bats from Jamaica,
according to Anderson and Nelson (1965), were larger than those from Cuba, and inter¬
mediate in size between Cuban and southern Bahaman and Hispaniolan populations (Ander¬
son and Nelson, 1965:21). Specimens from Oaxaca averaged significantly larger than those
from Morelos (region of the type locality) but were not as large as specimens from Hispan¬
iola and the southern Bahamas. Specimens from Oaxaca averaged larger than the western
Cuban specimens. A sample from Morelos, Guerrero, and Puebla were only slightly larger
in cranial size than a sample from Jalisco.

Buden (19756) stated that the statistical data he used were comparable to those of
Anderson and Nelson (1965) but concluded that a dendrogram, based on levels of
morphological differences, placed the northern Bahaman specimens with the Cuban
ones. An increase in specimen size from southwest to northeast throughout the West Indies
(western to eastern Cuba to northern Bahamas; and Jamaica, Hispaniola, to southern
Bahamas) was found. Ear length, however, did not show this pattern (Buden, 19756).
Buden (19756) also described an increase in size from western Cuba to eastern Cuba
as did Anderson and Nelson (1965). However, in contrast to Anderson and Nelson,
Buden did not find intra-island variation on Hispaniola.

Davis and Baker (1974) reported a general trend of size increase on the mainland
from north to south in all measurements. Their multivariate analyses showed that the
groups were nonclinally tied one to another with respect to geography.

Micronycteris behni (Peters, 1865)

Measurements of Micronycteris behni have been recorded as follows: Peters (1865 6),
external measurements of the holotype from Brazil; Dobson (1878a), external measure¬
ments of a specimen; Andersen (1906a), external measurements of two specimens and
cranial measurements of one specimen from Peru; Sanborn (1949a), range of forearm
length in the species.

Micronycteris brachyotis (Dobson, 1878)

Measurements of Micronycteris brachyotis have been recorded as follows: Dobson
(18786), external measurements of the male holotype of M. brachyotis from Cayenne; Miller
(1900c), forearm length for M. brachyotis ; Andersen (1906a), external and cranial measure¬
ments of the holotype of M. brachyotis (after Dobson 18786); Sanborn (1949a), external
and cranial measurements of the holotype and two topotypes of M. platyceps (= M.
brachyotis) and external measurements of four additional specimens from Trinidad; Hall
and Kelson (1959), external and cranial measurements of the holotype of M. platyceps, two
topotypes, and one female; Goodwin and Greenhall (1961), forearm measurements (range)
of 16 specimens from Trinidad, cranial measurements of one male and two females in¬
cluding the holotype of M. platyceps, and a comparison of external and cranial measure¬
ments of a large adult male from Trinidad and the holotype of M. brachyotis from Cayenne;
Davis et al. (1964), external and cranial measurements of a female from Chiapas; Jones
(1966), forearm and cranial measurements of a male from Guatemala; Villa-R. (1967),

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external measurements of one specimen from Oaxaca; Rick (1968), external and cranial
measurements of eight males and one female from Guatemala; Goodwin (1969), forearm
and cranial measurements of a male from Oaxaca; Marinkelle and Cadena (1972), forearm
measurement of one male from Colombia, and external and cranial measurements of one
female from Colombia; Starrett (1976), forearm measurements of a female, male, and
juvenile male from Costa Rica.

Geographic variation. — The holotype of M. brachyotis from Cayenne, an old male
with worn teeth, was larger than a series of specimens from Trinidad but not larger
than a speciment of M. platyceps from Nicaragua (Goodwin and Greenhall, 1961).

Micronycteris (= Barticonycteris) daviesi (Hill, 1964)

Measurements of Micronycteris daviesi have been recorded as follows: Hill (1964),
external and cranial measurements of the female holotype from Guyana; Tuttle (1970),
external measurements of two males and one female from Peru.

Micronycteris hirsuta (Peters, 1869)

Measurements of Micronycteris hirsuta have been recorded as follows: Peters (1869),
external measurements of the holotype; Dobson (1878a), external measurements of one
specimen; Elliot (1904), external measurements of one specimen from Costa Rica;
Andersen (1906a), external measurements of two specimens and cranial measure¬
ments of one from Costa Rica; Sanborn (1932), external and cranial measurements of a
female from Colombia; Goodwin (1946), external and cranial measurements of a male
and female from Costa Rica; Hershkovitz (1949), external and cranial measurements of
two males and one female from northern Colombia; Sanborn (1949a), range of forearm
and greatest length of skull for the species; Hall and Kelson (1959), external and cranial
measurements of a male and female from Costa Rica; Goodwin and Greenhall (1961),
forearm length (range) of 12 specimens, and cranial measurements of three males and
two females from Trinidad; Hill (1964), forearm and cranial measurements of one female
from Guyana; LaVal (1969), external and cranial measurements of a male and female
from Honduras; Gardner et al. (1970), external and cranial measurements of one male
from Costa Rica; Valdez and LaVal (1971), external and cranial measurements of two
males from Nicaragua; Baker et al. (1973), forearm and cranial measurements (mean,
se, range, CV) of two samples, one from Trinidad (four specimens) and the other from
Honduras (one specimen) and Nicaragua (four specimens).

Individual variation. — Coefficients of variation in forearm and cranial measurements
obtained from four specimens from Trinidad revealed little variation (CV, 0.8-2. 3),
whereas one specimen from Honduras and four from Nicaragua combined showed higher
values than those from Trinidad (CV, 1. 2-4.1) (Baker et al., 1973).

Geographic variation. — Valdez and LaVal (1971) recorded this species for the first
time from Nicaragua and showed that the two specimens obtained were smaller than
those from Costa Rica and other countries recorded by Goodwin (1946), Sanborn
(1949a), Goodwin and Greenhall (1961), and Gardner et al. (1970). However, these
Nicaraguan specimens proved to differ little from Honduran specimens (LaVal, 1969).
Forearm and cranial measurements of specimens from Trinidad averaged larger than
those for specimens from Honduras and Nicaragua, but only forearm and greatest length
of skull proved to be significantly different (Baker et al., 1973).

Micronycteris megalotis (Gray, 1842)

Measurements of Micronycteris megalotis have been recorded as follows: Dobson
(1878a), external and cranial measurements of one specimen; Miller (1898), external
measurements for specimens from Nicaragua (including the male holotype of M. m.

BIOLOGY OF THE PHYLLOSTOMATIDAE

microtis), Trinidad (one male), Margarita (one male and female), Colombia (two males and
females), Honduras (two males), Colima (four males and three females), Jalisco (two males
and three females), and Oaxaca (one female); Miller (1900c), forearm length for M.
m. microtis', Robinson and Lyon (1901), external measurements of five males and six
females from Venezuela; Elliot (1904), external and cranial measurements of one specimen
and external measurements of the holotype of M. m. microtis ; Rehn (1904), external
and cranial measurements of the holotype of Macrotus pygmaeus (= Micronycteris
megalot is) and one male from Yucatan; Andersen (1906a), external measurements of
the holotype of M. m. microtis (after Miller 1898), external and cranial measurements
(range) of 30 (18 cranial) specimens from Brazil, Peru, Guyana, Venezuela, Trinidad
and Tobago, and of 10 (nine cranial) specimens from Colombia, Guatemala, Honduras
and Mexico; Lyon (1906), ear measurements of the holotype of M. m. microtis and a
specimen from Venezuela; Lima (1926), external measurements of a male from Brazil;
Goodwin (1934), external measurements of one specimen from Guatemala; Martinez
and Villa-R. (1938), external measurements of one specimen from Morelos; Cunha
Vieira (1942), external measurements of four males and cranial measurements of two
males from Brazil; Goodwin (1942a), forearm and cranial measurements of two specimens
of unknown sex from Honduras; Goodwin (1946), external and cranial measurements of
two males from Costa Rica; Sanborn (1949a), range of forearm length of three subspecies;
Hershkovitz (1949), forearm measurement of one specimen and skull measurements
of another, both from Trinidad; Dalquest (1953a), external measurements of eight males
and 10 females, and cranial measurements of seven males and nine females from San
Luis Potosi; Goodwin (1953), external and cranial measurements of the holotype Macrotus
pygmaeus from Yucatan; Goodwin (1954), external measurements of a specimen from
Tamaulipas; Felten (1956a), external and cranial measurements of two males from El
Salvador; Felten (1956 d), external measurements (mean, range) of specimens from
El Salvador; Goodwin and Greenhall (1961), forearm measurements of three specimens
from Trinidad and three from Tobago (unsexed), and cranial measurements of a male
from Trinidad; Burt and Stirton (1961), range of forearm and cranial measurements of
eight males and five females combined from El Salvador; Husson (1962), external and
cranial measurements of six males and three females from Surinam; Tamsitt and
Valdivieso (1963a), mean and range of external and cranial measurements of three
males and four females combined from Colombia; Valdivieso (1964), mean and range
of external and cranial measurements of specimens from Colombia; Brosset (1965),
external and cranial measurements of two males from Ecuador; Villa-R. (1967), external
measurements of six males and 10 females, and cranial measurements of eight males
and seven females from Mexico; Pirlot (1968), forearm measurement of a male from
Peru; Goodwin (1969), forearm and cranial measurements of four males and five
females from Oaxaca; Gardner et al. (1970), wing and cranial measurements (mean,
range) of six males and one female combined from Costa Rica; Jones et al. (19716),
mean and range of forearm and cranial measurements of three males and five females from
westcentral Nicaragua, of three males and three females from Isla del Maiz Grande, and
of three males and three females from Rio Coco, and forearm and cranial measurements
of one male from Bonanza, Nicaragua, and cranial measurements of the M. m. microtis
| holotype (male) from Greytown, Nicaragua; Watkins et al. (1972), forearm and cranial
measurements of two males and females from Jalisco; Jones et al. (1973), forearm and
cranial measurements of three males from the Yucatan Peninsula; Birney et al. (1974),
forearm and cranial measurements of a female from Yucatan; Smith and Genoways
(1974), forearm and cranial measurements of a male and female from Margarita Island,
Venezuela; Taddei (1975a), external and cranial measurements (mean, se, range, CV)
of males and females combined (N= 10) from Brazil.

Individual variation. — Coefficients of variation for 10 specimens (sexes combined) from
Brazil were given for external and cranial measurements by Taddei (1975a). Cranial

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measurements showed little variation (CV, 0.66 to 3.18), whereas those for external measure¬
ments were more variable (CV, 1.77 to 5.48).

Geographic variation. — Variation in size in M. megalotis between two localities in
Costa Rica (Fila la Maquina, Cordillera Talamaca, 6600 to 8700 feet; Rincon and Tilaran,
below 700 feet) were discussed by Gardner et al. (1970). Those from the higher altitude
proved to be larger than those from the lower. Size differences were particularly evident
in wing dimensions; no difference in ear length was observable (see also Jones et al., 19716).
Although cranial measurements seemed to be more or less equal, specimens from the higher
altitude tended to be larger.

Jones et al. (1971 b) concluded that specimens from westcentral Nicaragua and Isla del
Mafz Grande were, on the average, considerably larger in skull and forearm measurements
than the holotype of M. m. microtis from Greytown, eastern Nicaragua. Specimens from
Rio Coco were intermediate between the two morphological types leading these authors
to suggest that intergradation occurred between them. No difference in ear length was
found. In the original description. Miller (1898) claimed that M. m. microtis was
characterized by much smaller ears. Lyon (1906) presented evidence that the ears of the
holotype were small and not damaged. Forearm measurements of four specimens previously
obtained from Isla del Maiz Grande (G. M. Allen, 1929) were also relatively big according
to J ones et al. (19716).

Micronycteris minuta (Gervais, 1856)

Measurements of Micronycteris minuta have been recorded as follows: Dobson (1878c/),
external measurements of one specimen from Brazil; Thomas (1901c), forearm measure¬
ments of the holotype as given by both Gervais and Dobson; Andersen (1906«), external
measurements of eight specimens (range) and cranial measurements of six specimens (range)
from Brazil; G. M. Allen (1908), external and cranial measurements of one female from
Brazil; Cunha Vieira (1942), external measurements of a male from Brazil; Sanborn (1949c/),
range of forearm length in the species, forearm and cranial measurements of one specimen
from Colombia; Goodwin (1953), external measurements of the female holotype of M.
hypoleuca ( = M. minuta ) from Colombia; Goodwin and Greenhall (1961), range of
forearm length of 12 specimens and cranial measurements of one male and two females
from Trinidad; Linares (1969), external and cranial measurements of a male and female
from Venezuela; Gardner et al. (1970), mean and range of external and cranial measure¬
ments of four specimens (three males, one female) from Costa Rica; Valdez and LaVal
(1971), external and cranial measurements of one male from Nicaragua and the range of
measurements of three males and one female from Costa Rica.

Geographic variation. — According to Sanborn (1949c/), specimens from Brazil appeared
to be larger than specimens from Colombia.

Micronycteris nicefori Sanborn, 1949

Measurements of Micronycteris nicefori have been recorded as follows: Sanborn (1949c/),
external and cranial measurements of the male holotype and the range of measurements
of four paratypes from Colombia; Goodwin and Greenhall (1961), forearm length of the
holotype, the range of this measurement in five specimens from Trinidad, and cranial
measurements of the holotype (male) and a male and female from Trinidad; Hill (1964),
forearm (two males) and cranial measurements of one specimen from Guyana; Baker and
Jones (1975), external and cranial measurements of a female from Nicaragua; Starred
(1976), external and cranial measurements of five males and cranial measurements of one
male from Costa Rica; LaVal (1977), forearm length, greatest length of skull, and weight of
a male from Costa Rica.

Geographic variation. — According to Starred (1976), his specimens from Costa Rica
agreed closely in most measurements with those given by Sanborn (1949c/) for specimens
from Colombia.

BIOLOGY OF THE PHYLLOSTOMATIDAE

Micronycteris pus ilia Sanborn, 1949

Measurements of Micronycteris pusilla have been recorded as follows: Sanborn (1949a),
external and cranial measurements of the male holotype from Brazil; Goodwin (1953),
forearm and cranial measurements of the holotype.

Micronycteris schmidtorum Sanborn, 1935

Measurements of Micronycteris schmidtorum have been recorded as follows: Sanborn
(1935), external and cranial measurements of the holotype and paratype (both males)
from Guatemala; Goodwin (1942a), external and cranial measurements of the holotype
from Guatemala; Sanborn (1949a), range of forearm measurements in the species; Hall
and Kelson (1959), external and cranial measurements of the holotype from Guatemala
and one male; Davis et al. (1964), external and cranial measurements of a male from
Nicaragua; Villa-R. (1967), external and cranial measurements of two specimens from
Yucatan; Starrett and Casebeer (1968), forearm (two males, mean and range of five females)
and cranial measurements (two males, two females) from Guanacaste, Costa Rica; Jones
et al. (1973), forearm and cranial measurements of one juvenile female from the Yucatan
Peninsula; Baker and Jones (1975), external and cranial measurements of a male from
Nicaragua.

Micronycteris sylvestris (Thomas, 1896)

Measurements of Micronycteris sylvestris have been recorded as follows: Thomas
(1896), external and cranial measurements of the male holotype from Costa Rica; Elliot
(1904a), external and cranial measurements of one specimen; Andersen (1906a), external
and cranial measurements of the male holotype from Costa Rica; Goodwin (1946), external
and cranial measurements of the male holotype from Costa Rica; Hall and Kelson (1959),
cranial measurements of the holotype of M. sylvestris and one male; Goodwin and Green-
hall (1961), forearm and cranial measurements (range) of four males from Trinidad and
four males from Veracruz; Villa-R. (1967), external measurements (mean, range) of nine
specimens and cranial measurements (mean, range) of five specimens from Colima and
Jalisco; Goodwin (1969), forearm and cranial measurements of two females from Veracruz;
Linares (1969), external and cranial measurements of a female from Venezuela.

Geographic variation. — Specimens from Trinidad were similar to Mexican and Central
American specimens; however, skulls of the material from Trinidad were relatively shorter
than those from Mexico (Goodwin and Greenhall, 1961).

Mimon bennettii (Gray, 1838)

Measurements of Mimon bennettii have been recorded as follows: Saussure (1860c),
external measurements of one specimen of Vampirus auriculas (= M. bennettii)', Peters
(18666), external measurements of a specimen from Brazil; Dobson (1878a), external measure¬
ments of one specimen; Lima (1926), external measurement of a specimen from Brazil;
Cunha Vieira (1942), external and cranial measurements of a female from Brazil; Dalquest
(1957), external and cranial measurements of one specimen from Brazil; Husson (1962),
external and cranial measurements of two females from Surinam; Hill (1964), forearm and
cranial measurements of a male from Brazil.

Mimon cozumelae Goldman, 1914

Measurements of Mimon cozumelae have been recorded as follows: Goldman (19146),
external and cranial measurements of the holotype from Cozumel Island off the east coast
of Yucatan; Elliot (1917), external and cranial measurements of the holotype; Sanborn
(1941), external measurements of two specimens from Yucatan; Goodwin (1942a, 1946),
external measurements of a male and female from Yucatan; Dalquest (1957), external
and cranial measurement (mean) of 10 specimens from Veracruz; Hall and Kelson (1959),

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forearm and cranial measurements of the holotype of M. cozumelae. Carter et al. (1966),
forearm measurements of a male and female from Chiapas; Villa-R. (1967), external
measurements of one male and one female from Yucatan and one male from Oaxaca, and
cranial measurements of the male and female from Yucatan; Goodwin (1969), forearm
and cranial measurements of five males and five females from Oaxaca; Gardner et al.
(1970), external and cranial measurements of one male from Costa Rica; Valdez and
LaVal (1971), external and cranial measurements of one female and the mean of two males
from Honduras; Marinkelle and Cadena (1972), forearm measurements of one male from
Colombia.

Geographic variation. — According to Gardner et al. (1970), their male from Costa Rica
closely resembled a male from Chiapas in cranial measurements.

Mimon crenulatum (E. Geoffroy St. -Hilaire, 1810)

Measurements of Mimon crenulatum have been recorded as follows: Peters (1866a),
external measurements of a specimen from Brazil; Dobson (1878a), external measure¬
ments of one (A/, longifolium) from Brazil, and a specimen from an unknown locality;
Thomas (1903c), external and cranial measurements of the male holotype of M. c. picatum
from Brazil; Cunha Vieira (1942), external and cranial measurements of two specimens
from Brazil; Sanborn (19496), forearm and cranial measurements of two males from Peru;
Handley (1960), external and cranial measurements of five males and four females from
Brazil, Trinidad, Venezuela, Panama, and Ecuador (including the holotype of M. c. keenani)',
Goodwin and Greenhall (1961), external and cranial measurements of a male from Trinidad;
Husson (1962), external and cranial measurements of two males from Surinam; Hill (1964),
forearm of two males and females and cranial measurements of one male from Guyana;
Jones (1964), external and cranial measurements of a female from Campeche and measure¬
ments available from the holotype of M. c. keenani from Panama; Gardner et al. (1970),
external and cranial measurements (mean, range) of four specimens (two males and
females) from Costa Rica; Gardner and Patton (1972), forearm and cranial measurements
(mean, range) of four males and three females from Peru.

Miimon koepckeae Gardner and Patton, 1972

Gardner and Patton (1972) recorded external and cranial measurements (mean, range)
of two males and one female and the measurements of the female holotype from Peru.

Phylloderma stenops Peters, 1865

Measurements of Phylloderma stenops have been recorded as follows: Peters (18666),
external measurements of one specimen from Cayenne; Dobson (1878a), external measure¬
ments of Guandira cayanensis from Cayenne; Goodwin (1940, 1946, 1953), external and
cranial measurements of the female holotype of P. stenops septentrionalis from Honduras;
Goodwin (1942a), external and cranial measurements of two specimens from Honduras;
Hall and Kelson (1959), external and cranial measurements of the P. septentrionalis holotype
and one female; Husson (1962), external and cranial measurements of the male holotype
from Cayenne; Hill (1964), external and cranial measurements of three females from
Guyana, one male from Brazil, and of the holotype of Guandira cayanensis (= P. stenops );
Carter et al. (1966), external and cranial measurements of a male from Chiapas; Gardner
(1976), external and cranial measurements of a female from Peru; LaVal (1977), forearm
length and weight of a female from Costa Rica.

Phyllostomus discolor (Wagner, 1 843)

Measurements of Phyllostomus discolor have been recorded as follows: Peters (18656)
external measurements of one specimen from Brazil; Dobson (1878a), external measure-

BIOLOGY OF THE PHYLLOSTOMATIDAE

merits of one specimen; Elliot ( 1 905 A; 1917), external and cranial measurements of the
holotype of P. verrucossum from Oaxaca; Miller (1932), forearm (range of five specimens)
and cranial measurements of a specimen from Barro Colorado Island, Canal Zone; Sanborn
(1936), forearm and condylobasal length of skull measurements (range) of specimens from
Brazil (discolor), and from Oaxaca, Veracruz, and Guatemala ( verruscosus)\ Cunha Vieira
(1942), external measurements of a male from Brazil and female from an unknown locality;
Goodwin (1942c/), external and cranial measurements of two males from Honduras; Goodwin
(1946), cranial measurements of two males from Honduras; Dalquest (1951), external
and cranial measurements of two males and one female from Trinidad; Felten (1956a),
external measurements (mean, range) of 185 males and 217 females, and cranial measure¬
ments (mean, range) of 35 males and 39 females from El Salvador; Burt and Stirton (1961),
forearm and cranial measurements (range) of 15 males and 12 females from El Salvador;
Goodwin and Greenhall (1961), forearm measurements (range) of four specimens
(two males and females) and cranial measurements of one female from Trinidad; Davis
and Carter (1962a), forearm and cranial measurements of one male from Costa Rica; Husson
(1962), external and cranial measurements of eight males and two females from Surinam;
Valdivieso and Tamsitt (1962), external measurements (range) of five males and three
females and cranial measurements of two specimens from Colombia; Tamsitt and Valdivieso
(1963a), external measurements (mean, range) of 11 specimens (seven males, four females)
and cranial measurements of one male and female from Colombia; Pirlot (1967), external
measurements of two specimens; Villa-R. (1967), external measurements of 13 specimens
(mean, sd, range) and cranial measurements (mean, sd, range) of 14 specimens from
Mexico; Goodwin (1969), forearm and cranial measurements of six males and three females
from Oaxaca; Power and Tamsitt (1973), forearm and cranial measurements (means) of
males and females from various localities in southern Mexico to South America; Smith and
Genoways (1974), external and cranial measurements of four females (mean, range) and
two males (means) from Margarita Island, Venezuela; Taddei (1975a), external (30 males,
30 females) and cranial measurements (mean, sd, range) of 15 males and females
from Brazil; Gardner (1976), external and cranial measurements of a male from Peru.

Individual variation. — Taddei (1975a) reported coefficient of variation values for external
measurements of Brazilian specimens to vary from 2.38 to 6.51, whereas CVs for cranial
measurements varied from 0.96 to 4.45.

Secondary sexual variation. — Taddei (1975a) found females averaged larger than males
in 17 external measurements and significantly so in three of these, length of ear, digit
Ill-phalanx 2, digit V-phalanx 2. Males averaged larger than females in 15 cranial measure¬
ments and significantly so in five of these, breadth across canines, breadth across molars,
zygomatic width, mastoid breadth, cranial depth. Power and Tamsitt (1973), performing
a manova, showed that males were significantly bigger than females, and a subsequent
discriminant function analysis revealed that mastoid width and zygomatic width contri¬
buted greatly to the separation of the sexes.

Geographic variation. — In forearm and condylobasal length of skull, specimens from
Barro Colorado Island, Canal Zone, were somewhat greater in size than three topotypes
of P. discolor from southern Mexico (Miller, 1932). Dalquest (1951), comparing cranial
measurements of Trinidad specimens with those from Venezuela, found no difference,
whereas forearm length appeared to be slightly less than in specimens from the mainland.
Davis and Carter (1962a) stated that the measurements considered to that time as an
expression of geographic variation were in reality due to individual variation. According
to Husson (1962), external and cranial measurements of Surinam specimens agree well
with those given by Sanborn (1936), Dalquest (1951), and Goodwin and Greenhall (1961)
for specimens from Trinidad and Venezuela. When comparing these data with those from
El Salvador (Felten, 1956a), Husson (1962) concluded that the cranial measurements were
larger in the specimens from El Salvador. Power and Tamsitt (1973) stated that populations

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west of the Andes in southwestern Ecuador, those near or within the Andes mountains in
central Colombia, and those east of the Andes in eastern Colombia were quite similar
and did not warrant subspecific recognition. Smith and Genoways (1974) found external
and cranial measurements of specimens from Margarita Island, Venezuela, comparable
to those given by Sanborn (1936) for specimens from Brazil, Venezuela, and French Guiana,
and by Goodwin and Greenhall (1961) for material from Trinidad.

Phyllostomus elongatus(E. Geoffroy St. -Hilaire, 1810)

Measurements for Phyllostomus elongatus have been recorded as follows: Peters (18656),
external measurements of a specimen from Brazil; Dobson ( 1 878z/), external measurements
of one specimen; Sanborn (1936), forearm and cranial measurements of a female from
Ecuador; Cunha Vieira (1942), external measurements of three males and one female and
cranial measurements of one male from Brazil; Husson (1962), external and cranial
measurements of four males and two females from Surinam; Butterworth and Starrett
(1964), external and cranial measurements of a male from Venezuela; Hill (1964), fore¬
arm measurements of a male and female and cranial measurements of a female from Guyana.

Geographic variation. — Measurements of six specimens from Surinam correspond well
to those given by Sanborn (1951) for specimens from Peru, and by Husson (1962) for
material from Guyana.

Phyllostomus hastatus (Pallas, 1767)

Measurements for Phyllostomus hastatus have been recorded as follows: Dobson
(1878a), external measurements of one specimen; Flower and Lydekker (1891), forearm
length of the species; Jentink (1893), forearm length of a male from Guyana; Robinson
and Lyon (1901), external measurements of five males and eight females from Venezuela;
J. A. Allen (1904), external and cranial measurements (range) of two males and four
females (including the female holotype of P. h. panamensis) from Chiriqui, Panama,
external and cranial measurements of the male holotype of P. h. caurae from Colombia,
and cranial measurements (mean, range) of two specimens from Trinidad and four from
eastern Venezuela; Elliot (1904), external and cranial measurements of one specimen;
G. M. Allen (1908), external measurements of three and cranial measurements of one
specimen from Brazil, and external measurements of five specimens from Costa Rica;
Miller (1912), external and cranial measurements of a male from Panama; Cabrera (1917),
external and cranial measurements of the male holotype of P. h. curaca and the range of
some of these measurements in three females from Ecuador; Lima (1926), external measure¬
ments of a male from Brazil; Cunha Vieira (1942), external measurements of eight males
and three females and cranial measurements of three males from Brazil; Dalquest (1951),
forearm and cranial measurements (mean) of four specimens from Trinidad; Goodwin
(1953), forearm and cranial measurements of the female holotype of P. h. panamensis
from Panama and of the holotype of P. h. caucae from Colombia; Hall and Kelson (1959),
external and cranial measurements of a male and female from Costa Rica; Goodwin and
Greenhall (1961), forearm measurements (range) of five specimens (two males, three
females) and cranial measurements of one female from Trinidad; Husson (1962), external
and cranial measurements of eight males and two females from Surinam; Taddei (1975a),
external measurements (mean, sd, range) of 20 males and 20 females and cranial measure¬
ments (mean, sd, range) of 15 males and 15 females from Brazil.

Individual variation. — Taddei (1975a) gave CV values for external measurements from
1.28 to 6.04 and for cranial measurements from 1.06 to 2.84.

Secondary sexual variation. — In all of the 15 cranial measurements taken by Taddei
(1975a), males proved to be significantly larger than females, this was also the case in eight
of the 17 external measurements.

BIOLOGY OF THE PHYLLOSTOMATIDAE

Geographic variation. — According to J. A. Allen (1904), specimens from Chiriqui,
Panama, were much larger than those from Trinidad and eastern Venezuela. Specimens
from Costa Rica seemed to correspond fairly well with the holotype of P. h. panamensis
from Chiriqui (G. M. Allen, 1908).

Phyllostomus latifolius Thomas, 1901

Measurements for Phyllostomus latifolius have been recorded as follows: Thomas
(1901 h), forearm and cranial measurements of the male holotype and external measure¬
ments of a second male from Guyana; Husson (1962), external and cranial measurements
of six paratypes (four males, two females) from Guyana; Marinkelle and Cadena (1972),
forearm and cranial measurements (means) of five females from Colombia.

Tonatia bidens(Spix, 1823)

Measurements for Tonatia bidens have been recorded as follows: Dobson (1878a),
external measurements of one specimen from Brazil; Lima (1926), external measure¬
ments of a specimen from Brazil; Sanborn (1936), external measurements (range) of three
males and cranial measurements of two males from Brazil; Cunha Vieira (1942), external
and cranial measurements of a female from Brazil; Goodwin (19426), external and cranial
measurements (range) of one male and five females from the Amazon basin, one male from
Venezuela, and two males and six females from Costa Rica; Goodwin (1946); forearm and
cranial measurements of a male and female from Costa Rica; Koopman and Williams
(1951), cranial measurements of the holotype and paratype of Tonatia bidens saurophila
from Jamaica and of one specimen of T. b. bidens from Costa Rica and another from
Guyana; Goodwin (1953), one cranial measurement of the holotype of T. b. saurophila
from Jamaica; Hall and Kelson (1959), forearm and cranial measurements of a male and
female from Costa Rica; Goodwin and Greenhall (1961), forearm and cranial measurements
of one male and one female from Trinidad; Hill (1964), forearm measurements of one male
and two females and cranial measurements of one female from Guyana; Carter et al. (1966),
external and cranial measurements of a female from Guatemala; Pirlot (1967), external
measurements of one specimen; Gardner et al. (1970), forearm and cranial measurements
of a female from Costa Rica; Valdez and LaVal (1971), external and cranial measurements
of one male and four females (mean, range) from Honduras; Gardner (1976), external
and cranial measurements (mean, range) of seven specimens from Peru.

Tonatia brasiliense (Peters, 1866)

Measurements for Tonatia brasiliense have been recorded as follows: Peters (18666),
external measurements of the holotype from Brazil; Dobson (1878«), external measure¬
ments of the holotype from Brazil; Cunha Vieira (1942), external measurements based on
Peters (18666); Goodwin (19426), external and cranial measurements of one male and
one female from Brazil and Peters’ measurements of the holotype; Goodwin and Green-
hall (1961:236), forearm and cranial measurements of the holotype; Gardner (1976),
external and cranial measurements of two males from Peru.

Tonatia carrikeri (J. A. Allen, 1910)

Measurements for Tonatia carrikeri have been recorded as follows: J. A. Allen (1910),
external measurements for the male holotype and five females and cranial measurements
of the holotype from Venezuela; Goodwin (19426), external and cranial measurements
of one male and one female from Venezuela; Goodwin (1953), external and cranial measure¬
ments of the holotype from Venezuela; Husson (1962), external and cranial measurements
of a male from Surinam; Gardner (1976), external and cranial measurements of two females
from Peru.

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Geographic variation. — Husson (1962) noted that a male from Surinam was smaller
than one reported by Goodwin (19426) from Venezuela and that it compared more favorably
with a female from Venezuela.

Tonatia minuta Goodwin, 1942

Measurements of Tonatia minuta have been recorded as follows: Goodwin (19426),
external and cranial measurements of the female holotype of T. nicaraguae from Nicaragua,
and the male holotype of T. minuta and two females from Ecuador; Goodwin (1946),
forearm and cranial measurements of the holotype of T. nicaraguae', Goodwin (1953),
external and cranial measurements of the holotype of T. minuta and T. nicaraguae'. Hall
and Kelson (1959), forearm and cranial measurements of the holotype of T. nicaraguae
and one female; Goodwin and Greenhall (1961), forearm and cranial measurements of a
male, female, and juvenile from Trinidad and the holotype of T. minuta', Davis and Carter
(1962 a), external and cranial measurements of a male and the female holotype of T.
nicaraguae from Nicaragua; Davis et al. (1964), external and cranial measurements of
one female from Panama; LaVal (1969), external and cranial measurements of one male
and the mean of two females from Honduras; Gardner et al. (1970), forearm and cranial
measurements of five males (mean, range) from Costa Rica; Jones et al. (19716), external
and cranial measurements of two males from Nicaragua; Ojasti and Naranjo (1974),
external and cranial measurements of one male from Venezuela.

Geographic variation. — LaVal (1969) noted that the three specimens (one male, two
females) he measured from Honduras were notably larger in some measurements (fore¬
arm, third metacarpal, length of skull) than those reported by Davis and Carter (1962a)
and Davis et al. (1964). According to Gardner et al. (1970), specimens from Costa Rica
were smaller than those reported from Honduras by LaVal (1969) but similar in size
to those reported by Davis and Carter (1962a) and Davis et al. (1964) from Nicaragua and
Panama. Jones et al. (19716) concluded that their specimens from Nicaragua resembled
material reported from Nicaragua by LaVal (1969) and averaged larger than other published
measurements (Goodwin, 19426; Davis and Carter, 1962a; Davis et al., 1964; Gardner
et al., 1970). A male collected in Venezuela was, according to Ojasti and Naranjo (1974),
slightly larger than the average size reported from Eucador (Goodwin 19426), Honduras
(LaVal, 1969), Costa Rica (Gardner et al., 1970), and Nicaragua (Jones et al., 19716).

Tonatia silvicola (D'Orbigny, 1836)

Measurements of Tonatia silvicola have been recorded as follows: Peters (18656),
external measurements of a specimen from Brazil; Dobson (1878a), external measure¬
ments of one specimen from Brazil; Elliot (1904), external and cranial measurements of
one specimen; Thomas (1910), external and cranial measurements of the holotype of
T. s. laephotis; Cabrera (1917), external measurements of a male and a female (T. amblyotis)
from Ecuador; Sanborn (1936), external and cranial measurements (range) of specimens
from Ecuador; Sanborn (1941), forearm and cranial measurements of one female from
Peru, one specimen from British Honduras, four specimens from Bolivia, and the range
of measurements of a series from Ecuador; Cunha Vieira (1942), external and cranial
measurements of a male from Brazil; Goodwin (1942a), forearm and cranial measurements
(range) of the species T. amblyotis (=7. silvicola); Goodwin (19426), external and
cranial measurements (range) of T. amblyotis from Bolivia, Ecuador, Colombia, and
Panama and cranial measurements of one specimen from British Honduras, and for T.
laephotis, external measurements of one male and one female from the lower Amazon,
and range of cranial measurements of 16 specimens from Brazil; Goodwin (1946),
external and cranial measurements (range) of the species; Goodwin (1953), external and
cranial measurements of the holotype of Chrotopterus columbianus ( = T. silvicola)
from Colombia; Husson (1962), external and cranial measurements of one male and two

BIOLOGY OF THE PHYLLOSTOMATIDAE

females from Surinam; Hill (1964), forearm measurements of two males and females
and cranial measurements of one female from Guyana; Jones (1964), external and
cranial measurements of a male from Campeche; Carter et al. (1966), external and cranial
measurements of a female from Guatemala; Villa-R. (1967), external and cranial measure¬
ments (range) of T. s. silvicola from Mexico; Villa-R. and Villa Cornejo (1969), external
measurements of one specimen from Argentina; Jones et al. (1973), forearm and cranial
measurements of a male from Campeche.

Geographic variation. — According to Carter et al. (1966), measurements of a female
from Guatemala approximated those given by Goodwin (19426) for South American
specimens but were slightly larger than those for a British Honduran specimen examined
by Goodwin. Sanborn (1941) noted that forearm and total length of skull of a specimen
from British Honduras were small for the species.

Tonatia venezuelae (Robinson and Lyon, 1901)

Measurements of Tonatia venezuelae have been recorded as follows: Robinson and
Lyon (1901), external measurements for the male holotype and two additional males from
Venezuela and cranial measurements of the holotype; Sanborn (1941), forearm measure¬
ments (range) in the original series; Goodwin (19426), external and cranial measurements
of a male and female from Venezuela (including cranial measurements of the holotype
from Venezuela); Goodwin and Greenhall (1961:236), forearm and cranial measurements
of a paratype; Ojasti and Naranjo (1974), external and cranial measurements of one specimen
from Venezuela.

Trachops cirrhosus(Spix, 1823)

Measurements of Trachops cirrhosus have been recorded as follows: Saussure (1860c),
external measurements of one specimen of Tylostoma mexicana ( = T. cirrhosus)', Peters
(1865c), external measurements of a specimen from Brazil; Dobson (1878u), external
measurements of one female from Bermuda; Elliot (1904), external measurements of one
specimen; Goldman (1925), external and cranial measurements of the female holotype
of T. cirrhosus coffin i from Guatemala; Lima (1926), external measurements of a male
from Brazil; Cunha Vieira (1942), external measurements of three males and three females
and cranial measurements of two females from Brazil; Goodwin (1942u), external and
cranial measurements of two females from Honduras and the holotype of T. c. coffin i
from Guatemala; Goodwin (1946), forearm and cranial measurements of one male from
Colombia; Herskovitz (1949), external and cranial measurements (range) of 20 specimens
(eight males, nine females, three unsexed) from northern Colombia; Felten (1956c/), exter¬
nal and cranial measurements of a male from El Salvador; Felten (19566), forearm and
cranial measurements of the female holotype and two paratypes (a male and female) of
T. c. ehrhardti from Brazil, and range of these measurements in two other subspecies,
coffini (Guatemala, Honduras, El Salvador) and cirrhosus (Colombia); Burt and Stirton
(1961), forearm and cranial measurements (range) of five males and 17 females from El
Salvador; Goodwin and Greenhall (1961), forearm measurements (range) of two males and
one female and cranial measurements of one male and one female from Trinidad; Davis
and Carter (1962 a), forearm and cranial measurements of a female from Costa Rica; Husson
(1962), external and cranial measurements of one male from Surinam; Villa-R. (1967), ex¬
ternal and cranial measurements of five specimens from Mexico; Starrett and Casebeer
(1968), forearm and cranial measurements of two females and means and ranges of four
males from Costa Rica; Goodwin (1969), forearm and cranial measurements of four males
and two females from Oaxaca.

Geographic variation. — Husson (1962), comparing external measurements of one male
from Surinam with 20 specimens from Colombia (Hershkovitz, 1949), concluded that the
Surinam specimen was large. The skull measurements, however, did not differ markedly.

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Davis and Carter (1962a) found measurements of their one female from Costa Rica within
the range of variation reported in this species from Colombia (Hershkovitz, 1949). These
authors also concluded that other published measurements (Goldman, 1925; Felten, 1956c/)
fell within the range of the Colombian series (Hershkovitz, 1949).

Vampyrum spectrum (Linnaeus, 1758)

Measurements of Vampyrum spectrum have been recorded as follows: Dobson (1878a),
external measurements of one specimen; Flower and Lydekker (1891), forearm length for
the species; Elliot (1904), external and cranial measurements of a specimen; Goldman
(1917) and Goodwin (1942a), external and cranial measurements of the male holotype
of V. s. nelson i from Veracruz; Sanborn (1941), external and cranial measurements of one
female from Trinidad; Cunha Vieira (1942), external measurements from Dobson (1878a);
Goodwin (1946), external and cranial measurements of one male from Nicaragua and of
the holotype of V. s. nelsonr, Hall and Kelson (1959), forearm and cranial measurements
of the holotype of V. s. nelsonr, Goodwin and Greenhall (1961), forearm measurements
(one male, one female) and cranial measurements (one male) from Trinidad; Husson
(1962), external and cranial measurements of three males, two females, and two unsexed
specimens from Surinam, one male and one female from Cayenne, and one male from
Guyana; Casebeer et al. (1963), external and cranial measurements of a male from Costa
Rica; Hall and Dalquest (1963), external and cranial measurements of the holotype from
Veracruz; Goodwin (1969), forearm and cranial measurements for two males, one from
Veracruz the other from Nicaragua; Peterson and Kirmse (1969), external and cranial
measurements of a female from Panama; Gardner et al. (1970), external and cranial measure¬
ments of one female from Costa Rica.

Geographic variation. — Casebeer et al. (1963) stated that their measurements corre¬
sponded closely with those given by Goldman (1917) for the male holotype of V. spectrum
nelsoni from Veracruz and were slightly smaller than measurements of specimens from
Trinidad (Goodwin and Greenhall, 1961). Peterson and Kirmse (1969), comparing their
female specimens from Panama with those reported by Husson (1962) from the Guianas,
found their specimen actually larger in most measurements than the mean of specimens
from near the type locality (Surinam).

Subfamily Glossophaginae
Anoura brev irostrum Carter, 1968

Measurements of Anoura brevirostrum have been recorded as follows: Carter (1968),
external and cranial measurements of the female holotype from Peru and (mean and range)
of five specimens (one male, four females) from Peru; Gardner (1976), external and cranial
measurements of a male from Peru.

Anoura caudifer(E. Geoffry St.-Hilaire, 1818)

Measurements of Anoura caudifer have been recorded as follows: Saussure (1860c),
external measurements of one specimen of A. ecaudata ( = A . caudifer ); Peters (1869),
external measurements of the holotype of Anoura wiedii from Brazil; Dobson (1878a),
external measurements of one specimen; Lonnberg (1921), external and cranial measure¬
ments of a male from Ecuador in the original description of A. c. aequatoris-, Lima (1926),
external measurements of a specimen of Lonchoglossa ecaudata ( A . caudifer) from Brazil;
Sanborn (1933), forearm and cranial measurements (range) of 11 specimens from Brazil;
Sanborn (1938), external measurements of two specimens and cranial measurements of one
specimen from Venezuela; Sanborn (1941), forearm measurements (range) of two males
from Venezuela and one male and four females from Brazil combined, and the forearm
measurement of one male from Peru; Cunha Vieira (1942), external measurements of five

BIOLOGY OF THE PHYLLOSTOMATIDAE

males and two females and cranial measurements of two males and two females from
Brazil; Hershkovitz (1949), external and cranial measurements (range) of four males and
one female combined, and these measurements for one young adult from Colombia; Husson
(1962), external and cranial measurements of a female from Surinam; Tamsitt and
Valdivieso (19666), external measurements of a male and female, cranial measurements of
a male from Colombia, and mean, sd, se, and range in measurements of specimens from
Andean and Amazonian populations; Taddei (19756), external measurements of 40 males
and 40 females and cranial measurements of 15 males and 15 females (mean, se, range)
from Brazil.

Individual variation. — In specimens from Brazil, coefficients of variation for external
measurements varied in 40 males from 2.64 to 5.88 and in 40 females from 2.09 to 7.44;
for cranial measurements in 15 males, CV values were from 1.37 to 4.27 and in 15 females
from 1.22 to 3.17 (Taddei, 19756).

Secondary sexual variation. — In material from Brazil, 17 external measurements showed
no secondary sexual differences. However, in three (breadth across canines, zygomatic
breadth, mastoid breadth) of 15 cranial measurements, males proved to be significantly
larger than females (Taddei, 19756).

Geographic variation. — Tamsitt and Valdivieso (19666) found specimens from an Andean
population to be generally larger in external measurements than those from an Amazonian
population — forearm measurements proved to be significantly different. Cranial
measurements were similar between the two populations and no geographic trend was
obvious.

Anoura cultrata Handley, 1960

Measurements of Anoura cultrata have been recorded as follows: Handley (1960),
external and cranial measurements of the female holotype from Panama; Carter et al.
(1966), external and cranial measurements of a male from Costa Rica; Carter (1968),
external and cranial measurements (mean, range) of 15 specimens from Panama and
Costa Rica; Gardner et al. (1970), forearm and cranial measurements (mean, range) of
five specimens (four males, one female) from Costa Rica; LaVal (1977), forearm length
and weight of a specimen from Costa Rica.

Anoura geoffroyi Gray, 1838

Measurements of Anoura geoffroyi have been recorded as follows: Peters (1868), external
measurements of the holotype of A. g. lasiopyga from Mexico; Dobson (1878u), external
measurements of the holotype of Lonchoglossa wiedii from Brazil, external measure¬
ments of the holotype of A. geoffroyi, and those of an immature specimen; Elliot (1904),
external and cranial measurements of one specimen; Anthony (1921), external and cranial
measurements of the female holotype of A. g. antricola from Ecuador; Lima (1926),
external measurements of a male from Brazil; Sanborn (1933), external and cranial measure¬
ments (range) of specimens from Veracruz, Tlaxcala, Jalisco, and El Salvador; Goodwin
(1934), external measurements of one specimen from Guatemala; Sanborn (1936), fore¬
arm and cranial measurements (range) of 1 1 males and two females from Guatemala;
Cunha Vieira (1942), external measurements of a male and three females and cranial
measurements of a male from Brazil; Goodwin (1942 a), external and cranial measurements
of one specimen; Goodwin (1953), external and cranial measurements of the female holotype
of A. g. antricola and the holotype of Glossophaga apolinari from Colombia; Sanborn
(1954), forearm measurements of one male and one female from Venezuela; Felten
(1956a), external measurements of five males and eight females (mean and range), and
cranial measurements of two males and one female from El Salvador; Anderson (1957),
external and cranial measurements (mean, sd, range) of 58 males and 42 females from

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Chiapas and of one specimen from Costa Rica; Baker (1960), external and cranial measure¬
ments of one male from Durango; Burt and Stirton (1961), forearm and cranial measure¬
ments of a specimen from El Salvador; Goodwin and Greenhall (1961), forearm measure¬
ments (range) of 15 males and cranial measurements of one male from Trinidad; Husson
(1962), external and cranial measurements of six males from Surinam and one male from
Cayenne; Valdivieso (1964), external measurements of a specimen from Colombia;
Tamsitt and Valdivieso (1966a), forearm and cranial measurements of one female from
Colombia; Villa-R. (1967), external measurements of 29 males and 10 females and cranial
measurements of 28 males and 10 females (mean, sd, range) from Mexico; Goodwin
(1969), forearm and cranial measurements of three males and four females from Oaxaca;
Spenrath and LaVal (1970), cranial measurements of two males from San Luis Potosf
and of seven males (mean, range) from Chiapas; Matson and Patten (1975), forearm
measurements of seven males (mean, range) and two females, and cranial measurements
of five males (mean, range) and two females from Zacatecas.

Secondary sexual variation. — Anderson (1957) found no significant differences in both
external and cranial measurements between 58 males and 42 females from Chiapas.

Geographic variation. — Anderson (1957) found a significant difference in forearm
length and length of skull between specimens from South America and Chiapas.

Anoura werckleae Starrett, 1969

Starrett (1969) recorded external and cranial measurements of the male holotype and
one female paratype from Costa Rica.

Choeroniscus godmani (Thomas, 1903)

Measurements of Choeroniscus godmani have been recorded as follows: Thomas
(1903a), external and cranial measurements of the male holotype from Guatemala; Elliot
(1904), external and cranial measurements of one specimen; Goodwin (1942a), external
and cranial measurements of the holotype from Guatemala and a male from Honduras;
Goodwin (1946), external and cranial measurements of one male and female from Costa
Rica; Sanborn (1954), forearm and cranial measurements (range) of three males from
Honduras, and two males, two females, and one unsexed specimen from Costa Rica combined;
Hall and Kelson (1959), external and cranial measurements of one male and two females
from Costa Rica; Burt and Stirton (1961), forearm and cranial measurements of one male
and female from El Salvador; Gardner (1962 ft), external and cranial measurements of
a female from Nayarit; Carter et al. (1966), external and cranial measurements of one female
from Veracruz and one from Guatemala; Villa-R. (1967), external and cranial measure¬
ments of one female from Oaxaca; Goodwin (1969), forearm and cranial measurements
of two males (subadult) and one female from Oaxaca; LaVal (1969), forearm and cranial
measurements (mean, range) of six males and six females from scattered localities in
Mexico and Central America; Gardner et al. (1970), forearm and cranial measurements
of one male and three females from Costa Rica.

Secondary sexual variation. — LaVal (1969), in a comparison of six males and six
females from scattered localities in Mexico and Central America, found females to be
generally larger than males. He found no overlap in greatest skull length between the sexes.
The rostrum was larger relative to the braincase in skulls from females.

Gardner et al. (1970) also noted in a collection of four specimens from Costa Rica,
that the skull of the one male was considerably shorter than those of the three females
from Costa Rica.

Sanborn (1954) stated, contrary to the above, that there is no great difference in size
between the sexes.

BIOLOGY OF THE PHYLLOSTOMATIDAE

Choeroniscus inca (Thomas, 1912)

Measurements of Choeroniscus inca have been recorded as follows: Thomas (19126),
external and cranial measurements of the male holotype from Peru; Sanborn (1954),
forearm and cranial measurements of the holotype (after Thomas), external measurements
of one male and two females, and cranial measurements of one male and three females
from Venezuela.

Choeroniscus interinedius(J. A. Allen and Chapman, 1893)

Measurements of Choeroniscus intermedins have been recorded as follows: J. A. Allen
and Chapman (1893), external measurements of the female holotype and two males from
Trinidad; Goodwin (1953), forearm and cranial measurements of the female holotype from
Trinidad; Sanborn (1954), forearm and cranial measurements of the holotype as given
by Goodwin (1953), forearm measurement of the holotype as in the original description,
and forearm length of an additional male from Trinidad; Goodwin and Greenhall (1961),
external and cranial measurements of the female holotype, a male, and a female from
Trinidad; Genoways et al. (1973), external and cranial measurements (mean, se, range)
of 10 males and 26 females from Trinidad.

Individual variation. — Coefficients of variation in external measurements ranged from
2.5 (total length for males) to 25.4 (length of tail vertebrae of females). CV values in cranial
measurements ranged from 1.9 (mastoid breadth for females) to 6.9 (postorbital constriction
for males). Females showed higher coefficients of variation than males in external measure¬
ments and lower values than males in cranial measurements (Genoways et al., 1973).

Secondary sexual variation. — Females proved to be significantly larger than males
in five (greatest length of skull, condylobasal length, mastoid breadth, breadth of brain-
case, length of maxillary toothrow) of 12 measurements tested. In two of the other seven
measurements, males averaged larger than females and in one they were equal (Genoways
etal. , 1973).

Choeroniscus minor (Peters, 1868)

Measurements of Choeroniscus minor have been recorded as follows: Peters (1868),
external measurements of the male holotype from Surinam; Dobson (1878a), external
measurements of one specimen from Surinam; J. A. Allen and Chapman (1893), external
measurements as given by Dobson (1878a); Elliot (1904), external measurements of one
specimen; Lima (1926), external measurements of a male from Brazil; Cunha Vieira (1942),
external and cranial measurements of a female from Brazil; Sanborn (1954), forearm
measurements of three specimens from Peru; Husson (1962), external and cranial measure¬
ments of the male holotype from Surinam; Valdivieso (1964), external and cranial measure¬
ments of one female from Colombia.

Choeroniscus periosus Handley, 1966

Handley (1966a) recorded external and cranial measurements of the female holotype
from Colombia.

Choeronycteris mexicana Tschudi, 1844

Measurements of Choeronycteris mexicana have been recorded as follows: Peters
(1868), external measurements of one specimen from Mexico; Dobson (1878a), external
measurements of a single specimen; J. A. Allen and Chapman (1893), external measure¬
ments as given by Dobson (1878a); Elliot (1904), external measurements of one specimen;
Goodwin (1934, 1942a, 1946), external measurements of a specimen from Guatemala;

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Dalquest (1953a), external and cranial measurements (mean) of four males from San Luis
Potosi; Baker (1956), external and cranial measurements (mean, range) of three males
and 10 females from Coahuila; Hall and Kelson (1959), external and cranial measurements
of a male and female from Morelos; Schaldach and McLaughlin (1960), external and
cranial measurements of two males and six females from Arizona, one female from
Sonora, and four males and a female from Oaxaca (mean, range); Axtell (1962), external
measurements of a male, female, and juvenile, and cranial measurements of the two adults
from Coahuila; Baker and Greer (1962), external and cranial measurements (mean, range)
of six males from Durango; Davis et al. (1964), external and cranial measurements of one
female from Honduras; Villa-R. (1967), external measurements (mean, range) of seven
males and females combined and cranial measurements (mean, range) of six males and
females combined from Mexico; Barbour and Davis (1969), range of forearm length
of the species; Goodwin (1969), forearm and cranial measurements of three males from
Oaxaca; Anderson (1972), external measurements of a specimen from Arizona and cranial
measurements of one from Sinaloa; Findley et al. (1975), external measurements (mean,
range) of 12 females from New Mexico.

Glossophaga alticola Davis, 1944

Measurements of Glossophaga alticola have been recorded as follows: Davis (1944),
external and cranial measurements of the male holotype and a female from Tlaxcala;
Davis and Russell (1952), external and cranial measurements (mean, range) of seven males
and six females from Morelos; Gardner (1962a), a graphic representation (mean, se, range)
of variation in forearm and cranial measurements in the species; Villa-R. (1963), com¬
parison of externa! and cranial measurements as in the original description of Glossophaga
morenoi, G. alticola , and G. commissarisi and external measurements of 19 males and
18 females and cranial measurements of 19 males and 19 females of G. morenoi (mixed
sample of G. alticola and G. commissarisi) from Mexico; Villa-R. (1967), external
measurements (19 males, 18 females) and cranial measurements (19 males, 19 females)
of G. morenoi (mixed sample of G. alticola and G. commissarisi from Mexico); Goodwin
(1969), forearm and cranial measurements of five females and one subadult male from
Oaxaca.

Glossophaga commissarisi Gardner, 1962

Measurements of Glossophaga commissarisi have been recorded as follows: Gardner
(1962a), external and cranial measurements of the male holotype from Chiapas and a
graphic representation (mean, se, range) of variation in forearm and cranial measure¬
ments in the species; Villa-R. (1963), comparison of external and cranial measurements
as in the original description of Glossophaga morenoi, G. alticola, and G. commissarisi,
external measurements of 19 males and 18 females and cranial measurements of 19 males
and 19 females of G. morenoi (mixed sample of G. alticola and G. commissarisi ) from
Mexico; Villa-R. (1967), external measurements (18 males, 19 females) and cranial measure¬
ments (19 males, 19 females) of G. morenoi (mixed sample of G. alticola and G. com¬
missarisi)-, Goodwin (1969), forearm and cranial measurements of a male, female, and three
unsexed specimens from Oaxaca; Jones et al. (1972), forearm and cranial measurements of
three females from Sinaloa.

Glossophaga longirostris Miller, 1898

Measurements of Glossophaga longirostris have been recorded as follows: Miller
(1898), external and cranial measurements of the female holotype from Colombia; Robinson
and Lyon (1901), external measurements and greatest length of skull for nine males and
four females from Venezuela; G. M. Allen (1908), external measurements (range) of ten
specimens from Carriacou, Lesser Antilles; Miller (1913a), external and cranial measure-

BIOLOGY OF THE PHYLLOSTOMATIDAE

ments of the male holotype of G. I. rostrata from Grenada, Lesser Antilles; Miller (1913 /?),
external and cranial measurements of nine males and one female from Venezuela, one
male and one unsexed specimen from Colombia, nine males from Grenada, three males,
two females, and three unsexed specimens from Carriacou, and ten males and ten females
from Curasao; Elliot (1917), external and cranial measurements of the holotype of G.
I. rostrum ; Hershkovitz (1949), external and cranial measurements (range) of five males
and two females combined from Colombia; Husson (1960), forearm measurements (range)
of 21 males and 42 females and cranial measurements (range) in 12 specimens from
Aruba, Curasao, and Bonaire islands; Goodwin and Greenhall (1961), forearm measure¬
ments of 10 females and cranial measurements of four females from Tobago, forearm
measurements of 14 females and cranial measurements of 10 females from Trinidad;
Tamsitt and Valdivieso (1963«) and Valdivieso (1964), external and cranial measurements
of a male and two females from Girardot, Colombia; Smith and Genoways (1974), fore¬
arm and cranial measurements of specimens from Curasao (20 from Miller, 19136),
Margarita Island (9), Venezuela (22), Trinidad (5), Grenada (9), and St. Vincent (10).

Geographic variation. — Smith and Genoways (1974) stated that a comparison of
measurements obtained from specimens from Margarita Island with those of the main¬
land and Antillean islands showed that the material from Margarita Island is well within
the range of variation of the mainland specimens and overlap those obtained from Antillean
material.

Glossophaga soricina (Pallas, 1766)

Measurements of Glossophaga soricina have been recorded as follows: Dobson (1878<v),
external measurements of a female; H. Allen (1895), external measurements of the holotype
of Glossophaga truer, Robinson and Lyon (1901), external measurements and greatest
length of skull of one male and three females from Venezuela; Rehn (1902rtX external and
cranial measurements of the female holotype of G. s. antillarum from Jamaica and one
specimen each from Guyana, Trinidad, and the Bahamas; Cabrera (1903), external
measurements for the species in Chile; Elliot (1904), external measurements of one
specimen from Tres Marias Islands and external and cranial measurements of two additional
specimens; G. M. Allen (1908), forearm measurements of three specimens from Peru;
G. M. Allen (1911), forearm and cranial measurements of a specimen from Jamaica;
Miller (1913 6), external and cranial measurements of nine individuals (eight females,
one male) from Brazil, one female from Guyana, seven (five females, one male, one un¬
sexed) from Venezuela, 10 (five females, five males) from Trinidad, five (two females,
two males, one unsexed) from Colombia, eight (three females, five males) from Moyobamba,
Peru, 1 1 (seven females, four males) from Paraguay, 20 specimens (nine females, 1 1 males)
from the mainland of Mexico, two (one female, one male) from Nicaragua, one male from
Costa Rica, five (three females, two males) from Chiriqui, Panama, 10 (five males, five
females) from Panama, 12 (six females, six males) from Tres Marias Islands, 14 (five females,
nine males) from Balsas, Peru, three (two females, one male) from Charapex, Peru, and
two females from Jamaica; Elliot (1917), external and cranial measurements (range) of
specimens from Nayarit to Panama; Lima (1926), external measurements of a male from
Brazil; Goodwin (1934), external measurements (mean) of five specimens from Guatemala;
Martinez and Villa-R. (1938), external measurements of one specimen and cranial
measurements of four specimens of G. morenoi ( — G. soricina ) from Morelos; Martinez
and Villa-R. (1941), external and cranial measurements (mean, variance, and correlation
between measurements) of 52 males and 25 females from Guerrero; Cunha Vieira (1942),
external measurements of nine males and one of unknown sex and cranial measurements
of three males from Brazil; Goodwin (1942 a), external and cranial measurements of two
specimens from Honduras; Goodwin (1946), external and cranial measurements of two
males from Costa Rica; Hershkovitz (1949), external and cranial measurements of three
females from Colombia; Dalquest (1951), forearm and cranial measurements of one

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specimen (sex unknown) from Trinidad; Davis and Russell (1952), external and cranial
measurements (mean, range) of seven males and 12 females from Morelos (G. s. leachi)',
Dalquest (1953a), external measurements (means) of seven males and 15 females and
cranial measurements (means) of nine males and seven females from San Luis Potosi;
Villa-R. (1953), external and cranial measurements (mean, range) of specimens from Tlaxcala
(1), Districto Federal (15), Morelos (12), and Guerrero (5); de la Torre (1954), external
and cranial measurements of one female from Tamaulipas; de la Torre (1955), forearm
measurements (mean, range) of nine specimens (six males, three females combined) from
Guerrero; Felten (1956a), external measurements (mean, range) of 286 males and 200
females and cranial measurements of 27 males and 38 females from El Salvador; Ryan
(1960), external measurements of two females from Guatemala; Burt and Stirton (1961),
forearm and cranial measurements (range) of 43 males and 32 females from El Salvador;
Goodwin and Greenhall (1961), forearm measurements (range) of 20 specimens and
cranial measurements of three females from Trinidad; Husson (1962), external and cranial
measurements of five males and five females from Surinam; Gardner (1962a), graphic
representation (mean, range, se) of variation in forearm and cranial measurements of the
species; Tamsitt and Valdivieso (1963a), external measurements (mean, range) of 51 specimens
from Colombia; Tamsitt and Valdivieso (19636), external measurements of one male and
one female from Colombia; Villa-R. (1963), comparison of external and cranial measure¬
ments as in the original description of Glossophaga morenoi, G. alticola, and G. commissar is i;
Starrett and de la Torre (1964), forearm measurements of two males and 14 females (mean,
range) from El Salvador, Honduras, Nicaragua, and Costa Rica; Valdivieso (1964),
external measurements (mean, range) of 77 specimens from Colombia; Aellen (1965),
external and cranial measurements of one male and one female from Peru; Villa-R. (1967),
external measurements (mean, se, range) of 70 males and 37 females and cranial measure¬
ments of 56 males and 25 females from Mexico; Pirlot (1968), forearm measurements of
a female from Peru; Goodwin (1969), forearm and cranial measurements of a female from
Peru; Goodwin (1969), forearm and cranial measurements of two females, one subadult
male, and three unsexed individuals from Oaxaca; Anderson (1972), external measurements
of two specimens and cranial measurements of one from Chihuahua; Jones et al. (1972),
forearm and cranial measurements (mean, range) of nine males and one female combined
from Sinaloa; Taddei (19756), external measurements (mean, se, range) of 59 males and 47
females and cranial measurements of 20 males and 20 females from Brazil.

Individual variation. — In specimens from Brazil, coefficients of variation for external
measurements varied in 59 males from 2.00 to 5.60 and in 47 females from 2.10 to 5.26;
and for cranial measurements in 20 males, CVs ranged from 1.75 to 3.44 and in 20 females
from 1.65 to 3.37 (Taddei, 19756).

Secondary sexual variation. — Taddei (19756) found females to be significantly larger
than males in four (head and body length, forearm length, fourth and fifth metacarpal) of
17 external measurements. In the case of cranial measurements, females were significantly
larger in two measurements (length of molar, mandibular toothrow) of 15 but significantly
smaller in five (breadth across canines, zygomatic breadth, braincase breadth, mastoid
breadth, cranial depth).

Hylonycteris underwood i Thomas, 1903

Measurements of Hylonycteris underwoodi have been recorded as follows: Thomas
(1903a), forearm and cranial measurements of the holotype and external measurements
of a second specimen from Costa Rica; Elliot (1904), external and cranial measurements of
one specimen; Goodwin (1942a, 1946), forearm and cranial measurements of the holotype
from Costa Rica; Hall and Kelson (1959), forearm and cranial measurements of the
holotype; Davis and Carter (1962a), external and cranial measurements of the holotype
and two additional specimens (sex unknown) from Costa Rica, one male and four females

BIOLOGY OF THE PHYLLOSTOMATIDAE

from Veracruz, and one male and one female from Oaxaca; Jones (1964), forearm and
cranial measurements of one male and one female from Oaxaca; Villa-R. (1967), external
and cranial measurements of one specimen from Tabasco; Goodwin (1969), forearm and
cranial measurements of two males and two females from Oaxaca; Gardner et al. (1970),
forearm and cranial measurements (mean, range) of 14 males and seven females from
Costa Rica; Phillips and Jones (1971), forearm and cranial measurements (mean, range)
of four males and six females combined, additional measurements of the female holotype of
H. u. minor from Jalisco, comparative measurements of a Veracruz male, and a male and
female from Oaxaca; Jones and Homan (1974), external and cranial measurements as
given by Gardner et al. (1970) and Phillips and Jones (1971).

Secondary sexual variation. — Females averaged larger than males throughout the range
of the species according to Phillips and Jones (1971).

Geographic variation. — Davis and Carter (1962a) noted that specimens from Oaxaca
appeared to be smaller than those from Veracruz and Costa Rica. However, Jones (1964)
found his Oaxacan male specimen to be larger than those previously reported and the
measurements of his female specimens fell among the largest known individuals of the
species.

Specimens from Jalisco and southern Oaxaca (Davis and Carter, 1962a) were included
in a subspecies by Phillips and Jones (1971). They concluded that these specimens were
smaller externally and cranially than H. it. underwoodi from northern Oaxaca, Veracruz,
and Guatemala.

Leptonycteris curasoae Miller, 1900

Measurements of Leptonycteris curasoae have been recorded as follows: Miller (19006),
external and cranial measurements of the male holotype from Curasao; Hoffmeister
(1957), external measurements of the holotype and three male topotypes; Husson (1960),
forearm measurements (range) of 21 specimens and cranial measurements (range) of 13
specimens from Aruba, Curasao, and Bonaire islands; Davis and Carter (19626), external
and cranial measurements of four males and two females combined (mean, range); Pirlot
(1965a), external and cranial measurements of the male holotype of L. c. tarlosti, a male,
and three females from Margarita Island; Marinkelle and Cadena (1972), external and
cranial measurements of two females from Colombia; Smith and Genoways (1974),
forearm and cranial measurements of 12 specimens from Margarita Island, two from
Aruba, five from Curasao, and one from Bonaire.

Geographic variation. — In his study of this genus, Hoffmeister (1957) considered L.
curasoae to be a subspecies of L. nivalis. However, Davis and Carter (19626) in their
review of the genus and subsequent authors have consider L. curasoae a distinct species.
Pirlot (1965a) recognized specimens from Margarita Island as a distinct subspecies, however,
Smith and Genoways (1974), after examining specimens from throughout the range of the
species, considered the subspecific status of the island forms unwarranted.

Leptonycteris nivalis (Saussure, 1860)

Measurements of Leptonycteris nivalis have been recorded as follows: Saussure
(1860c), external measurements of one specimen; Dobson (1878a), external measure¬
ments of one specimen; Miller (19006), external and cranial measurements of a male from
Colima; Elliot (1904), external and cranial measurements of one specimen; Martinez
and Villa-R. (1938), external measurements of one specimen; Martinez and Villa-R.
(1940), external and cranial measurements (mean, sd) of samples of males and females from
Guerrero; Goodwin (1942a, 1946), external and cranial measurements of a male from
Colima; Dalquest (1953a), external and cranial measurements of four males and one
female from San Luis Potosi; de la Torre (1955), forearm measurements of one male from

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Guerrero; Baker (1956), external and cranial measurements of two males and mean and
range of five females from Coahuila; Hoffmeister (1957), cranial measurements of the
holotype of L. n. nivalis (Veracruz), external and cranial measurements (mean) of six
males and eight females combined from Texas, and external measurements (mean) of 11
males and 29 females combined from Nuevo Leon; Stains (1957), external and cranial
measurements of the holotype and mean and range of the holotype and 22 topotypes of
L. n. longala from Coahuila (see also Jones, 1958); Hall and Kelson (1959), external and
cranial measurements (range) of a large series of specimens from Jalisco; Davis and
Carter (19626), external and cranial measurements of three males and seven females (mean,
range); Alvarez (1963), external and cranial measurements of five males and five females
combined (mean, range) from Tamaulipas; Baker and Cockrum (1966), external and
cranial measurements of two females from Sinaloa; Villa-R. (1967), external measure¬
ments of 50 specimens (mean, sd, range) and cranial measurements of 37 (mean, sd, range)
from Mexico; Goodwin (1969), external and cranial measurements of two males and
two females from Morelos, and one female from Veracruz; Barbour and Davis (1969),
range of forearm length in the species; Anderson (1972), external and cranial measurements
of one specimen; Matson and Patten (1975), forearm measurements (mean, range) of
seven males from Zacatecas.

Individual variation. — In specimens from Guerrero, coefficients of variation for
external measurements varied in males from 3.03 to 16.25 and in females from 1.04 to
16.58; CV values for cranial measurements in males ranged from 1.68 to 7.44 and in females
from 1.23 to 5.58 (Martinez and Villa-R., 1940).

Geographic variation. — Hoffmeister (1957) and Davis and Carter (19626) have recently
reviewed this genus. Davis and Carter (19626) gave characteristics by which the currently
recognized species can be distinguished.

Leptonycteris sanborni Hoffmeister, 1957

Measurements of Leptonycteris sanborni have been recorded as follows: Hoffmeister
(1957), external measurements of 22 females and cranial measurements of 21 females
from Arizona, external measurements (mean) of 10 males from Chihuahua, and the mean
of eight males from Colima; Davis and Carter (19626), external and cranial measure¬
ments (mean, range) of five males and five females; Baker and Cockrum (1966), external
and cranial measurements of one specimen from Sinaloa; Villa-R. (1967), external measure¬
ments (jV=51) and cranial measurements (N= 39) (mean, sd, range) of L. yerbabuenae
( = L. sanborni) from Mexico; Genoways and Jones (1968), forearm measurements
(mean) of 28 males from Zacatecas; Barbour and Davis (1969), range of forearm length
of the species; Anderson (1972), external measurements (mean, sd, range) of 24 specimens
from Chihuahua and external and cranial measurements of one specimen from Sonora;
Ramirez-Pulido and Alvarez (1972), external and cranial measurements of a lectotype and
external measurements of a male and female paralectotype of L. yerbabuenae ; Jones and
Bleier (1974), forearm and cranial measurements of one male from El Salvador; Matson
and Patten (1975), forearm and cranial measurements of five males (mean, range) and one
female from Zacatecas.

Geographic variation. — The species was originally described as a subspecies of L. nivalis
by Hoffmeister (1957). Davis and Carter (19626) demonstrated characteristics by which this
taxon could be distinguished from L. nivalis. Considerable controversy exists in the litera¬
ture over the relationships of this taxon and L. yerbabuenae. Because the holotype of
yerbabuenae has been lost and because the original series was a composite, Watkins et al.
(1972) considered yerbabuenae to be a nomen dubium. However, as recently as Ramirez-
Pulido and Alvarez (1972), authors have believed that the name yerbabuenae superceded
sanborni. The reader is warned to take great care in using measurements recorded in the
earlier literature concerning this genus because considerable confusion has existed in the
proper identification of the species.

BIOLOGY OF THE PHYLLOSTOMATIDAE

Lichonycteris degener Miller, 1931

Miller (1931) gave external and cranial measurements of the female holotype from Brazil.

Lichonycteris obscura Thomas, 1895

Measurements of Lichonycteris obscura have been recorded as follows: Thomas (1895),
external and cranial measurements of the female holotype from Nicaragua; Elliot (1904),
external and cranial measurements of one specimen from Nicaragua; Sanborn (1936),
external and cranial measurements of a female from Costa Rica and the holotype from
Nicaragua; Goodwin (1942, 1946), external and cranial measurements of two females from
Costa Rica; Hall and Kelson (1959), external and cranial measurements of two females
from Costa Rica; Husson (1962), external and cranial measurements of a male from Costa
Rica; Davis et at. (1964), external and cranial measurements of a female from Nicaragua;
Carter et al. (1966), external and cranial measurements of three females from Guatemala;
Gardner et al. (1970), external and cranial measurements (mean, range) of one male and
three females from Costa Rica; Jones et al. (19716), external and cranial measurements of
three males from Nicaragua; Marinkelle and Cadena (1972), forearm measurements (range)
of three females and one unsexed specimen from Colombia; Gardner (1976), external
and cranial measurements of two females from Peru.

Lionycteris spurrelli Thomas, 1913

Measurements of Lionycteris spurrelli have been recorded as follows: Thomas (1913),
external and cranial measurements of the immature male holotype from Colombia; Gold¬
man (19146), greatest length of skull of a specimen from Colombia; Sanborn (1941),
external measurements of one male and one female and cranial measurements of one
specimen from Guyana, and the measurements for the holotype from Colombia.

Lonchophylla concava Goldman, 1914

Measurements of Lonchophylla concava have been recorded as follows: Goldman
(1914a), external and cranial measurements of the male holotype from Panama; Elliot
(1917), external and cranial measurements of the holotype; Goodwin (1946), external
and cranial measurements of the holotype from Panama; Hall and Kelson (1959), external
and cranial measurements of the holotype of L. concava', Davis et al. (1964), external and
cranial measurements of one male and two females from Costa Rica; Pirlot (1968), forearm
measurements of one male from Peru; Gardner et al. (1970), external and cranial measure¬
ments (mean, range) of five specimens from Costa Rica; Marinkelle and Cadena (1972),
forearm measurements of two females from Colombia.

Lonchophylla hesperiaG. M. Allen, 1908

Measurements of Lonchophylla hesperia have been recorded as follows: G. M. Allen
(1908), external and cranial measurements of the male holotype and two additional
specimens from Peru; Gardner (1976), external and cranial measurements of one male
and one female from Peru.

Lonchophylla mordax Thomas, 1903

Measurements of Lonchophylla mordax have been recorded as follows: Thomas
(1903c), external and cranial measurements of the male holotype from Brazil; G. M. Allen
(1908), external and cranial measurements of the holotype from Brazil; Lima (1926),
external measurements of a specimen from Brazil; Sanborn (1941), forearm measurements
(range) of 18 males from Brazil; Cunha Vieira (1942), external measurements of a male
and a female and cranial measurements of a male from Brazil; Baker (1974), external and
cranial measurements of one female from Ecuador.

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Lonchophylla robusta Miller, 1912

Measurements of Lonchyphylla robusta have been recorded as follows: Miller (1912)
and Goodwin (1946), external and cranial measurements of the male holotype and a female
from Panama; Hall and Kelson (1959), external and cranial measurements of the holotype
and a female topotype; Walton (1963), external and cranial measurements (mean, sd, se,
range) of specimens from Panama (N=21) and Costa Rica (A=10); Valdivieso (1964),
external and cranial measurements of one female from Colombia; Tuttle (1970), external
measurements of one male and one female and cranial measurements of the female from
Peru; Gardner (1976), external and cranial measurements of a male from Peru.

Secondary sexual variation. — According to Walton (1963), no sexual dimorphism in size
was evident in specimens from Panama and Costa Rica.

Geographic variation. — Walton (1963) found specimens from Panama to be larger than
those from Costa Rica. Of seven external measurements, three (total length, length of hind
foot, ear length) proved to be significantly different, whereas in nine cranial measurements
there were four (skull length, skull width, interorbital width, width of rostrum at canines)
that showed significant differences.

Lonchophylla thomasiJ. A. Allen, 1904

Measurements for Lonchophylla thomasi have been recorded as follows: J. A. Allen
(1904), external and cranial measurements of the male holotype from Venezuela; Goodwin
(1953), forearm and cranial measurements of the holotype; Husson (1962), external and
cranial measurements of two males and one female from Surinam; Hill (1964), forearm
and cranial measurements of a male from Guyana; Gardner (1976), external and cranial
measurements (mean, range) of six males and six females combined from Peru.

Monophyllus plethodon Miller, 1900

Measurements of Monophyllus plethodon have been recorded as follows: Miller (1900a),
external and cranial measurements of the male holotype of M. plethodon from Barbados,
Lesser Antilles; Miller (1902a), external and cranial measurements of the male holotype
of M. p. luciae from St. Lucia, Lesser Antilles, and of the holotype of M. p. plethodon ; Elliot
(1904), external and cranial measurements of a specimen from Barbados and one from St.
Lucia; Anthony (1917), cranial measurements of the holotype and two additional specimens
(sub-Recent fossils) of M. frater from Puerto Rico; Anthony (1918, 1925), cranial measure¬
ments of three specimens of sub-Recent fossils from Puerto Rico; Goodwin (1953), a cranial
measurement of the holotype of M. p. frater (subfossil) from Puerto Rico; Hall and Kelson
(1959), external and cranial measurements of the holotype of M. plethodon, M. luciae,
and cranial measurements of the holotype and two topotypes of M. frater, Schwartz and
Jones (1967), external and cranial measurements of specimens from Angulla, Barbuda,
Antigua, Dominica, St. Lucia, and Barbados; Choate and Birney (1968), cranial measure¬
ments of six males and nine females from Dominica and of one sub-Recent fossil from Puerto
Rico; Koopman (1968), forearm and cranial measurements of one male from Dominica,
a specimen from Antigua, and one female from Anguilla; Homan and Jones (19756),
external and cranial measurements (range) of Lesser Antillean representatives of the species
(after Schwartz and Jones, 1967).

Geographic variation. — Schwartz and Jones (1967) have recently reviewed geographic
variation in Monophyllus plethodon. They recognized three subspecies occurring on Puerto
Rico and the Lesser Antilles. One subspecies was known only as a fossil from Puerto Rico.
Specimens of M. plethodon on Barbados were distinguished from all other Lesser Antillean
populations by overall small size.

BIOLOGY OF THE PHYLLOSTOMATIDAE

Monophyllus redmani Leach, 1821

Measurements of Monophyllus redmani have been recorded as follows: Gundlach (1872,
1877), external measurements of a Cuban specimen; Dobson ( 1 878a), external measure¬
ments of one male; Miller (1900a), external and cranial measurements of the male holotype
of M. portoricensis from Puerto Rico, the male holotype of M. clinedaphus from an unknown
locality, and a male from Jamaica, as well as external measurements of one male and three
females from Puerto Rico; Miller (1902a), external and cranial measurements of the male
holotype of M. cubanus from Cuba and cranial measurements of one male from Jamaica;
Elliot (1904), external and cranial measurements of one specimen each from Puerto Rico,
Cuba, and Jamaica; Miller (1904), external measurements of eight males and seven females
from Cuba; Anthony (1917), cranial measurements of a specimen from Puerto Rico;
Anthony (1918, 1925), external (18 specimens) and cranial (five specimens) measurements
(mean, range) of individuals from Puerto Rico; Hall and Kelson (1959), external and
cranial measurements of a male from Jamaica, the holotype of M. cubanus, and of the
holotype of M. clinedaphus, as well as cranial measurements of the holotype of M. portori¬
censis and the range in external measurements of five specimens from Puerto Rico;
Schwartz and Jones (1967), external and cranial measurements of the three recognized
subspecies from Jamaica, Cuba, Hispaniola, and Puerto Rico; Choate and Birney (1968),
cranial measurements of one fossil specimen from Puerto Rico; Silva-Taboada (1974),
measurements of fossil humeri, crania, and mandibles from Cuba; Buden (1975a), external
and cranial measurements (mean, range) of specimens from Jamaica, Cuba, Hispaniola,
Bahamas, and Puerto Rico; Homan and Jones (1975a), external and cranial measurements
(range) of specimens of the three recognized subspecies (after Schwartz and Jones, 1967;
Buden, 1975 a).

Geographic variation. — Schwartz and Jones (1967) have recently reviewed geographic
variation in Monophyllus redmani. They recognized three subspecies, all occurring in the
Greater Antilles. Specimens from Jamaica were characterized by large body and cranial
size but a relatively short forearm. On Cuba and Hispaniola, bats were characterized by
small body, moderate skull size, and relatively long forearms. Specimens of M. redmani
from Puerto Rico are of generally small size.

Musonycteris Harrison i Schaldach and McLaughlin, 1960

Measurements of Musonycteris harrisoni have been recorded as follows: Schaldach and
McLaughlin (1960), external and cranial measurements of the male holotype, 10 male
paratypes, and two female paratypes from Colima; Villa-R. (1967), external measurements
of nine specimens (mean, range), and cranial measurements (mean, range) of six specimens
from Colima; Goodwin (1969), forearm and cranial measurements of one male from
Guerrero and a male and female from Colima.

Platalina genovensium Thomas, 1928

Measurements of Platalina genovensium have been recorded as follows: Thomas
(1928a), external and cranial measurements of the male holotype from Peru; Sanborn
(1936), external and cranial measurements of the male holotype and a second male from
Peru; Sanborn (1943), forearm measurements (range) for the species from Peru; Aellen
(1965), external and cranial measurements of a male in addition to the holotype (Thomas,
1928a), and one male (Sanborn, 1936) from Peru.

Scleronyeteris ega Thomas, 1912

Thomas (19126) gave external and cranial measurements of the female holotype from
Ega, Brazil.

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Subfamily Carolliinae
Carollia brevicauda (Schinz, 1821)

Measurements of Carollia brevicauda have been recorded as follows: Peters ( 1 865 z/),
external measurements of one specimen; Dobson (1878a), external measurements of one
specimen; H. Allen ( 18906), external measurements of three males and six females; Robin¬
son and Lyon (1901), external measurements of two males from Venezuela; Goodwin
(1942a), external and cranial measurements of one male and one female from Honduras
(originally reported as C. castanea)', Dalquest (1953a), external measurements of one male
and six females (mean) and cranial measurements (mean) of six males and five females from
San Luis Potosl (originally reported as perspicillata)', Jones (1966), forearm and cranial
measurements (range) of 12 specimens from Guatemala (another specimen representing
C. subrufa is included in ranges); Goodwin (1969), external and cranial measurements
of a male and a female from Chiltepec, Oaxaca (these were originally listed as C. subrufa ,
but according to Pine, 1972, these two specimens are probably C. brevicauda)'. Pine (1972),
external measurements (mean, range) of four males and 10 females from San Luis Potosl,
11 males and 17 females from Veracruz, seven males and 23 females from Chiapas, 15
males and 10 females from Guatemala, one male and one female from Honduras, 20
males and 26 females from Panama, nine males and seven females from Ecuador, eight
females from Brazil, four females from Peru, one male from Bolivia, and cranial measure¬
ments (mean, range) of five males and nine females from San Luis Potosl, 11 males and 15
females from Veracruz, seven males and 23 females from Chiapas, 15 males and 10 females
from Guatemala, one male and one female from Honduras, 20 males and 26 females from
Panama, nine males and seven females from Ecuador, eight females from Brazil, five males
and eight females from Peru, and one male from Bolivia; Jones et al. (1973), external and
cranial measurements (mean, range) of 20 specimens from the Yucatan Peninsula.

Geographic variation. — According to Pine (1972), specimens from the northernmost
part of the geographic range of the species in Mexico are the largest.

Carollia castanea H. Allen, 1890

Measurements of Carollia castanea have been recorded as follows: H. Allen (18906),
external measurements of the young male holotype from Costa Rica; Elliot (1904), external
measurements of the holotype as given by H. Allen (18906) from Costa Rica; Hahn (1907),
external and cranial measurements of the holotype from Costa Rica; Goodwin (1946), fore¬
arm and cranial measurements of the holotype and a second male from Costa Rica;
Hershkovitz (1949), external and cranial measurements of one male and one female from
Colombia; Husson (1962), external and cranial measurements of two females from Surinam;
Pirlot (1968), external and cranial measurements discussed in conjunction with C. perspicillata'.
Pine (1972), external measurements of 10 males and four females from Honduras, five
males from Nicaragua, seven males and eight females from Costa Rica, 31 males and 20
females from Panama, five males from Colombia, three males and two females from Peru,
one female from Bolivia, and cranial measurements of 10 males and four females from
Honduras, five males from Nicaragua, seven males and eight females from Costa Rica,
31 males and 19 females from Panama, four females from Colombia, seven males and four
females from Peru, and one female from Bolivia.

Geographic variation. — Pine (1972) could detect no geographic trends in variation in this
species; therefore, he considered C. castanea to be monotypic.

Carollia perspicillata (Linnaeus, 1758)

Measurements of Carollia perspicillata have been recorded as follows: Saussure (1860c),
external measurements of one specimen; Peters (1866a), external measurements of one
specimen; Miller (1902a), external and cranial measurements of the female holotype of C.

BIOLOGY OF THE PHYLLOSTOMATIDAE

tricolor from Paraguay; Elliot (1904), external and cranial measurements of a single
specimen; Hahn (1907), external measurements (mean) of nine specimens from Paraguay,

10 from Brazil, 10 from Trinidad, two from Guyana, 10 from northern Ecuador, nine from
Colon, Panama, six from Panama, Panama, nine from Nicaragua, 13 from Veracruz,

1 1 from Oaxaca, two from Campeche, and 13 from Veracruz (Jaltipan), and cranial measure¬
ments (mean) of eight specimens from Paraguay, two from Sao Paulo, Brazil, five from
Naranhoa, Brazil, five from Trinidad, two from Guyana, four from Venezuela, nine from
Colombia, 10 from Ecuador, eight from Oaxaca, six from Veracruz, two from Costa Rica,
two from Campeche, three from Colon, Panama, six from Boqueron, Panama, and six from
Panama, Panama; Lima (1926), external measurements of a specimen from Brazil; Sanborn
(1932), forearm measurement of one specimen from Bolivia; Goodwin (1934), external
measurements of one specimen from Guatemala; Cunha Vieira (1942), external measure¬
ments of nine males and four females and cranial measurements of three males and one
female from Brazil; Goodwin (1942 a), external and cranial measurements of two males from
Honduras; Goodwin (1946), external and cranial measurements of a male and female
from Costa Rica; Hershkovitz (1949), forearm measurements (range) in 79 specimens from
northern Colombia and the mean of the greatest length of skull in this sample (some specimens
in this sample are brevicauda, see Pine, 1972); Dalquest (1951), forearm and cranial
measurements (mean) of 27 specimens of both sexes combined from Trinidad; Felten
(1956a), external measurements (mean, range) of 15 males and 28 females and cranial
measurements of 10 males and 19 females from El Salvador; Felten (1956*/), external
measurements (mean, range) of specimens from El Salvador; Ryan (1960), external measure¬
ments of one male from Guatemala; Goodwin and Greenhall (1961), forearm measurements
(range) of 30 specimens and cranial measurements of one male and one female from Trinidad;
Husson (1962), external and cranial measurements of five males and five females from
Surinam; Burt and Stirton (1961), forearm and cranial measurements (range) of 22 males
and 14 females combined from El Salvador; Pirlot (1963), external measurements of specimens
from Venezuela; Butterworth and Starrett (1964), cranial measurements of a male and
female from Venezuela; Starrett and de la Torre (1964), external and cranial measure¬
ments of one male from Nicaragua and two males and a female from Costa Rica; Tamsitt
and Valdivieso (1963a), external measurements (mean, range) of 28 specimens and cranial
measurements of 11 from Colombia; Tamsitt and Valdivieso (19636), external measure¬
ments (mean, range) of four males from Colombia; Valdivieso (1964), external and cranial
measurements (mean, range) of 19 specimens from Colombia; Brosset (1965), external and
cranial measurements of three males from Ecuador; Jones (1966), forearm and cranial
measurements (range) of specimens from Guatemala; Pirlot (19656), external measure¬
ments of 14 males and 10 females from Est du Venezuela and 19 males and 15 females from
Zulia, Venezuela; Pirlot (1968), external and cranial measurements discussed in conjunction
with C. castanea; Goodwin (1969), forearm and cranial measurements of nine males
and three females from Oaxaca; Pine (1972), external and cranial measurements (mean, range)
of males and females throughout the range of the species; Pirlot (1972), external measure¬
ments of a specimen from Brazil; Jones et al. (1973), external and cranial measurements
(mean, range) of 10 specimens from the Yucatan Peninsula; Smith and Genoways (1974),
forearm measurements of two specimens from Margarita Island, Venezuela; Taddei
(19756), external measurements (mean, se, range) of 30 males and 30 females, and cranial
measurements of 15 males and 15 females from Brazil.

Individual variation. — In specimens from Brazil, coefficients of variation for external
measurements varied in 30 males from 2.70 to 6.15 and in 30 females from 2.70 to 5.94;
CV values for cranial measurements in 15 males ranged from 1.78 to 4.01 and in 15 females
from 1.85 to 4. 11 (Taddei, 19756). According to Tamsitt and Valdivieso (1963a), specimens
from central Colombia were homogeneous in size.

Secondary sexual variation. — In specimens from Brazil, females generally averaged larger
than males in external measurements and in four (head and body, ear, forearm, metacarpal

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II) of 17 measurements they proved to be significantly so. Cranial measurements showed the
opposite in one of 15 measurements (mastoid breadth), males proved to be significantly
larger than females (Taddei, 1 975 />). Pine (1972) also found cranial measurements of males
to average slightly larger than those of females. However, Tamsitt and Valdivieso ( 1 963a)
reported that their males and females were of the same size in a sample of 16 males and 12
females from Colombia.

Geographic variation. — Tamsitt and Valdivieso (1963a) found individuals from localities
on each side of the East Andes not to differ in any way. Their specimens, although slightly
smaller, did not differ significantly from the range of measurements given by Hershkovitz
(1949) for northern Colombian specimens. According to Pine (1972), specimens from in
and around the Panama drainage are characteristically small. Dalquest (1951), comparing
forearm length and cranial measurements of his specimens from Trinidad with examples
from San Luis Potosi, concluded that they are alike (however, the material from San Luis
Potosi was probably C. hrevicauda).

Carollia subrufa (Hahn, 1905)

Measurements of Carollia subrufa have been recorded as follows: Hahn (1905), external
and cranial measurements of the male holotype from Oaxaca; Hahn (1907), external measure¬
ments of eight specimens from Oaxaca, seven from Colima, four from Campeche, and one
from Honduras, and cranial measurements of nine specimens from Oaxaca, four from
Colima, two from Campeche, and one from Honduras; Elliot (1917), external and cranial
measurements of the holotype; Goodwin (1934), external measurements of one specimen
from Guatemala; Goodwin (1942a), external and cranial measurements of two males from
Honduras; Felten (1956a), external measurements (mean, range) of 99 males and 99 females
and cranial measurements of 27 males and 33 females from El Salvador (as a subspecies of
C. castanea)', Felten ( 1 956z/), external measurements (mean, range) of specimens from El
Salvador; Hall and Kelson (1959), external measurements (range) of 198 (99 males, 99
females) specimens from El Salvador listed as C. castanea ; Ryan (1960), external measure¬
ments of one female from Guatemala; Burt and Stirton (1961), external and cranial measure¬
ments of four males from El Salvador (as a subspecies of castanea ); Starrett and de la Torre
(1964), external and cranial measurements of two males from El Salvador; Jones (1966),
forearm and cranial measurements of one male from Jocotan, Guatemala (others listed are
C. brevicauda); Villa-R. (1967), external measurements (mean, se, range) of 51 males and
females combined and cranial measurements (mean, se, range) of 38 males and females
combined; Goodwin (1969), forearm and cranial measurements of six males and one female
from Oaxaca, (also lists a male and a female from Chiltepec, but, according to Pine, 1972,
these are probably C. brevicauda)', Pine (1972), external measurements of one male from
Colima, two males and eight females from Oaxaca, 16 males from Chiapas, one male and
one female from Honduras, two males and seven females from Nicaragua, and cranial
measurements of two males and five females from Colima, two males and eight females
from Oaxaca, 16 males and 24 females from Chiapas, one male and one female from
Honduras, and two males and seven females from Nicaragua; Watkins et al. (1972),
external and cranial measurements of one female from Jalisco.

Geographic variation. — Pine (1972) found specimens from the northern part of the
geographic range of the species to be larger than those of the southernmost part of the
geographic range.

Rhinophylla alethina Handley, 1966

Handley (1966a) gave external measurements (mean, range) of six males and four females,
and cranial measurements of the male holotype from Colombia.

BIOLOGY OF THE PHYLLOSTOMATIDAE

Rhinophylla fischerae Carter, 1966

Measurements of Rhinophylla fischerae have been recorded as follows: Carter (1966),
external and cranial measurements of the female holotype from Peru, six additional females
and two males, all from the type locality except one female from Pucallpa, Peru; Marinkelle
and Cadena (1972), external measurements of a male and female from Colombia; Mumford
(1975), external and cranial measurements of an unsexed specimen from Ecuador.

Rhinophylla pumilio Peters, 1865

Measurements of Rhinophylla pumilio have been recorded as follows: Peters (1865 a),
external measurements of the holotype from Brazil; Dobson ( 1 878«), external measurements
of one specimen from Brazil; Sanborn (1936), external and cranial measurements of a male
and female from Ecuador; Husson (1962), external and cranial measurements of two females
from Surinam and two from Guyana; Hill (1964), forearm measurements of two males and
cranial measurements of one of these from Guyana; Carter (1966), external measurements
of 15 males and 10 females combined, and cranial measurements (mean, range) of 15 males
and 13 females combined from Venezuela, Brazil, Ecuador, and Peru; Marinkelle and
Cadena (1972), forearm and cranial measurements of a male (juvenile) and the range of three
females from Colombia.

Subfamily Stenoderminae
Ametrida centurio Gray, 1847

Measurements of Ametrida centurio have been recorded as follows: Peters ( 1 866c/),
external measurements of one specimen; Dobson (1878a), external measurements of the
female holotype from Brazil; H. Allen (18946), external and cranial measurements of the
male holotype of A. minor from Surinam (type locality according to Peterson, 1965) and
external measurements of a specimen of A. centurio ; Sanborn (1938), external and cranial
measurements of a male (female according to Peterson, 19656) from Brazil; Husson (1960),
cranial measurements of one specimen from Bonaire; Goodwin and Greenhall (1961),
forearm and cranial measurements of a male from Guyana, a female from Venezuela, and
a subadult from Trinidad; Husson (1962), external and cranial measurements of two males
and two females (see Peterson, 19656:3-4, on the question of the sex of one of these specimens)
from Surinam and one male from Bonaire; Peterson (1965), forearm and cranial measure¬
ments of 12 males from Brazil, Guyana, Surinam, Venezuela, Trinidad, and Bonaire
(including the holotype of A. minor from Surinam), 13 females from Brazil, Guyana,
Venezuela, Trinidad, and Surinam (including the holotype of A. centurio from Brazil), and
external measurements (mean, range) of males and females.

Secondary sexual variation. — Peterson (19656) described distinct differences in size
between the sexes with no overlap in forearm length or the following cranial measure¬
ments: condylobasal length; least interorbital width; breadth of palate (Ml-Ml); toothrow
length (C-M3).

Ardops nichollsi (Thomas, 1891)

Measurements of Ardops nichollsi have been recorded as follows: Thomas (1891a),
external and cranial measurements of the female holotype of A. n. nichollsi from Dominica;
Thomas (1894), external and cranial measurements of the male holotype of A. n. montser-
ratensis from Montserrat; Elliot (1904), external and cranial measurements of one specimen
from Monsterrat, one from Dominica, and one from St. Lucia; Miller (1902a), external
and cranial measurements of the female holotype of A. n. luciae from St. Lucia and of a
male from Dominica; Miller (1913a), external and cranial measurements of the female

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holotype of A. n. annectens and a male from Guadeloupe; Elliot (1917), external and
cranial measurements of the holotype of A. n. annectens , G. M. Allen (1942), forearm length
of taxa described at that time; Hall and Kelson (1959), externa] and cranial measurements of
the holotypes of A. n. monsterratensis, A. n. annectens , and A. n. luciae; Jones and Schwartz
(1967), forearm and cranial measurements of the female holotype of A. n. nichollsi, external
measurements (mean, range) of six males and seven females, cranial measurements (mean,
range) of eight males and seven females from Dominica, external and cranial measurements
of a male and a female from St. Eustatius, and the male holotype of A. n. monserratensis
from Montserrat, and the female holotype of A. n. luciae , cranial measurements of a female,
forearm measurements of one male and four females from St. Lucia, external measurements
of an adult male and the female holotype of A. n. annectens , cranial measurements of the
holotype, two males, and two females, forearm measurements of four females from
Guadeloupe, external measurements of the female holotype (A. n. koopmani), another
female, and two males, and cranial measurements of the female holotype and a male from
Martinique; Jones and Genoways (1973), some measurements as given by Jones and
Schwartz (1967).

Secondary sexual variation. — In individuals from Dominica, females were clearly larger
than males. This was also found to be true in one male and one female from Martinique
(Jones and Schwartz, 1967).

Geographic variation. — According to Jones and Schwartz (1967), specimens from
Dominica were the smallest of the species, whereas those from St. Eustatius and Montserrat
were the largest. Specimens from Martinique differed from those on adjacent islands,
Dominica to the north and St. Lucia to the south, in being considerably larger.

Ariteus flavescens(Gray, 1831)

Measurements of Ariteus flavescens have been recorded as follows: Peters (1876),
external measurements of a specimen of Peltorhinus achradophilus (= A. flavescens );
Dobson (1878c/), external measurements of the female holotype of Ariteus achradophilus
from Jamaica; Elliot (1904), external and cranial measurements of one specimen from
Jamaica; G. M. Allen (1942), external measurements for the species; Howe (1974), external
measurements of two males and two females from Jamaica.

Artibeus aztecus Andersen, 1906

Measurements of Artibeus aztecus have been recorded as follows: Andersen (19066),
external measurements of the male holotype of A. aztecus from Morelos; Andersen (1908),
external and cranial measurements (range) of four specimens from Morelos; Elliot (1917),
cranial measurements of the holotype; Dalquest (1953c/), external measurements of a male
and two females and cranial measurements of the male and one female from San Luis Potosf;
Lukins and Davis (1957), forearm and cranial measurement (range) for the species; Villa-R.
(1967), external and cranial measurements of one female from the state of Mexico; Koopman
(1961), forearm and cranial measurements (range) of four specimens (one male, three females)
from Sinaloa; Baker and Greer (1962), external and cranial measurements of a female from
Durango; Alvarez (1963), external and cranial measurements of three males and one female
from Tamaulipas; Jones (1964), forearm and cranial measurements (mean, range) of 15
specimens (10 males and five females) from Sinaloa; Davis (1969), external and cranial
measurements (mean, range) of 33 specimens from the Mexican highlands, 41 from the
Guatemalan highlands, and 18 from the Costa Rican highlands, and external and cranial
measurements of the male holotype of A. aztecus aztecus from Morelos, the male holotype
of A. a. minor from Guatemala, and the male holotype of A. a. major from Costa Rica;
Goodwin (1969), forearm and cranial measurements of four males and five females from
Oaxaca; Alvarez and Ramirez-Pulido (1972), external and cranial measurements of two

BIOLOGY OF THE PHYLLOSTOMATIDAE

males from Michoacan, and a female from Oaxaca; Jones et al. (1972), forearm and cranial
measurements as given by Jones (1964).

Geographic variation. — Artiheus aztecus, which occurs in the Middle American high¬
lands, was segregated into three recognizable populations — aztecus in the Mexican highlands,
minor from the Guatemalan highlands, and major of the Costa Rican highlands. With regard
to size, A. a. major is the largest, and minor is the smallest (Davis, 1969).

Artibeus cinereus (Gervais, 1855)

Measurements of Artibeus cinereus have been recorded as follows: Peters ( 1 865 r/),
external measurements of the holotype of A. quadrivittatum from Surinam; Dobson
( 1 878«), external measurements of a male and a female; Robinson and Lyon (1901), external
measurements of three males and six females from Venezuela; Andersen (19066), cranial
measurements (range) of eight specimens including the male holotype (Colombia) of A.
cinereus bogotensis from Colombia and Venezuela and seven additional specimens of A. c.
cinereusr, Andersen (1908), external measurements (mean, range) of 10 specimens and
cranial measurements (mean, range) of eight from Guyana, Trinidad, and Venezuela, external
and cranial measurements (mean, range) of eight specimens from Colombia and Venezuela
and the range of these measurements in three specimens of A. quadrivittatus from Surinam;
Lima (1926), external measurements of a male from Brazil; Sanborn (1932), forearm measure¬
ments of a female and a specimen of unknown sex and cranial measurements of the female
from Bolivia; Cunha Vieira (1942), external measurements of two females from Venezuela
and external measurements of a male from Ecuador; Hershkovitz (1949), external and cranial
measurements of a female from Colombia; Goodwin and Greenhall (1961), forearm and
cranial measurements of three males and one female from Trinidad; Burt and Stirton (1961),
forearm and cranial measurements (range) of four males and 14 females from El Salvador;
Husson (1962), external and cranial measurements of three males, four females, and the
unsexed holotype of A. quadrivittatus from Surinam; Tamsitt and Valdivieso (1963 a),
external measurements of four females from Colombia; Brosset (1965), external and cranial
measurements of a male from Ecuador; Tamsitt and Valdivieso (1966a), forearm and cranial
measurements of a male and female from Colombia (values for the female as given by
Hershkovitz, 1949); Davis (19706), external and cranial measurements (mean, range) of
36 specimens from Trinidad; Tuttle (1970), forearm measurements (range) of specimens
from east of the Andes in Peru; Pirlot (1972), external measurements of two males and one
female from Brazil (type description of A. c. solimoesi ).

Artibeus eoncolor Peters, 1865

Measurements of Artibeus eoncolor have been recorded as follows: Peters (1865a),
external measurements of the holotype from Surinam; Thomas (1892), forearm and cranial
measurements of the holotype; Andersen (1908), external and cranial measurements of a
female from Surinam and cranial measurements of the holotype from Surinam; Cabrera
(1917), external and cranial measurements of a female possibly from Brazil; Cunha Vieira
(1942), external measurements based on Andersen (1908); Husson (1962), external and
cranial measurements of a female as given by Andersen (1908) and measurements of the
holotype as given by Peters and Thomas; Hill (1964), forearm and cranial measurements
of one male from Guyana; Linares (1969), external measurements of a male and two females
from Venezuela; Gardner (1976), external and cranial measurements of a male from Peru.

Artibeus glaucus Thomas, 1893

Measurements of Artibeus glaucus have been recorded as follows: Thomas (1893), external
and cranial measurements of the female holotype from Peru; Andersen (1908), external and
cranial measurements of the holotype from Peru; Davis (1970a), cranial measurements
(mean, range) of nine specimens from Peru and Ecuador.

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Artibeus hirsutus Andersen, 1906

Measurements of Artibeus hirsutus have been recorded as follows: Andersen (19066),
forearm and cranial measurements (range) of eight specimens from Michoacan, Colima,
and Jalisco; Andersen (1908), external and cranial measurements (mean, range) of eight
specimens from Michoacan, Colima, and Jalisco; Elliot (1917), cranial measurements of the
holotype; Davis and Russell (1952), external and cranial measurements of one male and
five females (mean, range) from Morelos; Anderson (1960), external and cranial measure¬
ments (mean, range) of 28 specimens from Guerrero; Davis and Carter (1964), external and
cranial measurements (mean, range) of six females; Villa-R. (1967), external measurements
(mean, sd, range) of 55 specimens and cranial measurements of 46 specimens from Sonora,
Sinaloa, Nayarit, Jalisco, Morelos, and Guerrero; Genoways and Jones (1968), forearm
measurements (mean, range) of four young males and four females from Zacatecas; Goodwin
(1969), forearm and cranial measurements of two males from Guerrero and two from
Sonora; Anderson (1972), external and cranial measurements of three specimens from
Chihuahua; Jones et al. (1972), forearm and cranial measurements (mean, range) of 10
specimens (five males and five females) from Sinaloa.

Secondary sexual variation. — Anderson (1960) found no significant size differences
between sexes in four external and four cranial measurements in a sample of 28 specimens
from Guerrero.

Artibeus inopinatus Davis and Carter, 1964

Davis and Carter (1964) reported external and cranial measurements (mean, range) of
eight females from Honduras and forearm measurements of one male from Honduras and
one from Nicaragua. Although Davis and Carter did not examine the specimens reported
from El Salvador by Burt and Stirton (1961), under the name Artibeus hirsutus, they judged,
and we agree, from the published measurements that the specimens are referable to A.
inopinatus.

Artibeus jamaicensis Leach, 1821

Measurements of Artibeus jamaicensis have been recorded as follows: Saussure (18606),
external measurements of one specimen; Gundlach (1872, 1877), external measurements of
a specimen from Cuba; Dobson (1878a), external measurements for a male of A. perspicillatus
from Guatemala and a female; Cope (1889), external measurements of one male cotype
of Dermanura eva from St. Martin, Lesser Antilles; H. Allen (1894a), external measurements
from three specimens (two from Mexico, one locality unknown) and cranial measurements
(mean) of three specimens from an unspecified locality; J. A. Allen and Chapman (1897a),
forearm measurements of four specimens from Yucatan, 10 from Jamaica, 31 females and
20 males from Cuba; Rehn (1900), cranial measurements of the two male cotypes of
Dermanura eva Cope from St. Martin, Lesser Antilles, a specimen from Jamaica, and one
from Brazil; Robinson and Lyon (1901), external measurements of a male and two females
from Venezuela; Rehn (19026), external measurements of the unsexed holotype of A. hercules
( = A. jamaicensis) and the mean of external measurements for two additional specimens,
cranial measurements of a specimen from Peru, external measurements of the male holotype,
the mean for six specimens of A. parvipes ( — A. jamaicensis) from Cuba, and one specimen
of A. jamaicensis from Jamaica, the mean of six specimens and external measurements
(mean) of two specimens of A. planirostris and cranial measurements of one from Brazil;
J. A. Allen (1904), external and cranial measurements of the male holotype of A. insularis
from St. Kitts, Lesser Antilles, and the male holotype of A. j. yucatanicus from Yucatan;
Elliot (1904), external and cranial measurements of one specimen each of A. coryi, A.
jamaicensis, A.j. parvipes, and A. j. planirostris; Miller (1904), external measurements of 12
males and 13 females from Cuba; Elliot (1905a), external and cranial measurements of a
specimen from St. Kitts Island, Lesser Antilles; Andersen (1906), cranial measurements

BIOLOGY OF THE PHYLLOSTOMATIDAE

(mean) of 65 specimens of A. j. jamaicensis and external measurements (range) of three
specimens of A. j. praeceps from Guadeloupe; G. M. Allen (1908), external measurements
of three specimens and cranial measurements of one male from Brazil, and external measure¬
ments of one specimen from Jamaica; J. A. Allen (1908u), forearm measurements (range)
of four specimens from the Dominican Republic; J. A. Allen (19086), external and cranial
measurements of the male holotype of A. j. richardsoni from Nicaragua; Andersen (1908),
external and cranial measurements (range) of 16 specimens (11 cranial) from Brazil, three
from Venezuela, and three from Chiapas and Guerrero, median and range of the above
combined, 13 specimens (nine cranial) from Trinidad and Tobago, nine (eight cranial) from
Grenada, 41 (33 cranial) from Surinam, Cayenne, Guyana, and Lower Orinoco, 25 speci¬
mens (12 cranial) from Cuba, 14 (12 cranial) from Yucatan and Cozumel Island, 12 (nine
cranial) from Central America, 27 (23 cranial) from southern Mexico, 21 (11 cranial) from
Puerto Rico, three from Dominican Republic, one from St. Kitts Island, eight (five cranial)
from St. Andrews and Old Providence Island, and 95 (65 cranial) (median, range) of A. j.
jamaicensis (including much of the above data); Elliot (1917), external and cranial measure¬
ments of the holotype; Anthony (1919), cranial measurements of fossil material from Cuba;
Anthony (1924«), external and cranial measurements of the female holotype of A. j. frater-
culus from Ecuador, forearm measurements (mean) of 18 specimens and cranial measure¬
ments (mean, range) of 13 others; Anthony (1918, 1925), external measurements (mean,
range) of 24 specimens and cranial measurements (mean, range) of 10 specimens (five
males, five females) from Puerto Rico; Goodwin (1934), external measurements of one
specimen from Guatemala; Sanborn (1936), forearm measurements (range) of three males
and four females and cranial measurements (range) of three specimens (one male, two fe¬
males) from Barbados; Martinez and Villa-R. (1938), external measurements of five males
and nine females from Morelos; Cunha Vieira (1942), external and cranial measurements
of a male from Brazil; Goodwin (1942), forearm and cranial measurements of two males
from Honduras, and these measurements of another specimen; Goodwin (1946), external
and cranial measurements (range) for the species; Hall and Villa-R. (1949), external and
cranial measurements of one female from Michoacan; Hershkovitz (1949), external and
cranial measurements of a male and female (two males and a female for forearm) from
Colombia; Dalquest (1951), forearm and cranial measurements (mean) of four males and
eight females from Trinidad; Dalquest ( 1 95 3 zz), external measurements (mean) of eight males
and eight females and cranial measurements (mean) of two males and 1 1 females from San
Luis Potosf; Goodwin (1953), forearm and cranial measurements of the male holotype of
A. coryi from St. Andrews Island, the male holotype of A. insular is from St. Kitts, the male
holotype of A. j. richardsoni from Nicaragua, the male holotype of A. j. yucatanicus
from Yucatan, and the female holotype of A. j. fraterculus from Ecuador; de la Torre
(1955), forearm measurements (mean, range) of five specimens (three males, two fe¬
males) from Jalisco; de la Torre (1954), external and cranial measurements (mean,
range) of 23 specimens from Tamaulipas; Felten (1956u), external measurements (mean,
range) of 16 males and five females and cranial measurements of nine males (mean,
range) and one female from El Salvador; Felten (1956 d), external measurements (mean,
range) of specimens from El Salvador; Anderson (1960), external and cranial measure¬
ments (range) of three specimens from Sinaloa, and four from Jalisco; Husson (1960),
cranial measurements (mean, range) of specimens from Curasao and St. Martin; Burt
and Stirton (1961), forearm and cranial measurements (range) of 44 specimens (18
males, 26 females) from El Salvador; Goodwin and Greenhall (1961), forearm measure¬
ments (range) of 12 males and 18 females, and cranial measurements of one male and one
female from Trinidad; Baker and Greer (1962), external and cranial measurements of a male
and female from Durango; Pirlot (1963), forearm measurements (range) of 35 males and
20 females from Venezuela; Tamsitt and Valdivieso (1963«), external measurements of one
male and three females and cranial measurements of one female from Colombia; Davis and
Carter (1964), external and cranial measurements (mean, range) of eight females from Central
America; Hill (1964), forearm measurements of two males and three females and cranial

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measurements of two males and three females and cranial measurements of two males
and two females from Guyana; Valdivieso (1964), external measurements of one male and
two females and cranial measurements of one female from Colombia; Starrett and de la
Torre (1964), external and cranial measurements of one male from Nicaragua and one from
Costa Rica; Handley (1965), external and cranial measurements of the female holotype of
A. j. triomylus from Guerrero and mean and range of external measurements of 10 males
and nine females and cranial measurements of 12 females and 10 males from Guerrero;
Pirlot (19656), external measurements of 15 males and 33 females from Est du Venezuela
and of 35 males and 20 females from Zulia, Venezuela; Villa-R. (1967), external measure¬
ments of 46 specimens and cranial measurements of 43 specimens of A. j. triomylus from
Mexico, and external measurements of 76 specimens and cranial measurements of 71
specimens of A. j. yucatanicus from Mexico; Genoways and Jones (1968), mean and range
of forearm measurements of six young specimens (two males, four females) and individual
forearm measurements of two young males and one young female from Zacatecas; Koopman
(1968), forearm and cranial measurements of the holotype of A. praeceps (Guadeloupe)
and specimens (range) from Guadeloupe and Dominica; Pirlot (1968), forearm measure¬
ments of a female from Peru; Goodwin (1969), forearm and cranial measurements of four
males and three females of A. j. yucatanicus from Oaxaca and three males and three females
of A. j. triomylus from Oaxaca; Jones and Phillips (1970), forearm measurements (mean,
range) of seven specimens from Barbados, 11 from St. Lucia, 20 from St. Vincent, 23 from
Grenada, and 16 from Trinidad, and cranial measurements for 7, 15, 32, 15, and 1 1 specimens,
respectively; Davis (19706), external and cranial measurements of the male holotype of A.
j. richardsoni from Nicaragua, mean and range of 13 topotypes, means of 14 from Chiapas,
12 from Guatemala (Alta Verapaz), 20 from Guatemala (Puerto Barrios), 20 from Nicaragua
(Castillo), 20 from Honduras (coastal), 16 from Costa Rica (coastal), 20 from Panama
(Veraguas), 21 from Panama (Chepo) of A. j. richardsoni , external and cranial measure¬
ments of the male holotype of A. j. yucatanicus from Yucatan, mean and range of eight
topotypes, mean of 18 from Tamaulipas, 25 from San Luis Potosf, 19 from Veracruz, 14
from Campeche and Yucatan, four from British Honduras, 20 from Honduras (Bay Islands)
of A. j. yucatanicus, forearm and cranial measurements of the female holotype of A. j. trio¬
mylus from Guerrero, mean and range of 20 from near the type locality, external and cranial
measurements of the female holotype of A. j. paulus from El Salvador, means of 15 from
Chiapas (below 1000 feet), 20 from Guatemala, 20 from El Salvador, 20 from Honduras
(Nueva Ocotepeque), six from Honduras (Pacific lowlands), 11 from Nicaragua (San
Antonio), and four from Costa Rica (Guanacaste Lowlands) of A. j. paulus ; Tuttle (1970),
cranial measurements of a female from Peru, and range in forearm length of specimens east
of the Andes; Jones et al. (1972), forearm and cranial measurements (mean, range) of 10
specimens (five males, five females) from Sinaloa; Smith and Genoways (1974), forearm and
cranial measurements (mean, range) from four localities in Venezuela (sample sizes five,
22, 17, 22) and eight specimens from Trinidad.

Age variation. — According to Davis (19706), young individuals in which the cartilaginous
epiphyses of finger joints were readily discernable were consistantly smaller than adults in
all measurements. However, individuals in which the joint of the finger was only swollen and
in which the epiphyses and diaphyses appeared to be united were as large as adults in all
measurements.

Individual variation. — Within sample variation of cranial measurements was shown by
Davis (19706) to be usually less than 10 per cent of the minimum value of each variate tested.
Of six cranial measurements tested, length of skull was the least variable and breadth across
upper molars the most. Wing measurements varied more than cranial. Of four wing
measurements examined, length of forearm was the least variable and length of phalanx 1,
digit III the most.

Secondary sexual variation. — Davis (19706) found no significant secondary sexual
variation in four wing and eight cranial measurements.

BIOLOGY OF THE PHYLLOSTOMATIDAE

Geographic variation. — Both Koopman (1968) and Jones and Phillips (1970) noted a trend
toward slightly larger size in specimens from the southern part of the Lesser Antilles. Jones
and Phillips (1970) found A. jamaicensis from Grenada to approach those from Trinidad
and Tobago in size. They also found that specimens from St. Vincent averaged considerably
larger than specimens from any other Antillean population.

Davis (19706), studying geographic variation in Middle American populations of Artibeus
jamaicensis, recognized four areas of differentiation. The largest individuals occurred along
the Atlantic versant of Middle America (northern Chiapas to eastern Panama). Greatest
length of skull in this area averaged near 29 and forearm near 61. The population along the
Atlantic versant of Mexico (Tamaulipas to the Yucatan Peninsula and into British Honduras
and on the Bay Islands of Honduras) was characterized by small size. More than 90
per cent of the individuals had a skull length of less than 28.45 combined with a zygomatic
breadth of less than 17.05. Populations from the Pacific versant were also characterized
by small size — those from Oaxaca and Morelos northward into Sinaloa and Durango normally
possessed three upper molars and had a zygomatic breadth seldom less than 17.0. Populations
from Chiapas southward to Guanacaste, Costa Rica, lacked the upper third molar.

Smith and Genoways (1974) found their material from Margarita Island, Venezuela,
averaged slightly smaller in external and cranial measurements than specimens from the
adjacent Venezuelan mainland and Trinidad.

Artibeus lituratus(01fers, 1818)

Measurements of Artibeus lituratus have been recorded as follows: J. A. Allen and
Chapman (18976), external measurements of the male holotype of A. I. palmarum from
Trinidad and a female, mean external measurements for five females, and cranial measure¬
ments of one female from Trinidad; J. A. Allen (1897), external and cranial measurements
of the male holotype of A. lituratus intermedins from Costa Rica; Bangs (1899), external
and cranial measurements of the male holotype of Artibeus femurvillosum from Colombia;
Robinson and Lyon (1901), external measurements of five males and 15 females from
Venezuela; Rehn (19026), external measurements of the holotype of A. I. Hercules from
Peru, the average of these measurements for two additional specimens and cranial measure¬
ments for one; J. A. Allen (1904), external and cranial measurements of the male holotype of
A. rusbyi from Peru; Elliot (1904), external and cranial measurements of a specimen of A.
lituratus intermedins ; G. M. Allen (1908), external measurements of three specimens and
cranial measurements of one from Brazil and forearm measurements of the holotype of A. I.
intermedins and three additional specimens from Costa Rica; Andersen (1908), external
and cranial measurements (mean, range) of 12 specimens (six cranial) from Paraguay, 20
(19 cranial) from Brazil, and nine (eight cranial) from Ecuador and Colombia, means for
these measurements for 15 specimens (10 cranial) from Venezuela, four (three cranial) from
Trinidad and St. Vincent, 20 (15 cranial) from Central America (Panama, Costa Rica,
Nicaragua, Guatemala), four (three cranial) from Mexico (Veracruz, Jalisco, Oaxaca)
and a mean for these measurements from the latter localities, cranial measurements of six
specimens of A. I. aequatorialis from Ecuador, and external of seven and cranial measure¬
ments of six specimens (median, range) of A. 1. aequatorialis from Ecuador and Colombia;
Lima (1926), external measurements of a male and cranial measurements of an unsexed
individual from Brazil; Cunha Vieira (1942), external measurements of one male and four
females and cranial measurements of three males from Brazil; Goodwin (1942u), external
and cranial measurements of two females from Honduras; Hershkovitz (1949), external and
cranial measurements (range) of specimens from Colombia; Dalquest (1950), cranial measure¬
ments (mean) of three males and two females from San Luis Potosi; Dalquest (195 1), forearm
and cranial measurements (mean) of three males and six females from Trinidad; Dalquest
(1953u), external measurements of a male and two females (mean) and cranial measurements
(mean) of three males and two females from San Luis Potosi; Goodwin (1953), forearm and
cranial measurements of the male holotype of A. lituratus palmarum from Trinidad, the

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male holotype of A. lituratus intermedins from Costa Rica, and the male holotype of A. rushy i
from Peru; de la Torre (1954), external and cranial measurements of three specimens from
Tamaulipas; Felten (1956c), external measurements (mean, range) of six males and six
females and cranial measurements of five males and five females from El Salvador; Felten
( 1956c/), external measurements of specimens from El Salvador; Russell (1956), forearm
and cranial measurements of a female from Morelos; Lukens and Davis (1957), forearm
and cranial measurements (mean, range) of adult specimens, one juvenile female, and a
subadult female from Guerrero; Anderson (1960), external and cranial measurements (mean,
range) of 17 specimens from Sinaloa; Goodwin and Greenhall (1961), forearm measurements
(range) of 14 males and 18 females and cranial measurements of one male from Trinidad;
Tamsitt and Valdivieso (1963a), external and cranial measurements (mean, range) of 46
males and 30 females combined from Colombia; Tamsitt and Valdivieso (19636), external
measurements of a female from Colombia; Hill (1964), forearm and cranial measurements
of a female from Guyana; Starrett and de la Torre (1964), forearm measurements of a male
and female from El Salvador and a female from Costa Rica, other external and cranial
measurements of a male and female from Costa Rica; Valdivieso (1964), external and
cranial measurements (mean, range) of specimens from Colombia; Brosset (1965), external
and cranial measurements of five males (including the lectotype of A. fallax) and five females
from Surinam; Pirlot (19656), external measurements of eight males and eight females from
Est du Venezuela; Tamsitt and Valdivieso (1965a), forearm measurements (mean, range)
of monthly samples of males from Colombia; Tamsitt and Valdivieso (19656), external
measurements (mean, sd, se, range) of 80 adult and 18 young adult females from Colombia;
Tamsitt and Valdivieso (19666), external measurements (mean, range) of 14 specimens
(four males, 10 females) and cranial measurements of five females from Colombia; Villa-R.
(1967), external measurements of 46 specimens and cranial measurements of 34 specimens
from Mexico; Koopman (1968), forearm measurements (range) of seven specimens from
St. Vincent; Goodwin (1969), forearm and cranial measurements of four males and four
females from Oaxaca; Burt and Stirton (1969), forearm and cranial measurements (range)
of five specimens from El Salvador; Villa-R. and Villa Cornejo (1969), external and
cranial measurements (mean, range) of seven specimens from Argentina; Tuttle (1970),
forearm measurements (range) of specimens from east of the Andes in Peru; Jones et al.
(1972), forearm and cranial measurements (mean, range) of 10 specimens (five males, five
females) from Sinaloa; Pirlot (1972), external measurements of specimens from Brazil.

Age variation. — Lukens and Davis (1957) presented forearm and cranial measurements
of a juvenile female and a subadult female from Guerrero. Anderson (1960) gave external
and cranial measurements of an immature female from Sinaloa.

Secondary sexual variation. — Tamsitt and Valdivieso (1963a) found that females from
Colombia averaged larger than males in all body measurements and in four of nine cranial
measurements. Anderson (1960) found no significant differences in size between males
and females from Sinaloa.

Geographic variation. — San Luis Potosi material was found to be comparable in cranial
size to topotypes of A. 1. palmarum from Trinidad (Dalquest, 1950). Specimens from Girardot,
Mariquita, and Puente Nacional in the Magdalena River Valley, Colombia, averaged slightly
larger in body size than did those from two other localities: Mesitas del Colegio, at a higher
elevation on the western slope of the East Andes, and Villavicencio, at the base of the
eastern slope of the East Andes (Tamsitt and Valdivieso, 1963 a).

Artibeus phaeotis (Miller, 1902)

Measurements of Artibeus phaeotis have been recorded as follows: Miller (1902a), external
and cranial measurements of the female holotype from Yucatan; Elliot (1904), external and
cranial measurements of a single specimen; Andersen (19066), cranial measurements of the
female holotype of A. turpis (= A. phaeotis) from Tabasco and the female holotype of A. p.

BIOLOGY OF THE PHYLLOSTOMATIDAE

nanus from Guerrero; Andersen (1908), external and cranial measurements of the female
holotype of A. phaeotis from Yucatan, the holotype of A. jucundus ( — A. phaeotis) from
Veracruz, the female holotype of A. turpis ( = A. phaeotis) from Tabasco, and mean and range
of these measurements in eight specimens from Guerrero, Sinaloa, and Colima; Goodwin
(1934), external measurements of a specimen from Guatemala; Goodwin (1942 a), forearm
and cranial measurements of one specimen; Dalquest (1953/?), forearm and cranial measure¬
ments of a male and female from Veracruz; Jones and Lawlor (1965), external and cranial
measurements of a male and two females from Cozumel Island, Quintana Roo; Jones
(1966), forearm and cranial measurements (mean, range) of five specimens (three males, two
females) from El Peten, Guatemala, and for a male and female from Santa Rosa, Guatemala;
Villa-R. (1967), external measurements of 28 specimens and cranial measurements of 22 of
A. turpis turpis, which more or less include A. p. phaeotis and A. p. palatinus of Davis
(1970a), external measurements of 38 specimens and cranial measurements of 35 specimens
of A. p. nanus and two males and three females of A. cinerus phaeotis from Veracruz,
Oaxaca, and Tabasco; Rick (1968), external measurements of three females and one male,
and cranial measurements of three females, one male, and an unsexed specimen from
Guatemala; Goodwin (1969), forearm and cranial measurements of four males and nine
females from Oaxaca; Davis (1970a), cranial measurements (mean, range) of 135 specimens
from the Pacific versant of Sinaloa to Guerrero, 19 from Oaxaca to Chiapas, 37 from
Guatemala, El Salvador, and Nicaragua, 34 from the Pacific versant of Costa Rica and seven
from the Caribbean versant, 124 from the Caribbean versant of Guatemala and British
Honduras, 67 from Honduras and Nicaragua, and cranial measurements of the female
holotype of A. phaeotis phaeotis from Yucatan, the female holotype of A. p. nanus from
Guerrero, and the male holotype of A. p. palatinus from Guatemala; Jones et al. (1972),
forearm and cranial measurements (mean, range) of five males and five females combined
from Sinaloa.

Age variation. — Juveniles (cartilaginous epiphyses and unworn dental cusps) could not
be distinguished from adults on the basis of seven cranial measurements (Davis, 1970a).

Secondary sexual variation. — Davis (1970a) found no significant secondary sexual
dimorphism in four external and seven cranial measurements.

Geographic variation. — Davis (1970a) noted the following size variation throughout the
geographic range of this species. Members of the population in western Mexico (Sinaloa to
Guerrero) were generally the smallest for the species. The rostrum in this population was
short, which was reflected in the shortness of the palate. In the Pacific lowlands (Oaxaca
to Costa Rica), specimens had a longer palate, skull, and forearm; they were, however,
smaller than those from the Caribbean-Gulf versant. The population occupying the Caribbean-
Gulf versant (Veracruz to South America) was the largest in the species.

Artibeus toltecus(Saussure, 1860)

Measurements of Artibeus toltecus have been recorded as follows: Saussure (18606),
external measurements of a single specimen; Miller (1902a), external and cranial measure¬
ments of the male holotype of A. t. ravus from Ecuador and a specimen from Morelos;
Andersen (1908), external and cranial measurements (range) of three specimens from Costa
Rica, Nicaragua, and Guatemala, two (one cranial) from Oaxaca, nine (five cranial) from
Jalisco and Durango, and three from Veracruz, external measurements (mean, range) of
18 specimens (cranial of 13) from Costa Rica, Nicaragua, Guatemala, Jalisco, Durango,
Oaxaca, and Veracruz, and 11 specimens (mean, range) from Ecuador; Goodwin (1934),
external measurements of a specimen from Guatemala; Goodwin (1942a), external and
cranial measurements of two males from Honduras; Goodwin (1946), external and cranial
measurements (range) for the species; Dalquest (1953a), external measurements (mean) of
two males and cranial measurements (mean) of two males and five females from San Luis
Potosi; de la Torre (1954), external and cranial measurements (mean, range) of six specimens

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from Tamaulipas; de la Torre (1955), forearm measurements (mean, range) of five males
and three females combined from Jalisco; Felten (1956 d), external measurements of a
specimen from El Salvador; Jones et al. (1962), forearm and total length of skull (range) of
12 specimens from Mexico (Oaxaca 6, Tamaulipas 3, Jalisco 2, Sinaloa 1); Alvarez (1963),
external and cranial measurements of a male and two females from Tamaulipas; Jones and
Alvarez (1964), forearm measurements of a female and cranial measurements of this female
and a specimen of unknown sex from San Luis Potosf; Jones (1964), forearm and cranial
measurements of a specimen from Sinaloa; Jones (1966), forearm and cranial measurements
(mean, range) of six specimens (five males, one female) from Guatemala; Villa-R. (1967),
external measurements of 20 specimens and cranial measurements of 18 from Mexico;
Genoways and Jones (1968), forearm measurements of two males and four females from
Zacatecas; Davis (1969), forearm and cranial measurements (mean, range) of samples from
the Pacific versant including 14 from Sinaloa and Nayarit, 12 from Guerrero, 18 from
Chiapas, 18 from Guatemala, and 17 from the Honduran highlands, from the Atlantic
versant including nine from Tamaulipas and San Luis Potosi, eight from Veracruz, 16
from Chiapas, 14 from Guatemala, and 29 from the Costa Rican highlands, external and
cranial measurements of the male holotype of A. t. Hesperus from Guerrero and the male
neotype of A. t. toltecus from Veracruz; Goodwin (1969), forearm and cranial measurements
of four males and four females from Oaxaca; Jones et al. (19716), forearm and cranial
measurements (mean, range) of six specimens (three males, three females) from Departa-
mento de Matagalpa, Nicaragua, and external and cranial measurements of 10 specimens
(four males, six females) from Isla de Ometepe, Rivas, Nicaragua; Alvarez and Ramirez-
Pulido (1972), external and cranial measurements of two males and two females from
Morelos; Jones et al. (1972), forearm and cranial measurements (mean, range) of 10 specimens
(five males, five females) from Sinaloa.

Geographic variation. — According to Jones (1966), specimens from Guatemala averaged
larger than specimens from western Mexico. Davis (1969) showed that specimens from the
Pacific versant (El Salvador to Sinaloa) averaged smaller for almost all measurements
compared to those occupying the remainder of the species geographic range. Jones et al.
(19716) reported two size groups (subspecies) occurring in Nicaragua. Those of smaller size
from Isla de Ometepe, Rivas, and the others from Departamento de Matagalpa.

Artibeus watson i Thomas, 1901

Measurements of Artibeus watsoni have been recorded as follows: Thomas (1901 a),
forearm and cranial measurements of the male holotype and external measurements of
another male from Panama; Elliot (1904), external and cranial measurements of the holotype
(after Thomas, 1901); Elliot (1906), external and cranial measurements of the holotype of
Dermanura jucundum from Veracruz; Andersen (1908), external and cranial
measurements (mean, range) of nine specimens from Panama and Nicaragua; Sanborn
(1936), external measurements of two males and cranial measurements of one male from
Guatemala; Goodwin (1942u), external and cranial measurements of a single specimen;
Goodwin (19426), external and cranial measurements of the male holotype from Panama
and the range for these measurements in the species; Jones (1966), forearm and cranial
measurements of a male and female from Guatemala; Davis (1970a), cranial measurements
of the holotype, external and cranial measurements (mean, range) of 62 males and 46 females
from the Pacific versant of Costa Rica, and from the Atlantic versant 25 males and 19 fe¬
males from Costa Rica, 22 males and 17 females from Nicaragua, 11 males and four fe¬
males from Honduras, and eight males and four females from Guatemala, and cranial mea¬
surements (mean, range) of 120 specimens from southwestern Costa Rica (near type
locality).

Geographic variation. — Davis (1970a) considered Artibeus watsoni to be monotypic.

BIOLOGY OF THE PHYLLOSTOMATIDAE

Centurio senex Gray, 1 842

Measurements of Centurio senex have been recorded as follows: Lichtenstein and Peters
(1855), external measurements of the holotype of Centurio flavogularis’, Saussure (1860a),
external measurements of the female holotype of Centurio mexicanus from Mexico; H.
Allen (1861), external measurements of the holotype of Centurio mcmurtrii from Veracruz;
Dobson (1878a), external measurements of the female holotype; Ward (1891), external
measurements of the female holotype of Centurio minor from Veracruz and measure¬
ments given by Dobson ( 1 878c/); Rehn (1901), external measurements from the litera¬
ture including Dobson’s for C. senex , Lichtenstein's and Peters’ for C. flavogularis,
Saussure’s for C. mexicanus and Ward’s for C. minor, external measurements of five and
cranial of two specimens from Veracruz and external and cranial measurements of one
specimen from Costa Rica; Elliot (1904), external and cranial measurements of a specimen;
Sanborn (1936), external measurements (range) of 12 specimens and forearm and cranial
measurements (range) of 24 specimens from Guatemala; Goodwin (1942a), external and
cranial measurements (range) in the species; Goodwin (1946), forearm and cranial measure¬
ments (range) of 24 specimens from Guatemala (as given by Sanborn, 1936) and the holotype;
Felten (1956c), external and cranial measurements of a female from El Salvador; Felten
( 1 956z/), external measurements of a specimen from El Salvador; Hall and Kelson (1959),
forearm and cranial measurements (range) of specimens from Guatemala; Burt and Stirton
(1961), forearm and cranial measurements of a male from El Salvador; Goodwin and
Greenhall (1961), forearm measurements of four males and one female and cranial measure¬
ments of three males and one female from Trinidad; Alvarez (1963), external and cranial
measurements of a female from Tamaulipas; Villa-R. (1967), external and cranial measure¬
ments (mean, sd, range) of 10 specimens from Mexico; Paradiso (1967), forearm and cranial
measurements of the female holotype of C. s. greenhalli from Trinidad, forearm measurements
(mean, range) of 28 topotypes, cranial measurements of 1 1 topotypes, and forearm and
cranial measurements (mean, range) of 20 specimens of C. 5. senex from Panama, 1 1 from
Guatemala, and two from Oaxaca; Goodwin (1969), forearm and cranial measurements of
a male and female from Oaxaca; Jones et al. (19716), forearm and cranial measurements
(mean, range) of 11 specimens (seven males, four females) from Nicaragua; Jones et al.
(1972), external and cranial measurements of two males and one female from Sinaloa;
Watkins et al. (1972), forearm and cranial measurements of a male and five females (mean,
range) from Jalisco, and seven males and four females from Nicaragua.

Secondary sexual variation. — Females from Nicaragua averaged slightly larger than males
in both external and cranial measurements (Jones et al., 1971 6).

Geographic variation. — Specimens from Trinidad were clearly larger than those from
Panama, Guatemala, and Oaxaca in most measurements. No overlap in forearm measure¬
ments were found (Paradiso, 1967). Jones et al. (19716) reported that measurements of their
specimens from Nicaragua agreed in general with those given by Paradiso (1967) for
material from Panama. Specimens from Jalisco compare favorably in size with those from
the vicinity of the type locality (restricted by Goodwin, 1946) and elsewhere in Nicaragua
(Watkins et al., 1972).

Chiroderma doriae Thomas, 1891

Measurements of Chiroderma doriae have been recorded as follows: Thomas (18916),
forearm and cranial measurements for the species (material described by Dobson, 1878a,
as C. villosum is actually C. doriae and formed the basis for Thomas’ description); Goodwin
(1958), forearm and cranial measurements of the holotype from Brazil; Baker and Genoways
(1976), external and cranial measurements (mean, range) of 15 males and 21 females from
Brazil.

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Chiroderma iinprovisum Baker and Genoways, 1976

Baker and Genoways (1976) recorded external and cranial measurements of the male
holotype from Guadeloupe, Lesser Antilles.

Chiroderma salvini Dobson, 1878

Measurements of Chiroderma salvini have been recorded as follows: Elliot (1904), external
and cranial measurements of one specimen; Sanborn (1941), forearm measurements (range)
of 22 specimens and cranial measurements of three from Honduras; Goodwin (1942«),
external and cranial measurements of two males from Honduras; Goodwin (1946), external
and cranial measurements of two males from Honduras and one from Costa Rica; Goodwin
(1958), forearm and cranial measurements of a female from Costa Rica; Hall and Kelson
(1959), external and cranial measurements of a male from Costa Rica; Brosset (1965),
external and cranial measurements of a female from Ecuador; Handley (1965), external
and cranial measurements of two males and 1 1 females (mean, range) of C. s. scopaeum from
Chihuahua, Sinaloa, Nayarit, Jalisco, Colima, and Guerrero; Carter el al. (1966), external
and cranial measurements of a female from Guerrero and one from Honduras; Villa-R.
(1967), external and cranial measurements of a male from Costa Rica; Genoways and
Jones (1968), forearm measurements of five males from Zacatecas; Alvarez and Ramirez-
Pulido (1972), external and cranial measurements of one female from Puebla; Anderson
(1972), external and cranial measurements of two females from Chihuahua; Baker (1974),
forearm measurements of three specimens from Ecuador.

Geographic variation.- — Handley (1965) distinguished specimens from western Mexico
from typical members of the species in Costa Rica and Panama by their smaller size and
paler coloration.

Chiroderma trinitatum Goodwin, 1958

Measurements of Chiroderma trinitatum have been recorded as follows: Goodwin
(1958), external and cranial measurements of the female holotype from Trinidad; Handley
(1960), external and cranial measurements of the male holotype of C. gorgasi ( = C. trinitatum )
from Panama, a female paratype, and the female holotype of C. trinitatum from Trinidad;
Goodwin and Greenhall (1961), forearm and cranial measurements of the female holotype
from Trinidad; Ojasti and Linares (1971), external and cranial measurements of two females
from Venezuela; Pirlot (1972), forearm measurements of a single specimen from Brazil;
Gardner (1976), external and cranial measurements (mean, range) of two males and six
females from Peru.

Chiroderma villosum Peters, 1860

Measurements of Chiroderma villosum have been recorded as follows: Thomas (18916),
forearm and cranial measurements for the species; J. A. Allen (1900), external and cranial
measurements of the male holotype of C. villosum jesupi from Colombia; Miller (1912),
external and cranial measurements of the female holotype of C. isthmicum (=C. villosum
jesupi) from Panama; Elliot (1917), external and cranial measurements of the holotype of
C. isthmicunr, Sanborn (1936), forearm and cranial measurements of a male from Veracruz;
Goodwin (1946), external and cranial measurements of the female holotype of C. isthmicum ;
Goodwin (1953), forearm and cranial measurements of the male holotype of C. villosum
jesupi from Colombia; Goodwin (1958), forearm and cranial measurements of the holotype
of C. v. jesupi from Colombia, male holotype and female topotype of C. isthmicum from
Panama, and a male from Trinidad; Hall and Kelson (1959), external and cranial measure¬
ments of the holotype of C. isthmicum; Goodwin and Greenhall (1961), forearm and cranial
measurements of one male and three females from Trinidad; Husson (1962), external and
cranial measurements of a female from Surinam; Villa-R. (1962), cranial measurements of

BIOLOGY OF THE PHYLLOSTOMATIDAE

three specimens from Chiapas, two from Colima, and of the holotype of C. isthmicum ;
Davis et al. (1964), forearm measurements (range) of 12 females from Chiapas; Hill (1964),
forearm and cranial measurements of a female from Guyana; Villa-R. (1967), external and
cranial measurements of three females from Chiapas; Goodwin (1969), forearm and cranial
measurements of a female from Oaxaca; Gardner et al. (1970), forearm and cranial measure¬
ments of two males from Costa Rica; Birney et al. (1974), external and cranial measurements
of one male from Quintana Roo.

Geographic variation. — Husson (1962) found the measurements of his female from Surinam
to correspond well with those of the four specimens reported by Goodwin and Greenhall
(1961) from Trinidad. According to Birney et al. (1974), their male specimen corresponded
closely in size to a female reported by Goodwin (1969) from Oaxaca.

Ectophylla alba H. Allen, 1892

Measurements of Ectophylla alha have been recorded as follows: H. Allen (1892), external
measurements of the holotype from Nicaragua; H. Allen (1898), external measurements of
the holotype and of an Oldfield Thomas specimen; Goodwin (1946), external measurements
of the holotype from Nicaragua; Casebeer et al. (1963), external and cranial measurements
of three females from Costa Rica; Starrett and Casebeer (1968), forearm measurements of a
male and two females and cranial measurements of one male from Costa Rica; Gardner
et al. (1970), forearm measurements (eight males, two females) and cranial measurements
(mean, range) of seven males and two females from Costa Rica.

Enchisthenes hartii (Thomas, 1892)

Measurements of Enchisthenes hartii have been recorded as follows: Thomas (1892),
external and cranial measurements of the “slightly immature" male holotype from Trinidad;
Andersen (1908), external and cranial measurements of the male holotype from Trinidad;
Sanborn (1932), external and cranial measurements of a female from Venezuela; Goodwin
(1940, 1942, 1946), external and cranial measurements of a specimen from Honduras; de la
Torre (1955), forearm measurements (mean, range) of 12 specimens (eight males, four females),
and cranial measurements of one male and two females from Jalisco; Hall and Kelson (1959),
external and cranial measurements of a male from Honduras; Goodwin and Greenhall
(1961), forearm and cranial measurements of the holotype from Trinidad; Villa-R. (1967),
external measurements of a male and female from Jalisco; Baker and Lopez (1968), forearm
and cranial measurements of a male from Tamaulipas and a male and female from Trinidad;
Goodwin (1969), forearm and cranial measurements of a female from Oaxaca; LaVal (1969),
external and cranial measurements of one female from Honduras; Gardner et al. (1970),
forearm and cranial measurements (mean, range) of 13 specimens from Costa Rica; Gardner
(1976), external and cranial measurements of a female from Peru.

Geographic variation. — When comparing one male from Tamaulipas with a male and
female from Trinidad, Baker and Lopez (1968) concluded that no outstanding variation
was obvious.

Mesophy 11a (= Ectophylla) macconnelli Thomas, 1901

Measurements of Mesophylla macconnelli have been recorded as follows: Thomas
(19016), external measurements of the female holotype and one male and cranial measure¬
ments of the holotype from Guyana; Lima (1926), external measurements of a specimen from
Brazil; Cunha Vieira (1942), external and cranial measurements of a female from Brazil;
Sanborn (1951), forearm and cranial measurements of one specimen from Peru; Goodwin
and Greenhall (1962), external and cranial measurements of the female holotype of M. m.
flavescens from Trinidad, forearm and cranial measurements of one male and two females
(including the holotype of M. macconnelli) from Guyana, two males and three females
from Peru, one male from Brazil, and one male and two females from Ecuador; Starrett
and Casebeer (1968), forearm and cranial measurements of a female from Costa Rica.

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Phyllops falcatus (Gray, 1839)

Measurements of Phyllops falcatus have been recorded as follows: Gundlach (1872,
1877), external measurements of a specimen from Cuba; Dobson ( 1 878«), external measure¬
ments of the male holotype from Cuba; Elliot (1904), external and cranial measurements of
one specimen from Cuba; G. M. Allen (1942), external measurements for the species.

Phyllops haitiensis(J. A. Allen, 1908)

Measurements of Phyllops haitiensis have been recorded as follows: J. A. Allen (1908 a),
external measurements of the holotype of P. haitiensis from the Dominican Republic;
Elliot (1917), external and cranial measurements of the holotype; Sanborn (1941), external
measurements of two females and cranial measurements of one from Haiti; Goodwin
(1953), forearm and cranial measurements of the holotype from the Dominican Republic.

Pygoderma bilabiatum (Wagner, 1843)

Measurements of Pygoderma bilabiatum have been recorded as follows: Peters (1863),
external measurements of the holotype of Stenoderma ( Pygoderma ) microdon from Surinam;
Dobson (1878a), external measurements of one specimen; Elliot (1904), external and
cranial measurements of a single specimen; Lima (1926), external measurements of a specimen
from Brazil; Cunha Vieira (1942), external measurements of two females and two of unknown
sex and cranial measurements of a female from Brazil; Goodwin (1942, 1946), external
measurements of a specimen from Paraguay; Husson (1962), external and cranial measure¬
ments of two Brazilian specimens and several measurements of the male holotype of P.
microdon from Surinam, as given by Peters (1863).

Sphaeronycteris toxophyllum Peters, 1 882

Measurements of Sphaeronycteris toxophyllum have been recorded as follows: Peters
(1882), external measurements of the holotype from tropical America; Husson (1958),
external and cranial measurements of four males, five females, and one of unknown sex
from Venezuela.

Stenoderma rufum Desmarest, 1820

Measurements of Stenoderma rufum have been recorded as follows: Peters (1869), external
measurements of the holotype of S. r. rufum, Anthony (1918, 1925), cranial measurements
of fossil material from Puerto Rico; G. M. Allen (1942), cranial measurements of a single
specimen; Hall and Bee (1960), external measurements of the holotype from an unknown
locality and external and cranial measurements of a male and female from St. John Island;
Tamsitt and Valdivieso (1966c), external measurements of a female and her one-day-old
young (male) from Puerto Rico; Choate and Birney (1968), cranial measurements of 10
specimens of sub-Recent material from Puerto Rico (type description of S. r. anthonyi),
six specimens of Recent material from Puerto Rico, and two specimens from St. John;
Hall and Tamsitt (1968), external and cranial measurements of the female holotype of S. r.
darioi from Puerto Rico, and the mean and range of these measurements in three males and
four females; Jones et al. (1971a), external and cranial measurements (mean, sd, range) of
15 males and seven females from Puerto Rico, and one male and female from St. John;
Genoways and Baker (1972), external measurements (mean, range) of 14 males and six
females and cranial measurements of 15 males and seven females from Puerto Rico (from
Jones et al., 1971a).

Individual variation. — Forearm and cranial measurements of specimens with a greyish
pelage and unfused or incompletely fused phalangeal epiphyses (immature) were significantly
smaller than adults (Jones et al., 197 1 a).

BIOLOGY OF THE PHYLLOSTOMATIDAE

Secondary sexual variation. — According to Choate and Birney (1968), females were
larger than males in material from Puerto Rico and St. John Island. Indications also exist that
this was true in sub- Recent material. Jones et al. (1971 a) found females significantly larger
than males in all external and cranial measurements tested.

Geographic variation. — Hall and Bee (1960) stated that cranial dimensions of Puerto
Rican specimens were larger than those from St. John. Sub-Recent material from Puerto
Rico was larger throughout than the Recent material from Puerto Rico and St. John (Choate
and Birney, 1968).

Hall and Tamsitt (1968) assigned specimens from St. John Island and St. Thomas Island
to S. r. rufum because they closely resembled the holotype. They named a new subspecies
from Puerto Rico on the basis of external color, although they found no differences between
the two in overall size or shape and size of skull.

Jones et al. (1971a) confirmed that Stenoderma rufum was a polytypic species with three
distinct subspecies. Recent Puerto Rican specimens were characterized by marked secondary
sexual dimorphism and by darker color than the other Recent race from the Virgin Islands;
subfossil material from Puerto Rico was distinguished by larger size and several details of
dentition.

Sturnira aratathomasi Peterson and Tamsitt, 1968

Measurements of Sturnira aratathomasi have been recorded as follows: Peterson and
Tamsitt (1968), external and cranial measurements of the male holotype from Colombia and
a male and female from Ecuador; Thomas and McMurry (1974), external and cranial
measurements of the holotype and three males and three females from Colombia.

Sturnira bidens (Thomas, 1915)

Measurements of Sturnira bidens have been recorded as follows: Thomas (1915), external
and cranial measurements of the immature male holotype from Ecuador; Gardner and
O’Neill (1969), forearm and cranial measurements (mean, range) of six specimens from
Peru and the holotype from Ecuador; Gardner and O’Neill (1971), forearm and cranial
measurements (mean, range) of 11 specimens from Peru; Marinkelle and Cadena (1972),
forearm measurements (range) of two males and seven females and cranial measurements
(range) of two males and four females from Colombia.

Geographic variation. — Marinkelle and Cadena (1972) found that their specimens from
Colombia averaged slightly larger in cranial measurements than those from Peru reported
by Gardner and O'Neill (1969).

Sturnira erythromos(Tschudi, 1844)

Measurements of Sturnira erythromos have been recorded as follows: Gardner et al.
(1969), forearm and cranial measurements (mean, range) of 24 specimens from Peru; Tuttle
(1970), forearm measurement range in species.

Sturnira lilium (E. Geoffroy St. -Hilaire, 1810)

Measurements of Sturnira lilium have been recorded as follows: Dobson (1878a), external
measurements of one male; Cabrera (1903), external measurements for the species in Chile;
Elliot (1904), external and cranial measurements of a specimen; Goldman (1917), external
and cranial measurements of the female holotype S. I. parvidens from Guerrero; Lima
(1926), external measurements of a male from Brazil; Cunha Vieira (1942), external measure¬
ments of five males and three females and cranial measurements of four males from Brazil;
Goodwin (1942a), external and cranial measurements of the holotype of S. I. parvidens and
a male and female from Honduras; Goodwin (1946), external and cranial measurements of

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one male from Honduras, also given by Goodwin (1942a); Hershkovitz (1949), external and
cranial measurements of a male and female from northern Colombia; Dalquest (1953 a),
external measurements (mean) of three males and seven females combined, and cranial
measurements (mean) of three males and five females combined from San Luis Potosi;
de la Torre (1954), external and cranial measurements of two specimens from Tamaulipas;
Felten (1956c), external and cranial measurements of a female from El Salvador; Felten
(1956 d), external measurements of a specimen from El Salvador; Hall and Kelson (1959),
forearm measurements (mean) of 12 topotypes from Paraguay; Goodwin and Greenhall
(1961), forearm and cranial measurements of a male and female from Trinidad and two males
from Paraguay; Husson (1962), external and cranial measurements of one male and four
females from Surinam; Pirlot (1963), external measurements of seven males and seven females
from Venezuela and cranial measurements of one female; Tamsitt and Valdivieso (1963a),
external measurements of three males and one female and cranial measurements of one
female from Colombia; Tamsitt and Valdivieso (19636), external measurements of two
males from Colombia; Starrett and de la Torre (1964), external and cranial measurements
of a male and two females from El Salvador and one female from Nicaragua; Valdivieso
(1964), external and cranial measurements of a specimen from Colombia; de la Torre
(1966), external and cranial measurements of the male holotype and the mean and range
of four male and five female paratypes combined of S. 1. angeli from Dominica, Lesser Antil¬
les; de la Torre and Schwartz (1966), external and cranial measurements of the female holo¬
type of S. I. paulsoni from St. Vincent, Lesser Antilles; Villa-R. (1967), external and cranial
measurements (mean, sd, range) of nine specimens from Mexico; Pirlot (1968), forearm
measurement of a female from Peru; Goodwin (1969), forearm and cranial measurements
of four males and five females from Oaxaca; Villa-R. and Villa Cornejo (1969), external and
cranial measurements (mean, range) of 15 specimens from Argentina; Anderson (1972),
external measurements of one adult specimen and cranial measurements of two from
Chihuahua; Jones et al. (1973), greatest length of skull (mean, range) of three males and five
females combined from the Yucatan Peninsula; Taddei (19756), external measurements
(mean, se, range) of 20 males and 20 females and cranial measurements of 15 males and
15 females from Brazil; Jones and Phillips (1976), forearm and cranial measurements (mean
and range of sexes combined) from four Lesser Antillean islands — Dominica, two males
and 12 females; Martinique, four males and four females; St. Lucia, four males and three
females; and St. Vincent, three males.

Individual variation. — In specimens from Brazil, coefficients of variation for external
measurements varied in 20 males from 2.85 to 5.86 and in 20 females from 2.48 to 7.08;
CV values for cranial measurements in 15 males ranged from 1.47 to 3.57 and in 15
females from 1.75 to 3.01 (Taddei, 19756).

Secondary sexual variation. — Although males generally averaged larger than females in
specimens from Brazil, no significant differences in external measurements were found. How¬
ever, in 15 cranial measurements, only two (braincase breadth, cranial depth) did not differ
significantly (Taddei, 19756).

Geographic variation. — Comparing Mexican material with species from Paraguay,
Goldman (1917) concluded that the forearm was shorter in most of the specimens available
from Mexico and that the skull was narrower. Goodwin (1942a) stated that size in a Honduran
series, including both males and females, was smaller than specimens from Mexico.
Jones et al. (1973) noted that the greatest length of skull of a specimen from La Tuxpena,
Campeche, which Goldman reported (1917) to be abnormally small, fell within the range
of that observed for three males and five females combined from the Yucatan Peninsula —
their specimens averaged only slightly smaller than specimens from adjacent Chiapas and
Guatemala. Jones and Phillips (1976) stated that Antillean S. I ilium generally fell within the
size range of populations of this species from Middle and South America. They did find
some variation between insular samples, although no clinal geographic trend could be

BIOLOGY OF THE PHYLLOSTOMATIDAE

demonstrated. Bats from St. Vincent tended to be the largest cranially among Antillean
populations, whereas specimens from Martinique had proportionally broader zygomatic
arches and longer maxillary toothrows. Forearm length in specimens from Dominica
averaged slightly larger than did specimens from other islands. No other differences in
external proportions were demonstrated.

Sturnira ludovici Anthony, 1924

Measurements of Sturnira ludovici have been recorded as follows: Anthony (1924/?),
external and cranial measurements of the male holotype from Ecuador; Shamel (1927),
external and cranial measurements of the female holotype of S. I. bogotensis ( = S. ludovici)
from Colombia; Goodwin (1940), external and cranial measurements of the female holotype
of 5. hondurensis (=5. ludovici) from Honduras; Goodwin (1942a), external and cranial
measurements of two specimens from Honduras; Goodwin (1946), forearm and cranial
measurements of the holotype of S. hondurensis, and a male from Costa Rica; Hershkovitz
(1949), external and cranial measurements of the holotype of 5. /. bogotensis and the range
of these measurements in two males and two females combined from Colombia; de la Torre
(1952), external and cranial measurements of a male and female from Michoacan; Dalquest
(1953 a), external measurements (mean) of three males and cranial measurements of one of
unknown sex, from San Luis Potosi; Goodwin (1953), external and cranial measurements
of the holotypes of S. ludovici and S. hondurensis ; Lukins and Davis (1957), external and
cranial measurements of a female from Guerrero; Baker and Greer (1962), external and
cranial measurements of one male and two females from Durango; Tamsitt and Valdivieso
(1963a), external and cranial measurements (mean, range) of six males and six females
combined from Colombia; Jones and Phillips (1964), external and cranial measurements
of the female holotype of S. I. occidentalis from Sinaloa, mean and range of these measure¬
ments for specimens from Durango and Jalisco (S. /. occidentalis), Puebla, Michoacan,
Oaxaca, Honduras, Colombia (after Hershkovitz, 1949), and Ecuador (S. /. ludovici)-,
Starrett and de la Torre (1964), external and cranial measurements of a male and female from
Costa Rica; Valdivieso (1964), external and cranial measurements (mean, range) of specimens
from Colombia; Jones and Dunnigan (1965), forearm and cranial measurements of 12
males and 15 females (mean, range) from Oaxaca; Villa-R. (1967), external and cranial
measurements of five specimens from Mexico; Goodwin (1969), forearm and cranial
measurements of eight males and one female from Oaxaca; Jones et al. (19716), external
and cranial measurements of one male from Nicaragua; Jones et al. (1972), forearm and
cranial measurements of the female holotype of S. 1. occidentalis and three males from
Sinaloa.

Secondary sexual variation. — Jones and Dunnigan (1965), examining the mean and
extremes of forearm and six cranial measurements, suggested that males average slightly
larger than females.

Geographic variation. — Lukins and Davis (1957) concluded that their female specimens
from Guerrero were somewhat smaller than those recorded by Hershkovitz (1949) from
Colombia and Dalquest (1953a) from San Luis Potosi but corresponded closely to one
regarded as S. hondurensis from Costa Rica (Goodwin, 1946). Jones and Phillips (1964)
found specimens in the northern part of the range of the species to be smaller than speci¬
mens from Central America and northern South America and described them as S. I.
occidentalis.

Sturnira niagna de la Torre, 1966

Measurements of Sturnira magna have been recorded as follows: de la Torre (1966),
external and cranial measurements of the male holotype and mean and range of five male
and three female paratypes from Peru; Peterson and Tamsitt (1968), external and cranial

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measurements of the male holotype, mean and range of five males and three females (after
de la Torre, 1966), and two females from Peru; Marinkelle and Cadena (1972), external
measurements of one specimen from Colombia; Baker (1974), forearm measurement
of a female from Ecuador; Gardner (1976), external and cranial measurements (mean, range)
of one male and three females from Peru.

Sturnira mordax (Goodwin, 1938)

Measurements of Sturnira mordax have been recorded as follows: Goodwin (1938, 1946),
external and cranial measurements of the male holotype from Costa Rica; Hall and Kelson
(1959), external and cranial measurements of the holotype; Davis et al. (1964), external
and cranial measurements of six males and two females from Costa Rica; Gardner et al.
(1970), forearm and cranial measurements (mean, range) of 12 specimens from Costa Rica.

Sturnira nana Gardner and O’Neill, 1971

Gardner and O’Neill (1971) recorded external and cranial measurements of the female
holotype and forearm and cranial measurements (mean, range) of five other specimens
from Peru.

Sturnira tildae de la Torre, 1959

Measurements of Sturnira tildae have been recorded as follows: de la Torre (1959),
external and cranial measurements of the male holotype and a female paratype from
Trinidad; Goodwin and Greenhall (1961), forearm and cranial measurements of two males
and two females from Trinidad; Hill (1964), external and cranial measurements of two
females from Guyana; Marinkelle and Cadena (1971), external measurements of 60 males
and 60 females from Colombia (mean, range), male holotype and female paratype from
Trinidad (after de la Torre, 1959), two females from Guyana (after Hill, 1964), and cranial
measurements of 50 males and 50 females from Colombia (mean, range), one male and
five females from Guyana, holotype, paratype, and three females from Trinidad.

Geographic variation. — Marinkelle and Cadena (1971) found external measurements
of Colombian specimens generally averaged larger than the holotype and paratype from
Trinidad.

Sturnira thomaside la Torre and Schwartz, 1966

Measurements of Sturnira thomasi have been recorded as follows: de la Torre and
Schwartz (1966), external and cranial measurements of the male holotype from Guadeloupe,
Lesser Antilles; Genoways and Jones (1975), external and cranial measurements of the
male holotype (after de la Torre and Schwartz, 1966) and four females (including one juve¬
nile) from Guadeloupe; Jones and Genoways (1975), external and cranial measurements
(after Genoways and Jones, 1975); Jones and Phillips (1976), external and cranial measure¬
ments of the same individuals as given by Genoways and Jones (1975).

Uroderma bilobatum Peters, 1866

Measurements of Uroderma bilobatum have been recorded as follows: Peters (1866a),
external measurements of a single specimen; Dobson (1878a), external measurements of
one specimen; Rehn (1900), cranial measurements of a specimen from Brazil; Lyon (1902a),
external and cranial measurements of the female holotype of U. b. convexum from Panama
and a specimen from Brazil; Elliot (1904), external and cranial measurements of the holo¬
type of U. b. convexum (after Lyon, 1902a) from Panama; Andersen (19066), measure¬
ments (range) of two specimens, including the male holotype of U. b. thomasi, from

BIOLOGY OF THE PHYLLOSTOMATIDAE

Bolivia; Andersen (1908), external and cranial measurements (range) of one specimen from
Brazil, one from Amazonas, two from Peru, one from Ecuador, one from Cali, Colombia,
three from Santa Marta, Colombia, and Valencia, Venezuela, two from Colon, Panama,
two from Chiriqui, Panama, nine (eight cranial) from the islands off Panama, and one from
Costa Rica; Lima (1926), external measurements of a male from Brazil; Cunha Vieira
(1942), external and cranial measurements of a male from Peru; Goodwin (1946), external
and cranial measurements of two males from Costa Rica; Hershkovitz (1949), external
and cranial measurements (range) of specimens from Colombia; Sanborn (1951), greatest
length of skull of one female from Peru; Felten ( 1956c), external measurements of a male
and four females and cranial measurements of one male and two females from El Salvador;
Felten (1956^0, external measurements (mean, range) of specimens from El Salvador; Hall
and Kelson (1959), external and cranial measurements of two males from Costa Rica; Burt and
Stirton (1961), forearm and cranial measurements (range) of 16 males and 13 females
from El Salvador; Goodwin and Greenhall (1961), external measurements of a subadult
male and four females and cranial measurements of the subadult male and two females
from Trinidad; Husson (1962), external and cranial measurements of four females from
Surinam; Tamsitt and Valdivieso (1963 a), external measurements (mean, range) of nine
males and five females combined from Colombia; Valdivieso (1964), external and cranial
measurements (mean, range) of one male and nine females combined from Colombia;
Brosset (1965), external and cranial measurements of one female from Ecuador; Villa-R.
(1967), external measurements (mean, sd, range) of 22 specimens and cranial measure¬
ments of 20 from Chiapas; Davis (1968), forearm and cranial measurements of the holotype
(juvenile, unsexed) of U. b. bilobatum from Brazil, 18 males and 30 females from Bolivia,
eastern Brazil, Cayenne, Guyana, and Venezuela, external and cranial measurements of the
male holotype of U. b. trinitatum, mean and range of eight males, and five females from
Trinidad, a male paratype of U. b. thomasi from Bolivia, 21 males and 14 females from
Ecuador, Peru, and western Bolivia, the female holotype (young) of U. b. convexum from
Panama, 77 males, and 124 females from western Venezuela, Colombia, Panama (exclusive
of the Bocas del Toro region), the Pacific versant of Middle America as far as Oaxaca,
the male holotype of U. b. molaris from Chiapas, 36 males and 58 females from the
Atlantic versant of Middle America from the Bocas del Toro region of Panama north¬
ward to southern Veracruz; Goodwin (1969), forearm and cranial measurements of one
male and two females from Oaxaca and one subadult male and two females of Uroderma sp.
from Oaxaca; Baker and McDaniel (1972), forearm and cranial measurements of the female
holotype of U. b. davisi from El Salvador, forearm and cranial measurements (mean,
sd) of 16 males and 10 females from Chiapas and El Salvador (U. b. davisi), 33 males
and 29 females from Nicaragua, Costa Rica, and Colombia ( U. b. convexum), and 25
males and 26 females from Tabasco, Honduras, Nicaragua, and Costa Rica ( U. b. molaris).

Secondary sexual variation. — Baker et al. (1972c;) described sexual dimorphism in this
species with males larger than females.

Geographic variation. — According to Davis (1968), specimens from Trinidad ( U . b.
trinitatum) were noticeably larger than those from the adjacent mainland ( U. b. bilobatum)
but were difficult to separate from specimens from Ecuador, Peru, and western Bolivia
( U. b. thomasi). Specimens from western Bolivia were larger than specimens from Colombia
and the Pacific versant of Central America ( U. b. convexum). U. b. convexum, again, was
smaller in most measurements than specimens from Bolivia, eastern Brazil, the Guianas,
and Venezuela ( U. b. bilobatum). Specimens from the Atlantic versant of Middle America
(U. b. molaris) from Bocas de Toro, Panama, northwest to Veracruz, Mexico, were of
moderate size for the species. Uroderma b. davisi from the Pacific versant of Middle Ameri¬
ca (Chiapas, El Salvador, Honduras) averaged smaller both externally and cranially than
either convexum or molaris (Baker and McDaniel, 1972).

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Uroderma magnirostrum Davis, 1968

Measurements of Uroderma magnirostrum have been recorded as follows: Davis
(1968), external and cranial measurements of the male holotype from Honduras and
26 males and 51 females (mean, range) from Oaxaca, Chiapas, El Salvador, Honduras,
Nicaragua, Panama, Colombia, Peru, Bolivia, Venezuela, and Brazil; Jones et al. (1971 6),
external and cranial measurements of one male and two females from Nicaragua.

Geographic variation. — Davis (1968) found little evidence of geographic variation but
his findings were based on relatively small sample sizes of U. magnirostrum.

Vampyressa bidens (Dobson, 1878)

Measurements of Vampyressa bidens have been recorded as follows: Dobson (1878 z/),
external measurements of the female holotype from Peru; Sanborn (1936), forearm
measurements (range) of two males and one female, wing measurements of one male and
one female from Ecuador, cranial measurements of a male and female from Ecuador and
the range of these measurements in three males and one female from Peru; Cunha Vieira
(1942), external measurements of a male and female and cranial measurements of a male
from Brazil; Hill (1964), forearm and cranial measurements of four males and one female
from Guyana; Marinkelle and Cadena (1972), external and cranial measurements of one
female from Colombia; Davis (1975), external and cranial measurements of 13 males
and 10 females (mean, sd, range) from Peru.

Individual variation. — Coefficients of variation, as given by Davis (1975), varied from
1.28 in greatest length of skull in females to 3.27 in postorbital constriction of females.
The two external measurements, which were tested, fell within this range.

Secondary sexual variation. — Comparing two external and eight cranial measurements
of 13 males with those of 10 females showed no significant differences. Females generally
averaged larger than males (Davis, 1975).

Vampyressa brocki Peterson, 1968

Measurements of Vampyressa brocki have been recorded as follows: Peterson (1968),
external and cranial measurements of the female holotype from Guyana; Baker et al.
(19726), external and cranial measurements of three females from Colombia; Peterson
(1972), external and cranial measurements of the holotype and a male from Guyana;
Davis (1975), forearm and cranial measurements (range) of published data.

Vampyressa melissa Thomas, 1926

Measurements of Vampyressa melissa have been recorded as follows: Thomas (1926),
external and cranial measurements of the female holotype from Peru; Goodwin (1963),
forearm and cranial measurements of the female holotype; Peterson (1968), forearm and
cranial measurements of one specimen; Gardner (1976), external and cranial measurements
of four specimens (one male, three females) from Peru.

Vampyressa nymphaea Thomas, 1909

Measurements of Vampyressa nymphaea have been recorded as follows: Thomas (1909),
forearm and cranial measurements of the male holotype from Colombia; Hall and Kelson
(1959), forearm and cranial measurements of the holotype and external measurements of a
specimen from Panama; Goodwin (1963), forearm and cranial measurements of two males
from Colombia and two females from Panama; Peterson (1968), forearm and cranial
measurements (range) in specimens of the species; Gardner et al. (1970), forearm and
cranial measurements (mean, range) of five specimens (three males, two females) from
Costa Rica; Jones et at. (19716), external and cranial measurements of one female from
Nicaragua.

BIOLOGY OF THE PHYLLOSTOMATIDAE

Vampyressa pusilla (Wagner, 1843)

Measurements of Vampyressa pusilla have been recorded as follows: Peters ( 1 866«),
external measurements of a specimen from Brazil; Dobson (1878a), external measurements
of one specimen from Brazil; Thomas (1909), forearm and cranial measurements of the male
holotype of V. p. thy one from Colombia; Miller (1912), external and cranial measurements
of the immature female holotype of V. minuta (= V. pusilla) from Panama; Elliot (1917),
external and cranial measurements of the holotype of V. minuta-, Cunha Vieira (1942),
external measurements of a specimen from Brazil; Goodwin (1946), external and cranial
measurements of the female holotype of V. minuta from Panama and those of a male from
Costa Rica; Hershkovitz (1949), external and cranial measurements of one female from
Colombia; Sanborn (1953), forearm and cranial measurements (range) of two males and
one female from Peru; Hall and Kelson (1959), cranial measurements of the holotype of
V. p. thyone; Davis et al. (1964), external and cranial measurements of a female from Chiapas;
Goodwin (1963), external and cranial measurements of the male holotype of V. pusilla from
Brazil, the male holotype of V. nattereri (— V. pusilla ) from Brazil, and forearm and cranial
measurements of the female holotype of V. p. venilla from Peru, three females from Panama,
two males from Costa Rica, one male and three females from Colombia, two males and
one female from Ecuador, five males and five females from Peru, and one female from
Venezuela; Starrett and de la Torre (1964), external and cranial measurements of one female
from Nicaragua; Peterson (1965a), external and cranial measurements of a female from
British Honduras; Tamsitt and Valdivieso (1966a), forearm and cranial measurements of a
male and female from Colombia (the latter as given by Hershkovitz, 1949); Rick (1968),
external and cranial measurements of one male and female from Guatemala; Gardner
et al. (1970), forearm and cranial measurements (mean, range) of five specimens (one male,
four females) from Costa Rica; Jones et al. (19716), forearm and cranial measurements of
two males and mean and range of six females from Nicaragua; Baker et al. (1973), external
and cranial measurements of 36 specimens from Colombia, Ecuador, and Venezuela, four
specimens from the Darien of Panama, 14 from the remainder of Panama, and seven from
Nicaragua; Jones et al. (1973), external and cranial measurements of one female from
Campeche.

Individual variation. — Baker et al. (1973) found coefficients of variation for forearm
and cranial measurements in four samples from Central and South America ranged
between 1.5 and 7.2. Lowest values were for breadth across upper molars in the sample
from the Darien of Panama and postorbital breadth in the sample from Nicaragua; the
highest CV value was for postorbital breadth in the sample from the Darien of Panama.
All samples had coefficients of variation exceeding 4.0 for palatal length.

Geographic variation. — Goodwin (1963), in his review of the genus, recognized three
subspecies of V. pusilla. These were based primarily on minor details of coloration and
slight size differences. Handley (19666) believed that the subspecific variations noted by
Goodwin could be attributed to variation with age and chose to consider V. pusilla as
being monotypic. Two years later, Peterson (1968) recognized two subspecies — one from
southeastern Brazil and the other occupying the remainder of the geographic range of
the species in South and Central America. He did not give, however, the characteristics
used to distinguish them.

Starrett and de la Torre (1964) concluded that their female specimen from Nicaragua
was similar in size to measurements given by Goodwin (1946) for the holotype of V. minuta
(= V. pusilla) from Panama and for a specimen from Costa Rica. They also found their
specimen from Nicaragua indistinguishable from three specimens from Peru.

Baker et al. (1973) found no significant differences in forearm and cranial measurements
of specimens from four geographic areas including Colombia, Ecuador, Venezuela, the
Darien and remainder of Panama, and Nicaragua.

Jones et al. (1973) followed Handley (19666) in considering V. pusilla monotypic when
assigning their specimen from Campeche.

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Vampyrodes caraccioli (Thomas, 1889)

Measurements of Vampyrodes caraccioli have been recorded as follows: Thomas (1889),
external and cranial measurements of the holotype from Trinidad; G. M. Allen (1908),
external and cranial measurements of the female holotype of V. major from Panama;
Sanborn (1936), forearm and cranial measurements (range) of two males, one female,
and one unsexed specimen, and wing measurements of one male from Guatemala; Sanborn
(1941), external and cranial measurements of a male from Trinidad; Goodwin (1942«),
external and cranial measurements of the female holotype of V. major from Panama;
Goodwin (1946), external and cranial measurements of the holotype of V. major (as in
Goodwin, 1942) and of one specimen from Nicaragua; Husson (1954), external and cranial
measurements of four males from Tobago; Hall and Kelson (1959), cranial measurements
of a male from Guatemala; Goodwin and Greenhall (1961), forearm and cranial measure¬
ments of the unsexed holotype from Trinidad and a female from Tobago; Villa-R. (1967),
external and cranial measurements of two males and one female from Veracruz; Starrett
and Casebeer (1968), forearm measurements of three males and nine females, and cranial
measurements of three males and two females from Costa Rica; Goodwin (1969), forearm
and cranial measurements of one male from Oaxaca; Linares (1969), external and cranial
measurements of one specimen from Venezuela; Gardner et al. (1970), forearm measure¬
ment of a female from Costa Rica.

Geographic variation. — According to Sanborn (1936), his series of specimens from
Guatemala agreed closely in measurements with the original description of V. major
from Danama. Gardner et al. (1970) noted that the forearm length of their female from
Costa Rica greatly exceeded the range for three males and nine females recorded by Starrett
and Casebeer (1968) from Costa Rica.

Vampyrops aurarius Handley and Ferris, 1972

Measurements of Vampyrops aurarius have been recorded as follows: Handley and Ferris
(1972), external and cranial measurements of the male holotype from Venezuela; Carter
and Rouk (1973), forearm and cranial measurements of the male holotype from Venezuela
and the mean and range for Peruvian specimens.

Vampyrops brachycephalus Rouk and Carter, 1972

Measurements of Vampyrops brachycephalus have been recorded as follows: Rouk and
Carter (1972), external and cranial measurements of the male holotype from Huanuco,
Peru and mean and range for 13 specimens from Loreto, Peru, six from Huanuco, Peru,
three from Colombia, and 13 from Venezuela; Gardner and Carter (19726), external and
cranial measurements of the male holotype and measurements (mean, range) of 13 specimens
from Loreto and six specimens from Huanuco, Peru (see also Rouk and Carter, 1972);
Handley and Ferris (1972), external and cranial measurements of the male holotype of
V. latus 1 = V. brachycephalus) from Peru and similar measurements for the male holotype of
V. latus saccharus from Venezuela; Carter and Rouk (1973), forearm and cranial measure¬
ments of the holotype of V. latus and V. latus saccharus as well as mean and range of these
measurements for 13 specimens from Loreto, Peru, and an unspecified number of specimens
from Tingo Maria, Peru.

Vampyrops dorsalis Thomas, 1900

Measurements of Vampyrops dorsalis have been recorded as follows: Thomas (1900),
external and cranial measurements of the holotype from Ecuador; Lyon (19026), external
and cranial measurements of the female holotype of V. umbratus from Colombia; Thomas
(1914), external and cranial measurements of the male holotype of V. oratus from Venezuela;
Sanborn (1951), forearm and cranial measurements of the holotype and a male from Peru;

BIOLOGY OF THE PHYLLOSTOMATIDAE

Sanborn (1955), external measurements of two males and cranial measurements (range) of
10 specimens (eight males, one female, one unsexed) from Colombia, Ecuador, Peru,
and Venezuela; Tamsitt and Valdivieso (1966a), forearm and cranial measurements (range)
of four males from Colombia, and those given by Sanborn (1955), Handley and Ferris
(1972), external and cranial measurements of the female holotype of V. aquilus from
Panama; Gardner and Carter (19726), external and cranial measurements of the immature
male holotype from Ecuador and mean and range for one specimen from Ecuador and
eight from Peru; Carter and Rouk (1973), forearm and cranial measurements of the holo¬
type of V. aquilus (= V. dorsalis) as reported by Handley and Ferris (1972) and mean and
range for specimens from Peru of V. dorsalis reported by Gardner and Carter (19726).

Vampyrops helleri Peters, 1866

Measurements of Vampyrops helleri have been recorded as follows: Peters (1866a),
external measurements of the holotype from Mexico; Dobson (1878a), measurements of
one specimen from Mexico; H. Allen (1891), external and cranial measurements of the
female holotype of Vampyrops zarhinus from Brazil (holotype now considered to be from
Panama according to Jones and Carter, 1976); Robinson and Lyon (1901), external measure¬
ments of four females from Venezuela; Elliot (1904), external and cranial measurements of
one specimen; Thomas (1912a), external and cranial measurements of the male holotype
of V. incarum from Peru; Cunha Vieira (1942), external measurements of a male and female
and cranial measurements of a male of Vampyrops zarhinus (=V. heller i) from Brazil;
Goodwin (1942a), external and cranial measurements of a single specimen; Goodwin
(1946), forearm and cranial measurements of one female from Costa Rica; Sanborn
(19496), forearm measurement of one female and cranial measurements of two females
from Peru; Sanborn (1955), external and cranial measurements (range) of specimens from
Oaxaca, Honduras, Costa Rica, Panama, Cayenne, Trinidad, Brazil, Venezuela, Colombia,
and Peru; Sherman (1955), external measurements of a male from Paraguay; Hall and
Kelson (1959), forearm and cranial measurements of one female from Costa Rica; Goodwin
and Greenhall (1961), external and cranial measurements of one male and three females
from Trinidad; Husson (1962), external and cranial measurements of eight males from
Surinam; Tamsitt and Valdivieso (1963a), external measurements of three males and one
female and cranial measurements of three males from Colombia; Starrett and de la Torre
(1964), external and cranial measurements of a female from Costa Rica; Davis et al.
(1964), external and cranial measurements (mean, range) of six specimens from Chiapas
and Central America; Valdivieso (1964), external measurements of one specimen from
Colombia; Villa-R. (1967), external and cranial measurements of a male and two females
from Oaxaca, Chiapas, and Tabasco; Rick (1968), external and cranial measurements of
a male and female from Guatemala; Goodwin (1969), forearm and cranial measurements
of one female from Oaxaca; Gardner and Carter (19726), external measurements of the
holotype (sex unknown) from Mexico, and external and cranial measurements (mean,
range) of four specimens from Peru; Rouk and Carter (1972), forearm and cranial measure¬
ments (mean, range) of four specimens from Peru, one from Ecuador, nine from Colombia,
three from Venezuela, one from Panama, two from Costa Rica, 20 from Nicaragua, and
12 from Honduras.

Vampyrops infuscus Peters, 1880

Measurements of Vampyrops infuscus have been recorded as follows: Peters (1880),
external measurements of the holotype from Peru; Miller (1902a), external and cranial
measurements of the female holotype of V. fumosus from Brazil; Sanborn (1936), forearm
and cranial measurements (range) of three males and one female from Ecuador; Cunha
Vieira (1942), external measurements of the holotype of V. fumosus based on Miller
(1902a); Sanborn (1951), forearm measurements of the holotype of V. infuscus from Brazil

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and a series of specimens from Peru, Ecuador, and Colombia; Marinkelle (1970), external
and cranial measurements of the female holotype of V. intermedius from Colombia and the
range of these measurements in the paratypes (five males, ten females); Gardner and Carter
(19726), external and cranial measurements of the adult male neotype of V. infuscus and
the mean and range of several external and cranial measurements of six specimens, including
the neotype from Peru.

Secondary sexual variation. — Marinkelle (1970) found no significant differences in size
between five males and 10 females from Colombia.

Vampyrops lineatus(E. Geoffroy St.-Hilaire, 1810)

Measurements of Vampyrops Uneatus have been recorded as follows: Dobson (1878 z/),
external measurements of the holotype; H. Allen (1891), external and cranial measurements
of one specimen; Elliot (1904), external measurements of a single specimen; Lima (1926),
external measurements of a specimen from Brazil; Cunha Vieira (1942), external measure¬
ments of three males, three females, and one unsexed specimen, and cranial measurements
of three males and one female from Brazil; Goodwin (1946), external and cranial measure¬
ments of a male from Paraguay; Hershkovitz (1949), external measurements of four males
and a female and cranial measurements of one male from Colombia; Sanborn (1955),
external measurements of one male and seven females and cranial measurements of an
unspecified number of specimens from Brazil, Paraguay, and Bolivia.

Vampyrops nigellus Gardner and Carter, 1972

Gardner and Carter ( 1 927 c/, 19726) gave external and cranial measurements of the male
holotype from Peru and mean and range of measurements of 17 specimens from Peru.

Vampyrops recifinus Thomas, 1901

Measurements of Vampyrops recifinus have been recorded as follows: Thomas (1901c),
external and cranial measurements of the male holotype from Brazil; Cunha Vieira
(1942), external measurements of a male and a female from Brazil; Sanborn (1955),
external and cranial measurements (range) of specimens from Brazil and Guyana.

Vampyrops vittatus( Peters, 1859)

Measurements of Vampyrops vittatus have been recorded as follows: Dobson (1878u),
external measurements of one specimen; Goodwin (1946), external and cranial measure¬
ments of a specimen from Costa Rica; Sanborn (1955), forearm and cranial measurements
(range) of specimens from Venezuela, Colombia, Brazil, Ecuador, and Peru (he considered
V. vittatus and V. fuscus nonspecific); Hall and Kelson (1959), external and cranial measure¬
ments of a single specimen from Colombia; Davis et al. (1964), external and cranial measure¬
ments of a male and two females from Costa Rica; Gardner et al. (1970), forearm and
cranial measurements (mean, range) of six males and nine females from Costa Rica; Gardner
and Carter (19726), external and cranial measurements of the male holotype from Venezuela
and several of these measurements (mean, range) for six specimens from Peru.

Geographic variation. — According to Gardner and Carter (19726) measurements
of six specimens from Peru were much the same as those reported by Gardner et al. (1970)
for 19 specimens from Costa Rica.

Subfamily Brachyphyllinae
Brachyphylla cavernarum Gray, 1834

Measurements of Brachyphylla cavernarum have been recorded as follows: Gray (1834),
external measurements of the holotype from St. Vincent; Dobson (1878u), external measure-

BIOLOGY OF THE PHYLLOSTOMATIDAE

ments of one specimen; Miller (1902//), cranial measurements of a male topotype from
St. Vincent; Miller (19026), external measurements of a female specimen; Elliot (1904),
external and cranial measurements of one specimen; Miller (1913 c/), external and cranial
measurements of the female holotype of B. c. minor from Barbados and cranial measure¬
ments for an additional male; Elliot (1917), external and cranial measurements of the holo¬
type of B. c. minor, Anthony (1918, 1925), external measurements (mean, range) of 11
specimens (2 males, 9 females) and cranial measurements of 10 specimens (3 males, 7 females)
from Puerto Rico; Hall and Kelson (1959), external and cranial measurements (range) of
10 specimens and external and cranial measurements of the holotype of B. 6. minor from
Barbados; Husson (1960), forearm and cranial measurements (range) of 18 specimens from
St. Martin and Saba; Choate and Birney (1968), cranial measurements of two samples of
sub-Recent material from Puerto Rico; Koopman (1968), cranial measurements of a male
and female from Barbados (as given by Miller, 1913 a) and the range of a series of males
from Anguilla and females from St. Martin; Buden (1977), forearm measurements (mean,
range) of three males and eight females, cranial measurements of four males and eight
females from Puerto Rico, forearm measurements (mean, range) of seven males and three
females, and cranial measurements of 1 1 males and four females from St. John.

Geographic variation. — Buden (1977) treated all members of the genus as a single species.
Within the species, he recognized several areas of morphological variation. Individuals
from Puerto Rico, Virgin Islands, and most of the Lesser Antilles were the largest. Specimens
from Barbados in the Lesser Antilles were small compared to populations on adjacent
islands. Specimens from Cuba, Hispaniola, and the Bahamas were also small, with Cuban
material being distinguished by deeper and more robust zygomatic arches. However, Silva-
Taboada (1976), after examining this group, concluded that it contained two species, each
with two subspecies.

Initially, populations from Barbados (minor) and the remainder of the Lesser Antilles
(caver narum) were considered two separate species. Koopman (1968), however, showed that
there was overlap in size among both males and females and concluded from this that the
two were subspecies of B. cavernarum.

Brachyphylla nana Miller, 1902

Measurements of Brachyphylla nana have been recorded as follows: Gundlach (1872,
1877), external measurements of a specimen from Cuba; Miller (1902c/), cranial measure¬
ments of the holotype from Cuba; Miller (19026), external measurements of one female from
Cuba; Elliot (1904), external and cranial measurements of a single specimen; Miller (1918),
cranial measurements of the holotype and an additional specimen of B. nana pumila from
the type locality on Haiti; Miller (1929), cranial measurements of one specimen from Haiti;
Goodwin (1933), external measurements of five males from the Dominican Republic
and one female from Cuba; Sanborn (1941), external measurements of three females (range)
and cranial measurements of one female from Haiti; Hall and Kelson (1959), cranial
measurements of the holotype of Brachyphylla nana and B. pumila:, Silva-Taboada (1974),
measurements of fossil humeri, crania, and mandibles from Cuba; Buden (1977), forearm
measurements (mean, range) of eight males and 13 females, cranial measurements (mean,
range) of five males and nine females from Cuba, forearm measurements of seven males
and three females, and cranial measurements of 10 males and three females from Hispaniola
and of seven males and 12 females from Middle Caicos, Bahamas.

Geographic variation. — Buden (1977), considering B. nana and B. cavernarum
conspecific, found populations from Middle Caicos, Cuba, and Hispaniola (nana) to be
distinctly smaller than individuals from Puerto Rico, Virgin Islands, and the remainder
of the Lesser Antilles (cavernarum). Many characters of specimens from Caicos and
Hispaniola overlap broadly, but Buden distinguished specimens from the two areas by the
deeper and more robust zygomatic arch of specimens from Cuba.

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Erophylla bonibifrons( Miller, 1899)

Measurements of Erophylla bombifrons have been recorded as follows: Miller (1899),
external and cranial measurements of the male holotype from Puerto Rico; Elliot (1904),
external and cranial measurements of the holotype from Puerto Rico as given by Miller
(1899); Elliot (19056), external and cranial measurements of the holotype of E. h. santacristo-
balensis from the Dominican Republic; Elliot (1917), external and cranial measure¬
ments of the holotype of E. b. santacristobalensis , Anthony (1918, 1925), external measure¬
ments (mean, range) of six specimens and cranial measurements (mean, range) of three
specimens from Puerto Rico; Miller (1929), cranial measurements of three specimens from
Haiti and three from Puerto Rico; Hall and Kelson (1959), forearm and cranial measure¬
ments of the holotype of E. b. bombifrons ; Buden (1976), external and cranial measure¬
ments (mean, sd, range) of 49 specimens (21 cranial) from Hispaniola and 47 (18 cranial)
from Puerto Rico.

Individual variation. — Coefficients of variation in external measurements of specimens
from Hispaniola and Puerto Rico varied from 1.98 to 4.94 and in cranial measurements
from 1.84 to 3.45 (Buden, 1976).

Geographic variation. — Buden (1976) treated the two recognized species ( bombifrons
and sezekorni) of the genus as conspecifics and relegated them to subspecific status. Differences
between many of the currently recognized taxa were considered slight. Skull shape was
considered the main diagnostic factor in distinguishing bombifrons and sezekorni.

Erophylla sezekorni (Gundlach, 1861)

Measurements of Erophylla sezekorni have been recorded as follows: Gundlach (1877),
external measurements of a specimen from Cuba; Dobson ( 1 878c/), external measurements
of a single specimen; Miller (1899), external and cranial measurements of the male holotype
of E. s. plantifrons from the Bahamas; Elliot (1904), external and cranial measurements of
two specimens; G. M. Allen (1917), external and cranial measurements of the male holotype
from Jamaica; Shamel (1931), external and cranial measurements of the male holotype of
E. s. mariguanensis from Mariguana Island, southern Bahamas, cranial measurements
(range) of eight additional specimens, and eight from the northern Bahamas; Buden (1976),
external and cranial measurements (mean, sd, range) of 50 specimens (19 cranial) from New
Providence, Bahamas, 35 (six cranial) from Mayaguana, Bahamas, 88 (44 cranial) from
Cuba, and 66 (29 cranial) from Jamaica.

Individual variation. — Coefficients of variation in external measurements of specimens
from the Bahamas, Cuba, and Jamaica varied from 2.06 to 4.40 and in cranial measurements
from 1.58 to 2.93 (Buden, 1976).

Geographic variation. — See geographic variation in E. bombifrons.

Phyllonycteris aphylla (M iller, 1 898)

Measurements of Phyllonycteris aphylla have been recorded as follows: Miller (1898),
external and cranial measurements of the male holotype from Jamaica; Elliot (1904),
external and cranial measurements of one specimen; G. M. Allen (1942), external and
cranial measurements for the species; Hall and Kelson (1959), external and cranial measure¬
ments of the holotype; Henson and Novick (1966), external measurements of a female from
Jamaica; Howe (1974), external measurements of three females from Jamaica.

Phyllonycteris mjyor Anthony, 1917

Measurements of Phyllonycteris major have been recorded as follows: Anthony (1917,
1918, 1925), cranial measurements of the holotype and eight additional specimens
(sub-Recent fossils) from Puerto Rico; G. M. Allen (1942), cranial measurements for the

BIOLOGY OF THE PHYLLOSTOMATIDAE

species; Goodwin (1953), cranial measurements of the holotype from Puerto Rico; Choate
and Birney (1968), measurements (mean, range) of partial crania and partial lower jaws
from Puerto Rico.

Phyllonycteris poeyi Gundlach, 1861

Measurements of Phyllonycteris poeyi have been recorded as follows: Gundlach (1872,
1877), external measurements of a specimen from Cuba; Dobson (1878c/), external measure¬
ments of one specimen from Cuba; Elliot (1904), external and cranial measurements of a
single specimen from Cuba; Miller (1904), external measurements of a single specimen from
Cuba; Miller (1904), external measurements of 12 males and 13 females from Cuba; Anthony
(1917, 1918, 1925), cranial measurements of two specimens from Cuba; Miller (1929),
cranial measurements of the holotype of P. p. ohtusci and an additional specimen from
Haiti; G. M. Allen (1942), cranial measurements for P. p. obtusa; Hall and Kelson (1959),
cranial measurements of the holotype of P. p. obtusa and two specimens of P. p. poeyi;
Silva-Taboada (1974), measurements of fossil humeri, crania, and mandibles from Cuba.

Subfamily Desmodontinae
Desmodus rotundus (E. Geoffroy St.-Hilaire, 1810)

Measurements of Desmodus rotundus have been recorded as follows: Dobson (1878c/),
external measurements of one specimen; Flower and Lydekker (1891), forearm length of
the species; Jentink (1893), external measurements probably of a female from Guyana; H.
Allen (1896), cranial measurements of a single specimen; Cabrera (1903), external measure¬
ments for the species in Chile; Elliot (1904), external and cranial measurements of one
specimen; J. A. Allen (1906), external measurements (mean, range) of five specimens
from Jalisco; Miller (1912), external and cranial measurements of a female from Taboga
Island, Panama; Lima (1926), external and cranial measurements of a specimen from
Brazil; Goodwin (1934), external measurements of one specimen from Guatemala; Martinez
and Villa-R. (1940), external and cranial measurements of males and females combined
from Guerrero; Cunha Vieira (1942), external measurements of four males and four females
and cranial measurements of three males and one female from Brazil; Goodwin (1942 a),
external and cranial measurements of two females from Honduras; Osgood (1943), fore¬
arm measurements of two specimens from Chile; Goodwin (1946), external and cranial
measurements of a male and female from Costa Rica; Hershkovitz (1949), external and
cranial measurements (range) of 14 females and a large male obtained in a sample from
Colombia; Dalquest (1953//), external measurements (mean) of 10 males and 10 females
and cranial measurements of one male and one female from San Luis Potosi; de la Torre
(1954), external and cranial measurements of a female from Tamaulipas; de la Torre
(1955), forearm measurements of one male and one female from Guerrero; Felten (1956c),
external measurements (mean, range) of 33 males and 23 females and cranial measurements
(mean, range) of 19 females and eight females from El Salvador; Felten (1956//), cranial
measurements of a single specimen from El Salvador; Jones (1958), cranial measurements
(mean, range) of three males and seven females (combined) from Tamaulipas; Koopman
(1958), cranial measurements of a sub-Recent fossil from Cuba and the range of these
measurements in seven specimens from Tamaulipas; Hall and Kelson (1959), external and
cranial measurements of a male and female from Costa Rica; Burt and Stirton (1961), fore¬
arm and cranial measurements (range) of 14 males and 23 females; Goodwin and Green-
hall (1961), forearm measurements (range) of 15 males and 16 females and cranial measure¬
ments of one male and one female from Trinidad; Husson (1962), external and cranial
measurements of a male and five females from Surinam; Tamsitt and Valdivieso (1962),
external measurements of a male from Colombia and a large male reported from
Colombia by Hershkovitz (1949); Tamsitt and Valdivieso (1963//), external measurements
of one male and one female from Colombia; Valdivieso (1964), external measurements of a

SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

specimen from Colombia; Aellen (1965), forearm measurements of two males, the range of
eight females, and cranial measurements of one male from Peru; Brosset (1965), external
measurements of two males and a female and cranial measurements of a male and female from
Ecuador; Tamsitt and Valdivieso ( 1 966c/), forearm and cranial measurements of one male
and the range of four females from Colombia; Villa-R. (1967), external measurements
(mean, sd, range) of 53 specimens and cranial measurements (mean, sd, range) of 42 specimens
from Mexico; Genoways and Jones (1968), forearm measurements (mean, range) of 10
young specimens (seven males, three females) from Zacatecas; Goodwin (1969), forearm
and cranial measurements of seven males and seven females from Oaxaca; Anderson (1972),
external measurements (mean, sd, range) of 21 specimens and cranial measurements
(mean, sd, range) of six specimens from Chihuahua; Smith and Genoways (1974), external
and cranial measurements of a male from Margarita Island, Venezuela, and mean and
range of four males from the adjacent mainland; Woloszyn and Mayo (1974), cranial measure¬
ments of the holotype of the sub-Recent D. r. puntajudensis from Cuba, one sub-Recent
specimen from Mexico, 10 Recent specimens (mean, range) from Mexico, and measure¬
ments after Koopman (1958) and Husson (1962).

Individual variation. — In specimens from Guerrero, coefficients of variation for external
measurements of sexes combined varied from 2.51 to 16.80 and for cranial measurements
from 1.48 to 4.41 (Martinez and Villa-R., 1940).

Secondary sexual variation. — Hershkovitz (1949) noted that males were smaller than
females, and Husson (1962) concluded from published accounts that males were smaller than
females.

Geographic variation.— Measurements of individuals from Surinam agreed well,
according to Husson (1962), with those from Colombia (Hershkovitz, 1949) and Trinidad
(Goodwin and Greenhall, 1961).

Diaemus youngii (Jentink, 1893)

Measurements of Diaemus youngii have been recorded as follows: Jentink (1893),
external measurements of the male holotype of D. y. youngii from Guyana; Thomas (19286),
external and cranial measurements of the female holotype of D. y. cypselinus from Peru;
Cunha Vieira (1942), external and cranial measurements of a male and female from Brazil;
Sanborn (1949), external and cranial measurements of one specimen from Venezuela and
another from Peru; Goodwin and Greenhall (1961), forearm measurements of one male
and two females and cranial measurements of one male and female from Trinidad; Husson
(1962), external and cranial measurements of the holotype from Guyana; Lay (1962),
external and cranial measurements of a male and female from Tabasco; Villa-R. (1965),
external and cranial measurements of a female from Tamaulipas; Villa-R. (1967), external
and cranial measurements of a specimen from Mexico; Gardner et al. (1970), external and
cranial measurements of a male from Costa Rica; Smith and Genoways (1974), external
and cranial measurements of one specimen from Margarita Island, Venezuela, three males
(mean, range) and one female from the adjacent mainland, and the holotype of D. youngii.

Geographic variation. — Gardner et al. (1970) reported that measurements of their
Costa Rican specimen were much larger than the holotype of D. y. youngii from Guyana but
that it agreed closely with the holotype of D. y. cypselinus from Peru and with a specimen
from Tamaulipas recorded by Villa-R. (1965). Measurements of two specimens from
Tabasco (Lay, 1962) were somewhat larger than those of a specimen from Costa Rica
(Gardner et al., 1970).

Diphylla ecaudata Spix, 1823

Measurements of Diphylla ecaudata are recorded as follows: Dobson (1878a), external
measurements of a specimen from Brazil; H. Allen (1896), external measurements of two

BIOLOGY OF THE PHYLLOSTOMATIDAE

specimens and cranial measurements of one from Mexico; Thomas (1903/)), external and
cranial measurements of the male holotype of D. e. centralis from Panama; Elliot (1904),
external and cranial measurements of the male holotype of D. e. centralis from Panama
(after Thomas, 1903 6) and another specimen; Lima (1926), external measurements of a
specimen from Brazil; Sanborn (1936), external and cranial measurements of one female
from Ecuador; Cunha Vieira (1942), external and cranial measurements of a male from
Brazil; Goodwin (1942a), external and cranial measurements of two males from Honduras;
Goodwin (1946), external and cranial measurements of two males from Honduras
(as given by Goodwin, 1942a) and the holotype of D. e. centralis from Panama; Dalquest
(1950), cranial measurements (mean) of seven males and three females from San Luis
Potosi; Dalquest (1953a), external measurements (mean) for two males and 13 females
and cranial measurements (mean) of seven males and three females from San Luis
Potosi; de la Torre (1954), external and cranial measurements of a male from Tamaulipas;
Felten (1956c), cranial measurements of five males from El Salvador; Felten ( 1 956c/),
external measurements of one specimen from El Salvador; Hall and Kelson (1959), external
and cranial measurements of the holotype of D. e. centralis , Burt and Stirton (1961),
forearm and cranial measurements (range) of six males and nine females from El Salvador;
Villa-R. (1967), external measurements of 20 specimens and cranial measurements of 19
from Mexico; Reddell (1968), external measurements of one female from Texas; Goodwin
(1969), forearm and cranial measurements of a male from San Luis Potosi and a female
from Yucatan; Ojasti and Linares (1971), forearm measurements (mean, se, range) of 16
males and 10 females and cranial measurements of 10 males and nine females from
Venezuela; Starrett (1976), forearm measurement of a single female from Costa Rica.

Geographic variation. — Ojasti and Linares (1971) compared length of forearm and
length of skull of specimens of Diphylla ecaudata from Central and South America. They
concluded that these populations were sufficiently distinct to warrant recognition as
separate subspecies.

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BIOLOGY OF THE PHYLLOSTOMATIDAE

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Sao Paulo I-Phyllostominae. Ciencia e Cultura, 27:621-632.

- . 19756. Phyllostomidae (Chiroptera) do Norte-ocidental do Estado de Sao

Paulo II-Glossophaginae; Carolliinae; Sturnirinae. Ciencia e Cultura, 27:723-734.

Tamsitt, J. R., and D. Valdivieso. 1962. Desmodus rotundus rotundus from a high
altitude in southern Colombia. J. Mamm., 43:106-107.

- . 1963a. Records and observations on Colombian bats. J. Mamm., 44: 168-180.

- . 19636. Notes on bats from Leticia, Amazonas, Colombia. J. Mamm., 44:263.

- . 1965a. The male reproductive cycle of the bat Artibeus lituratus. Amer.

Midland Nat., 73:150-160.

- . 19656. Reproduction of the female big fruit-eating bat, Artibeus lituratus

palmarum, in Colombia. Caribbean J. Sci., 5:157-166.

- . 1966a. Bats from Colombia in the Swedish Museum of Natural History, Stock¬
holm. Mammalia, 30:97-104.

- . 19666. Taxonomic comments on Anoura caudifer, Artibeus lituratus and

Molossus molossus. J. Mamm., 47:230-238.

- . 1966c. Parturition in the red fig-eating bat, Stenoderma rufum. J. Mamm.,

47:352-353.

SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

Thomas, M. E., and D. N. McMurray. 1974. Observations on Sturnira aratathomasi
from Colombia. J. Mamm., 55:834-836.

Thomas, O. 1889. Description of a new stenodermatous bat from Trinidad. Ann. Mag.
Nat. Hist., ser. 6, 7:167-170.

- . 1891a. Descriptions of three new bats in the British Museum Collection. Ann.

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species of the genus. Ann. Mus. Civ. Genova, 2a. ser., 10:881-883.

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Mag. Nat. Hist., ser. 6, 10:408-410.

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- . 19016. On a collection of mammals from Kanuku Mountains, British Guiana.

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descriptions of new species. Ann. Mag. Nat. Hist., ser. 7, 12:455-464.

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species. Ann. Mag. Nat. Hist., ser. 8, 4:230-242.

- . 1910. Mammals from the River Supinaam, Demerara, presented by Mr. F. V.

McConnell to the British Museum. Ann. Mag. Nat. Hist., ser. 8, 6:184-189.

- . 1912a. Three small mammals from S. America. Ann. Mag. Nat. Hist., ser. 8,

9:408-410.

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10:403-411.

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ser. 8, 14:410-414.

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by Mr. R. W. Hendee in the Chachapoyas Region of north Peru. Ann. Mag.
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BIOLOGY OF THE PHYLLOSTOMATIDAE

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Mamm., 52:247-250.

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Valdivieso, D., and J. R. Tamsitt. 1962. First records of the pale spear-nosed bat in
Colombia. J. Mamm., 43:422-423.

Villa-R., B. 1953. Mamiferos silvestres del Valle de Mexico. An. Inst. Biol., Mexico,
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(J. A. Allen) y Chiroderma isthmicum Miller en Mexico. An. Inst. Biol.,
Mexico, 33:379-384.

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siricoteros de Mexico, genero Glossophaga. An. Inst. Biol., Mexico, 34:381-391.

- . 1965. Diaemus youngi (Jentink) el vampiro, overo, en el sur de Tamaulipas,

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Biol., xvi + 491 pp.

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Walton, D. W. 1963. A collection of the bat Lonchophylla robusta Miller from Costa
Rica. Tulane Studies, Zool., 10:87-90.

Ward, H. L. 1891. Descriptions of three new species of Mexican bats. Amer. Nat.,
25:743-753.

Watkins, L. C., J. K. Jones, Jr., and H. H. Genoways. 1972. Bats of Jalisco, Mexico.
Spec. Publ. Mus., Texas Tech. Univ., 1:1-44.

Woloszyn, B. W., and N. A. Mayo. 1974. Postglacial remains of a vampire bat
(Chiroptera: Desmodus) from Cuba. Acta Zool Cracoviensia, 19:253-265.

SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

Appendix 1. — Selected measurements of phyllostomatid bats. Museum acronyms used are as
follows: AMNH, American Museum of Natural History ; AS, Albert Schwartz Collection;
BMNH, British Museum ( Natural History), CM, Carnegie Museum of Natural History,
COLU, Department of Biology, Colgate University; FHKSC, Museum of the High Plains,
Fort Hays Kansas State College; KU, Museum of Natural History, University of Kansas;
LACM, Natural History Museum of Los Angeles County; LSU, Museum of Zoology,
Louisiana State University; ROM, Royal Ontario Museum; TCWC, Texas Cooperative
Wildlife Collection, Texas A&M University; TTU, The Museum, Texas Tech University;

USNM, National Museum of Natural History.

Museum, catalogue

no., and sex Locality

_L_

'oo

C/5

—

C/5

tfl

—

C/5

■o

•u

C/5

-J

Phyllostomatinae

Chrotopterus auritus

KU 23661 2

Veracruz

78.9

36.7

31.2

19.2

5.9

13.4

12.9

12.0

KU 93385 2

Yucatan

78.7

36.2

31.1

18.5

6.3

13.3

12.7

11.5

USNM 305204 2

Panama

83.1

37.8

31.8

19.5

6.2

14.0

13.5

12.4

USNM 335156 2

Panama

82.5

37.1

31.7

19.6

6.2

14.5

13.2

12.6

TTU 9339 6

Veracruz

81.1

35.7

30.4

18.5

5.9

12.9

13.0

12.0

KU 23622 d

Veracruz

79.1

35.7

30.6

18.1

6.1

12.9

13.0

11.6

KU 93383 S

Yucatan

80.8

36.0

31.0

18.2

6.3

13.3

12.9

11.6

KU 105962 6

Nicaragua

79.8

36.2

31.0

18.2

6.0

13.2

12.6

11.3

Lonchorhina aurita

USNM 305186 2

Panama

50.5

20.0

17.9

10.4

4.8

8.4

6.3

7.0

TTU 5320 V

Trinidad

47.1

20.8

19.1

10.4

4.9

8.9

6.7

7.1

TTU 5322 2

Trinidad

50.3

20.5

18.6

10.8

4.9

8.8

6.6

7.0

TTU 8984 2

Trinidad

51.1

20.6

18.7

10.8

4.9

8.9

6.6

7.0

TTU 5321 6

Trinidad

49.0

20.7

18.9

10.5

4.9

8.7

6.6

7.0

TTU 5323 J

Trinidad

50.0

20.4

19.0

10.8

4.8

8.7

6.6

7.1

TTU 9827 6

Trinidad

49.9

20.7

18.7

10.4

5.0

8.7

6.6

7.1

TTU 9829 <3

Trinidad

49.8

20.5

18.7

10.4

4.9

8.7

6.6

7.0

Lonchorhina orinocensis

USNM 373254 2

Venezuela

42.3

19.0

16.4

9.3

4.0

8.0

5.9

6.0

USNM 373255 2

Venezuela

42.2

19.1

16.4

9.6

4.1

8.2

5.9

6.0

USNM 373256 2

Venezuela

43.7

19.2

16.8

9.6

4.2

8.0

6.0

USNM 373260 2

Venezuela

41.5

18.3

16.2

9.1

3.8

8.2

5.7

USNM 373248(3

Venezuela

41.4

19.5

17.0

9.7

4.2

8.3

6.0

5.9

USNM 373249(3

Venezuela

42.6

19.5

17.0

9.6

4.0

8.0

6.3

6.1

USNM 373257d

Venezuela

41.5

19.3

17.0

9.7

4.0

8.3

6.0

5.9

USNM 373258(3

Venezuela

43.0

19.5

16.8

9.8

4.0

8.1

6.1

Macrophylt um macrophyll ton

AMNH 177666 2

Nicaragua

36.9

17.1

14.5

9.2

3.4

8.2

5.5

6.1

AMNH 177669 2

Nicaragua

36.2

17.0

14.7

9.5

3.1

7.8

5.7

6.4

AMNH 177670 2

Nicaragua

37.4

17.4

14.7

9.5

3.0

8.0

5.5

6.2

AMNH 177671 2

Nicaragua

37.4

17.1

14.7

9.4

3.0

8.1

5.6

6.2

KU 70478 3

Nicaragua

35.6

16.6

13.6

9.2

3.0

7.8

5.2

6.1

USNM 311944(3

Panama

35.0

16.8

14.2

8.9

3.2

7.8

5.5

6.1

USNM 312963 6

Panama

37.2

17.7

14.9

9.8

3.2

8.0

5.7

6.7

USNM 315212(3

Panama

34.3

17.2

14.2

10.0

3.2

8.0

5.7

6.8

Macrolus californicus

FHKSC 2442 2

Arizona

49.0

22.7

19.9

10.4

3.3

8.1

8.9

7.4

TTU 10529 2

Sonora

49.4

22.3

19.9

10.8

3.5

8.1

9.0

7.0

TTU 10584 2

Sonora

51.8

22.9

20.7

11.2

3.5

9.0

8.8

7.1

BIOLOGY OF THE PHYLLOSTOMATIDAE

Appendix 1. — Continued.

TTU 10588 2

Sonora

51.7

23.0

20.4

11.2

3.7

8.4

8.9

7.0

FHKSC 1994 6

Arizona

50.7

23.2

20.5

11.4

3.8

8.5

8.7

7.5

TTU 10582 6

Sonora

49.7

23.8

20.1

11.7

3.5

8.5

9.5

7.4

TTU 10585 6

Sonora

48.3

22.6

20.2

10.6

3.6

8.1

8.9

7.0

TTU 10587 6

Sonora

50.0

23.2

20.1

11.1

3.6

8.3

9.2

7.3

Macrotus waterhousii

TTU 10566 2

Sonora

49.2

22.2

19.5

10.6

3.9

8.5

8.4

7.2

TTU 21470 2

Jamaica

53.9

25.3

21.5

12.2

4.1

9.2

9.4

7.8

TTU 21471 2

Jamaica

55.0

25.8

21.9

12.5

4.2

9.2

9.7

8.0

TTU 21505 2

Jamaica

54.1

26.0

22.0

12.0

4.2

8.8

9.6

7.5

TTU 6267 6

Sonora

47.2

23.2

20.0

11.1

4.2

8.6

8.9

7.5

TTU 10564 6

Sonora

49.6

23.0

20.0

11.2

4.1

8.6

8.8

7.4

TTU 10565 6

Sonora

48.3

22.4

19.4

10.9

4.1

8.5

8.6

7.4

TTU 21501 6

Jamaica

54.7

26.4

22.0

12.4

4.4

9.5

9.7

7.9

Micronycteris behni

BMNH 69.5.13.3 2

Peru

4.7

8.1

7.2

Micronycteris brachyolis

USNM 323059 2

Panama

42.2

22.4

19.3

11.4

5.0

9.0

8.3

7.3

TTU 5237 2

Trinidad

39.4

21.3

18.5

10.2

5.0

8.6

8.1

6.7

TTU 5315 2

Trinidad

40.9

21.6

19.0

10.4

5.1

8.4

8.6

6.9

AMNH 175633 2

Trinidad

40.3

21.4

18.7

10.5

5i0

8.5

8.1

6.9

USNM 245153 (3

Guatemala

40.9

21.7

19.3

10.7

5.1

8.7

8.2

6.9

USNM 306546<3

Panama

40.8

22.8

19.9

11.1

5.2

8.9

8.2

7.0

USNM 323060<3

Panama

40.2

21.3

18.7

10.8

4.8

8.4

7.9

6.9

TTU 5314<3

Trinidad

39.4

21.9

19.2

10.5

5.2

8.9

8.2

7.0

Micronycteris daviesi

BMNH 64.767 2

Guyana

57.1

27.3

23.7

13.3

6.5

10.8

11.0

9.3

USNM 335104(3

Panama

54.0

27.3

23.5

13.2

6.7

10.5

10.7

9.2

USNM 460089(3

Brazil

54.7

26.1

22.8

12.8

6.2

10.6

10.5

9.1

M icronycteris h irsuta

TTU 13158 2

Nicaragua

42.5

23.8

20.6

11.8

5.0

8.4

9.4

7.3

CM 2659 2

Colombia

42.9

23.0

19.9

11.3

4.7

8.9

8.8

6.9

USNM 418876 2

Venezuela

42.2

24.0

20.6

11.6

4.8

8.6

9.3

7.5

TTU 5299 2

Trinidad

43.0

23.8

20.6

11.8

5.2

8.8

9.4

7.5

TTU 13155(3

Nicaragua

39.5

22.8

19.4

11.0

4.7

8.5

8.7

7.2

TTU 5410(3

Trinidad

42.1

24.0

20.2

11.6

5.0

8.9

9.2

7.4

TTU 5449 (3

Trinidad

42.3

23.7

20.3

11.3

4.9

8.7

8.9

7.2

TTU 101 16 6

Trinidad

42.7

24.3

20.7

11.5

5.0

8.5

9.2

7.3

Micronycteris megalotis

KU 70474 2

Nicaragua

38.0

20.2

17.4

9.7

4.2

8.0

7.5

6.5

KU 97407 2

Nicaragua

33.6

19.6

17.2

9.2

4.1

7.7

7.1

6.2

KU 97409 2

Nicaragua

36.7

19.6

17.1

9.0

3.9

7.7

7.3

6.3

KU 114772 2

Nicaragua

34.7

18.6

16.3

9.1

4.0

7.4

7.0

6.0

TTU 5438 (3

Trinidad

32.6

18.4

16.0

8.9

4.0

7.5

6.9

6.0

TTU 5446 (3

Trinidad

35.5

19.1

16.3

8.7

3.9

7.5

7.0

5.9

TTU 5495 (3

Trinidad

32.2

18.7

16.2

8.6

3.8

7.4

6.9

6.0

TTU 9788 (3

Trinidad

33.3

18.8

16.1

8.8

4.1

7.5

7.1

5.9

Micronycteris minuta

TTU 5226 2

Trinidad

36.5

18.5

15.9

8.6

4.3

7.6

6.6

5.7

TTU 5437 2

Trinidad

35.6

18.6

15.8

8.6

4.0

7.5

6.5

5.7

TTU 5443 2

Trinidad

34.6

18.4

16.0

8.4

4.0

7.6

6.5

5.5

TTU 5444 2

Trinidad

36.5

18.8

16.4

8.5

4.0

7.5

6.7

5.6

TTU 5225 (3

Trinidad

35.3

18.8

16.5

8.6

4.2

7.4

6.8

5.6

TTU 5239 (3

Trinidad

35.2

18.7

16.5

8.7

4.1

7.6

6.4

5.5

TTU 5294 J

Trinidad

34.9

18.3

16.0

8.7

4.2

7.5

6.7

5.5

TTU 5295 (3

Trinidad

35.5

19.0

16.2

8.3

4.0

7.4

6.7

5.5

SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

Appendix 1. — Continued.

Micronycteris nicefori

TTU 5257 9

Trinidad

40.2

22.0

19.6

9.8

4.5

8.3

7.8

6.3

TTU 5297 9

Trinidad

40.0

21.4

19.3

9.5

4.1

8.2

7.7

6.0

TTU 5298 9

Trinidad

38.8

21.2

19.9

9.4

4.2

7.6

6.3

TTU 8954 9

Trinidad

38.4

21.1

19.0

9.8

4.4

8.3

7.5

6.3

TTU 8963 c?

Trinidad

36.8

21.1

18.9

9.5

4.0

8.0

7.6

6.2

TTU 8964 6

Trinidad

37.1

20.7

18.2

9.5

4.3

8.3

7.2

6.3

TTU 8965 c?

Trinidad

36.3

20.8

18.7

9.6

4.2

8.2

7.6

6.1

TTU 8966 c?

Trinidad

39.1

20.8

18.4

9.3

4.0

8.5

7.5

6.3

Micronycteris pnsilla

AMNH 78830 c?

Brazil

34.3

15.2

8.9

4.2

7.6

6.7

5.7

AMNH 78831 c?

Brazil

33.7

17.8

15.4

4.5

7.8

6.8

6.0

Micronycteris schmidtorum

USNM 388704 9

Venezuela

34.7

19.5

16.9

9.3

4.2

7.5

6.1

USNM 388713 9

Venezuela

34.2

19.8

17.1

9.2

4.4

7.7

7.4

6.0

USNM 407257 9

Venezuela

33.9

19.4

17.1

9.4

4.2

7.9

7.2

6.1

USNM 415210 9

Venezuela

35.1

18.9

17.0

9.1

4.1

7.4

5.9

AMNH 130715 c?

Venezuela

36.9

20.0

17.7

9.7

4.2

7.8

6.5

AMNH 130718 c?

Venezuela

36.0

20.1

17.4

9.5

4.0

7.6

7.5

6.4

AMNH 130725 c?

Venezuela

36.8

20.2

17.8

9.8

4.1

7.6

7.8

6.7

USNM 444235 <?

Venezuela

37.3

20.2

17.7

9.6

4.2

7.6

7.9

6.6

Micronycteris

sylvestris

KU 96970 9

Nayarit

43.8

21.2

18.7

10.2

4.9

8.5

8.4

6.7

KU 23646 9

Veracruz

42.1

20.5

18.7

10.4

4.8

8.5

8.4

7.0

USNM 396399 9

Panama

42.0

19.8

10.7

4.5

8.7

7.9

7.2

KU 23651 3

Veracruz

41.2

21.0

18.7

10.2

4.9

8.6

8.3

7.0

KU 23653 £

Veracruz

40.3

20.7

18.5

10.1

4.9

8.4

8.1

6.9

KU 29594 £

Veracruz

38.3

21.0

18.7

10.2

4.9

8.8

8.5

6.9

BMNH 96.10.1.2 £

Costa Rica

39.5

20.1

17.1

9.9

4.5

8.2

7.9

6.8

Minton bennettii

BMNH 3.7.1.153 9

Brazil

56.6

26.1

22.8

13.8

4.6

9.8

9.4

9.1

BMNH 3.7.1.155 9

Brazil

56.0

25.4

21.8

4.6

9.8

9.0

8.6

USNM 391027 9

Brazil

58.4

26.3

22.9

14.1

4.7

10.0

9.4

9.5

BMNH 65.618 c?

Guyana

53.1

25.6

22.0

13.9

4.5

9.6

9.0

9.5

USNM 123393 c?

Brazil

53.7

24.5

21.8

4.7

9.1

9.3

Mimon coz

umelae

KU 23658 9

Veracruz

57.5

26.7

23.3

14.3

4.8

10.1

9.5

9.7

KU 32092 9

Veracruz

55.1

26.1

22.6

14.2

4.4

10.0

9.5

9.8

KU 91548 9

Yucatan

59.0

27.0

23.0

13.8

4.5

9.6

9.5

9.3

KU 93380 9

Yucatan

54.9

25.2

22.1

13.6

4.4

9.8

9.1

8.8

KU 19171 c?

Veracruz

55.1

26.5

22.6

13.3

4.5

9.9

9.4

KU 23656 c?

Veracruz

55.4

26.0

22.2

13.8

4.6

9.9

9.4

9.7

KU 91546 c?

Yucatan

56.4

25.6

22.5

13.1

4.4

9.5

9.3

8.8

TTU 9340 c?

Yucatan

56.6

26.0

22.7

13.8

4.7

9.8

9.1

Mimon crenulutum

USNM 371497 9

Venezuela

48.8

21.3

18.5

11.7

3.9

8.7

7.6

8.4

USNM 371503 9

Venezuela

50.4

21.9

18.9

12.1

3.8

8.8

7.7

7.9

TTU 5340 9

Trinidad

45.9

22.0

19.2

12.5

4.0

8.4

7.9

9.0

TTU 5374 9

Trinidad

48.7

22.6

19.5

12.0

4.3

8.6

7.7

8.4

TTU 5264 <3

Trinidad

47.0

21.6

18.7

11.6

3.9

7.9

7.5

8.5

TTU 5375 <?

Trinidad

51.0

22.4

19.6

11.8

4.2

8.7

7.6

8.6

TTU 5379 c?

Trinidad

47.3

21.6

18.5

11.7

3.9

8.4

7.7

8.4

TTU 5460 c?

Trinidad

48.2

21.9

19.0

11.7

4.1

8.3

7.7

8.5

Mimon koepckeae

LSU 1 6447 9

Peru*

47.4

21.9

18.9

11.6

4.2

8.6

7.5

8.2

LSU 15675 c?

Peru

46.3

21.7

18.5

11.4

4.1

8.3

7.1

8.0

LSU 15676 c?

Peru

49.8

21.5

18.9

11.7

3.9

8.2

7.4

8.1

BIOLOGY OF THE PHYLLOSTOMATIDAE

Appendix 1. — Continued.

USNM 388843 9

Venezuela

Phylloderma stenops

71.8 30.9 26.0

14.9

8.9

12.7

9.7

USNM 388844 9

Venezuela

71.6

32.2

28.3

16.0

9.2

13.4

10.0

10.2

USNM 388848 9

Venezuela

71.2

30.3

26.3

14.8

9.0

12.9

9.7

9.4

TTU 5318 9

Trinidad

65.8

30.0

25.3

14.6

8.9

12.1

9.6

9.8

USNM 335144 <J

Panama

73.4

31.8

27.3

15.6

9.1

13.0

10.2

9.6

USNM 388842 6

Venezuela

72.2

31.2

27.4

14.7

8.9

12.7

9.8

9.6

USNM 388845 6

Venezuela

69.5

30.3

26.5

14.7

8.7

12.7

9.6

9.4

USNM 388846 i

Venezuela

73.3

30.8

26.5

16.3

9.0

13.1

10.3

10.6

KU 114811 9

Nicaragua

Phyllostomus discolor

61.7 30.8 27.1

15.0

6.5

12.1

9.5

10.3

KU 114812 9

Nicaragua

64.9

31.8

26.8

16.1

6.6

12.2

9.9

10.5

KU 1 148 13 9

Nicaragua

61.6

29.5

26.2

15.3

6.2

11.7

9.5

10.0

TTU 5452 9

Trinidad

59.9

29.7

25.6

15.4

6.2

12.1

9.0

9.6

KU 110701 6

Nicaragua

62.9

31.0

27.3

15.5

6.6

11.8

9.8

10.0

KU 1 10702 c?

N icaragua

64.3

32.1

28.5

16.4

6.4

12.6

10.0

10.4

KU 1 14800 6

Nicaragua

61.8

31.7

28.5

16.1

6.6

12.3

9.8

10.4

TTU 5412 6

Trinidad

60.9

30.5

27.0

15.4

6.3

11.9

9.6

9.8

Phyllostomus elongalus

USNM 364304 9

Peru

67.5

30.6

26.3

16.1

5.4

10.9

10.6

11.2

USNM 364306 9

Peru

66.2

30.0

25.2

16.7

5.4

11.1

10.3

11.4

USNM 364310 9

Peru

64.3

29.0

25.2

16.6

5.3

10.9

10.2

11.3

USNM 499015 9

Peru

64.6

29.0

25.4

16.3

5.4

10.8

10.0

11.3

USNM 483339 6

Colombia

60.8

28.9

24.7

15.4

5.1

10.5

10.3

11.0

USNM 361515 6

Brazil

64.5

30.5

25.6

16.5

5.3

10.9

10.3

11.5

USNM 364303 <J

Peru

67.2

30.2

25.8

16.9

5.7

11.2

10.4

1 1.3

USNM 364305 6

Peru

67.7

29.1

25.5

16.5

5.6

10.8

10.1

11.0

Phyllostomus hastatus

KU 1 10716 9

Nicaragua

90.4

39.3

33.9

21.0

7.3

14.9

13.2

13.5

KU 110717 9

Nicaragua

92.6

40.7

34.3

21.3

7.1

14.6

14.1

14.2

KU 110720 9

Nicaragua

88.2

40.8

34.5

21.9

7.3

15.1

13.9

14.4

CM 2667 9

Colombia

84.7

38.9

32.5

20.7

7.2

14.2

13.0

14.0

KU 110718d

Nicaragua

91.1

41.5

35.8

22.4

7.7

15.2

14.3

KU 110719d

Nicaragua

94.4

43.1

36.1

23.2

7.7

15.5

13.8

14.3

ROM 31469(3

Trinidad

82.8

37.6

32.1

19.9

6.8

13.8

12.8

ROM 50233 d

Brazil

86.9

39.0

32.5

21.0

7.4

14.1

12.9

13.9

Phyllostomus latifolius

BMNH

1.6.4.44 9

Guyana

59.6

28.0

23.5

15.1

4.9

10.3

10.0

10.3

BMNH

1.6.4.45 9

Guyana

59.8

28.4

24.0

15.1

5.2

10.4

9.9

10.7

BMNH

1.6.4.40(3

Guyana

58.7

28.3

24.1

15.5

5.1

10.3

10.0

10.5

BMNH

1.6.4.41 (3

Guyana

59.2

28.3

24.4

4.8

10.3

10.4

10.6

BMNH

1.6.4.42(3

Guyana

58.9

28.6

24.5

15.8

5.1

10.4

10.3

11.2

BMNH

1.6.4.43(3

Guyana

58.5

28.2

24.1

15.8

5.0

10.5

10.0

10.9

Tonatia hidens

USNM 315218 9

Panama

58.8

28.9

24.0

14.0

5.7

10.7

9.7

8.9

TTU 5260 9

Trinidad

55.8

27.9

23.7

13.7

5.3

10.2

9.6

8.3

TTU 9774 9

Trinidad

54.8

28.3

24.1

14.3

5.4

10.6

9.3

8.4

TTU 9778 9

Trinidad

55.2

28.3

23.5

14.0

5.2

10.4

9.6

8.6

TTU 13108(3

Nicaragua

57.0

28.9

24.5

14.5

5.7

10.6

10.4

9.4

TTU 5261 (3

Trinidad

55.1

28.6

24.2

14.5

5.6

10.6

9.6

8.5

TTU 5338 (3

Trinidad

54.9

27.6

23.1

13.9

5.1

10.7

9.4

8.4

TTU 5339(3

Trinidad

51.5

27.3

23.1

13.8

5.6

10.6

9.5

8.6

Tonatia hrasiliense

AMNH 95497 9

Peru

35.7

20.0

16.6

9.4

3.3

8.1

6.8

6.5

AMNH 95498 6

Brazil

35.8

20.0

17.0

9.5

3.2

7.9

6.8

6.3

LSU 16440(3

Peru

37.9

20.8

17.0

9.4

3.2

8.3

6.8

6.1

SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

Appendix 1. — Continued.

Tonatia carrikeri

AMNH 30180 9

Venezuela

46.0

25.2

20.3

11.5

3.6

9.4

8.0

7.8

AMNH 30183 9

Venezuela

46.8

24.8

20.0

10.8

3.6

9.3

8.3

7.5

AMNH 209322 9

Bolivia

45.6

24.5

20.2

11.1

3.6

9.4

8.1

7.7

AMNH 30181 6

Venezuela

48.4

25.8

21.5

12.2

3.9

9.7

8.6

8.0

ROM 67468 6

Guyana

43.9

23.9

19.5

11.5

3.7

9.4

7.8

7.3

Tonatia minutu

USNM 314221 9

Panama

33.3

18.9

15.8

8.8

2.9

7.6

6.7

5.8

USNM 362457 9

Panama

34.0

19.2

16.3

9.0

2.9

7.8

6.8

6.2

USNM 362458 9

Panama

35.0

19.2

16.0

9.2

3.0

7.6

6.7

6.1

TTU 5238 9

Trinidad

35.8

20.1

16.9

9.6

2.9

8.0

6.9

6.2

TTU 5222 c?

Trinidad

36.3

20.2

16.8

9.6

3.2

8.4

7.0

6.1

TTU 5309 <?

Trinidad

34.5

20.2

16.8

9.6

3.1

8.2

6.7

6.4

TTU 5422 c?

Trinidad

35.5

20.6

17.6

10.0

3.4

8.5

7.0

6.7

TTU 10119c?

Trinidad

35.2

20.8

17.3

10.0

3.3

8.4

6.9

6.4

Tonatia silvicola

USNM 306549 9

Panama

51.6

27.0

22.8

12.9

3.9

10.5

8.9

8.1

USNM 309357 9

Panama

50.0

26.4

22.2

12.7

3.9

10.1

9.5

8.7

USNM 323068 9

Panama

53.3

26.7

22.6

12.9

3.9

10.2

9.0

8.3

USNM 364278 9

Peru

55.0

28.7

23.6

13.1

4.0

10.4

9.8

9.0

USNM 323074 c?

Panama

54.7

27.9

23.1

13.5

4.1

10.6

9.3

8.8

USNM 323076c?

Panama

53.5

27.8

23.3

13.3

4.1

10.5

9.2

8.7

USNM 407291 6

Venezuela

51.4

28.3

23.6

13.7

4.3

11.1

9.7

8.6

USNM 364275 c ?

Peru

55.2

30.4

24.8

14.1

4.1

10.8

10.4

9.6

Tonatia venezuelae

USNM 102919 9

Venezuela

39.8

21.5

17.9

10.5

3.1

8.3

7.5

6.9

USNM 142567 9

Venezuela

38.9

21.7

17.9

10.0

3.2

8.3

7.4

6.7

BMNH 11.5.25.41 c?

Venezuela

39.1

21.5

17.7

10.6

3.4

8.6

7.4

7.0

Trachops cirrhosus

KU 93381 9

Campeche

57.9

27.8

24.1

13.6

5.0

11.1

9.7

TTU 13172 9

Costa Rica

60.1

28.0

24.2

13.8

5.3

11.5

10.3

9.7

TTU 9777 9

Trinidad

60.1

29.0

25.4

14.8

5.3

11.9

10.8

10.4

TTU 9780 9

Trinidad

61.2

29.6

25.7

14.9

5.2

11.7

11.4

10.5

TTU 6077 c?

Oaxaca

59.3

28.2

24.5

13.8

5.0

11.5

10.0

9.7

TTU 6115 c?

Chiapas

59.5

28.2

24.5

13.5

4.9

11.1

10.1

9.8

KU 114818c?

Nicaragua

57.3

27.6

24.2

13.9

4.8

11.4

9.8

9.4

TTU 9779 c?

Trinidad

60.7

30.4

26.4

15.4

5.5

12.1

11.1

10.8

Vampyrum spectrum

USNM 335161 9

Panama

106.0

51.9

43.1

24.5

8.5

15.9

20.2

14.8

USNM 335162 9

Panama

107.1

53.6

43.7

25.4

7.9

15.6

19.8

15.0

TTU 5357 9

Trinidad

102.0

51.2

42.9

23.3

8.0

15.8

20.9

14.5

TTU 9837 9

Trinidad

103.3

51.6

44.0

23.4

7.7

15.7

21.1

15.4

AMNH 28993 c?

Nicaragua

105.7

50.4

42.3

24.5

8.0

15.6

19.7

14.4

KU 88190 c?

Costa Rica

110.4

50.7

43.0

23.4

8.1

15.8

19.9

14.3

TTU 9836 c?

Trinidad

106.1

52.4

44.1

24.2

7.8

15.8

21.1

15.2

TTU 11439 c?

Trinidad

107.1

52.0

43.2

25.2

8.4

16.4

20.6

15.4

Glossophaginae

Anoura brevirostrum

AMNH 214324 9

Peru

39.8

23.5

22.5

9.6

5.0

9.4

8.3

5.7

AMNH 233263 9

Peru

38.9

23.3

22.3

9.4

4.9

9.1

8.0

5.4

TCWC 11881 9

Peru

38.0

23.1

22.3

9.4

4.8

9.2

7.7

5.3

TCWC 11882 9

Peru

40.0

23.1

22.0

9.5

4.6

9.2

8.2

5.6

LSU 17941 c?

Peru

40.2

23.3

22.6

10.3

5.0

9.3

8.0

5.7

TCWC 11880 c?

Peru

39.6

23.3

22.5

10.0

4.8

9.3

8.1

5.4

Anoura caudifer

USNM 373705 9

Venezuela

38.5

24.6

23.8

9.5

4.6

9.3

9.1

5.5

USNM 373761 9

Venezuela

36.0

22.0

21.3

8.9

4.3

8.6

8.0

5.3

BIOLOGY OF THE PHYLLOSTOMATIDAE

Appendix 1. — Continued.

USNM 389076 9

Venezuela

35.1

22.1

21.4

9.1

4.6

8.9

8.2

5.6

USNM 389108 9

Venezuela

36.4

23.0

22.3

9.2

4.6

8.9

8.3

5.3

USNM 370109 6

Venezuela

37.2

24.0

23.2

9.4

4.6

8.8

5.3

USNM 373704 3

Venezuela

37.4

24.3

23.4

10.1

4.6

9.2

8.9

5.8

USNM 385771 <?

Venezuela

36.8

22.0

21.5

9.3

4.5

9.0

8.0

5.3

USNM 385773 c?

Venezuela

37.2

22.0

21.3

9.3

4.4

9.0

8.0

5.4

Anoura cull rata

USNM 309400 9

Panama

43.8

26.4

25.5

10.9

5.0

10.0

9.3

6.0

USNM 319249 9

Panama

41.7

26.3

25.5

10.3

5.1

10.0

9.2

6.0

USNM 419465 9

Venezuela

41.4

25.4

24.6

10.6

5.0

9.8

8.9

6.2

USNM 419466 9

Venezuela

41.1

25.6

24.5

10.4

5.0

10.0

8.7

6.2

USNM 309396 c?

Panama

43.0

26.4

25.6

10.7

5.3

10.3

9.1

5.7

USNM 309397 c?

Panama

44.3

26.6

25.8

11.0

5.3

10.0

9.4

6.1

USNM 309398 c?

Panama

43.6

26.7

26.0

11.1

5.3

10.0

9.3

6.1

USNM 337991 c?

Panama

42.3

26.2

25.4

11.0

5.3

10.0

9.3

6.3

Anoura geoffroyi

USNM 362594 9

Panama

43.7

26.3

25.7

11.0

4.9

9.8

10.1

6.3

USNM 385802 9

Venezuela

42.7

25.0

24.1

10.7

4.9

9.7

9.5

6.2

TTU 5825 9

Trinidad

42.7

25.0

24.2

10.8

4.8

9.7

9.5

6.3

TTU 8977 9

Trinidad

42.0

24.8

24.1

10.6

5.1

9.8

9.3

6.2

USNM 385852 c?

Venezuela

42.0

25.3

25.1

10.8

4.8

9.7

9.5

6.0

TTU 5370 c?

Trinidad

41.0

24.9

24.1

10.8

5.1

9.6

9.2

6.1

TTU 5823 c?

Trinidad

43.0

24.7

24.2

11.3

5.1

9.8

9.2

6.3

TTU 5826 c?

Trinidad

40.5

24.5

24.0

11.0

4.9

9.8

9.0

6.3

Anoura werckleae

LACM 25438 9

Costa Rica

43.1

26.1

25.3

10.5

5.2

10.1

9.3

6.0

LACM 15186 c?

Costa Rica**

40.7

25.8

25.1

10.8

5.3

10.2

9.0

6.1

Choeroniscus godmani

KU 90650 9

Sinaloa

33.8

20.7

20.0

3.2

8.2

7.3

4.2

AMNH 186162 9

Oaxaca

35.1

20.8

20.1

3.2

8.4

7.3

4.2

USNM 337550 9

Nicaragua

34.4

21.2

20.6

3.3

8.1

7.7

4.3

USNM 337551 9

Nicaragua

33.8

20.6

20.4

3.5

8.0

7.6

4.2

KU 102370 c?

Chiapas

33.4

19.7

18.8

2.9

7.9

6.9

4.2

AMNH 172778 c?

Oaxaca

32.6

19.3

18.8

3.0

8.3

6.7

4.0

AMNH 172779 c?

Oaxaca

33.1

18.9

18.2

2.9

8.0

6.7

3.9

AMNH 208869 c?

Oaxaca

32.3

19.2

18.5

2.9

8.3

6.4

4.0

Choeroniscus inca

AMNH 140471 9

Guyana

37.3

24.5

24.1

3.8

8.5

8.3

4.7

BMNH 12.9.5.2 9

Peru

33.1

3.8

8.5

7.6

4.5

Choeroniscus intermedins

TTU 5319 9

Trinidad

34.2

23.1

22.8

3.8

8.5

7.8

4.6

TTU 5496 9

Trinidad

34.9

23.2

22.8

3.5

8.3

8.0

4.5

TTU 9006 9

Trinidad

34.8

22.6

22.5

3.5

8.4

8.1

4.4

TTU 9007 9

Trinidad

36.0

23.6

23.0

4.0

8.7

8.1

4.6

TTU 8994 c?

Trinidad

34.1

22.8

21.8

3.3

8.8

7.6

4.3

TTU 8995 c ?

Trinidad

35.0

21.7

21.3

3.2

8.4

7.1

4.4

TTU 8998 c?

Trinidad

35.4

21.2

20.7

3.2

8.2

7.5

4.2

TTU 8999 c?

Trinidad

35.7

22.4

21.9

3.6

8.2

7.8

4.7

Choeroniscus

minor

AMNH 69152 9

Guyana

36.0

22.7

21.6

3.7

8.2

7.7

4.2

USMN 361573 9

Brazil

33.7

23.2

22.5

3.6

8.2

8.6

4.6

USNM 361574 9

Brazil

35.7

22.7

22.2

3.4

8.3

4.4

USMN 460100 9

Brazil

35.7

23.6

22.8

3.6

8.7

8.2

4.5

Choeroniscus periosus

AMNH 217038 9

Colombia

40.4

30.0

29.2

4.8

9.3

10.5

5.0

USNM 344918 9

Colombia

41.2

30.2

29.5

4.9

9.8

10.9

5.3

SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

Appendix 1. — Continued.

Choeronycteris mexicana

TTU 6288 9

Sonora

45.8

30.8

29.8

4.0

9.9

11.4

5.6

TTU 6360 9

Sonora

46.3

29.8

28.6

3.9

10.0

11.0

5.9

TTU 6447 9

Sonora

42.4

29.7

28.8

4.2

10.0

11.5

5.6

TTU 10122 9

Tamaulipas

45.9

29.4

28.5

4.0

9.6

11.0

5.7

KU 31863 S

Jalisco

45.6

30.3

29.0

4.1

9.6

11.5

5.6

KU 38250 <3

Jalisco

45.3

29.5

28.6

4.0

9.4

11.3

5.3

KU 107192d

Jalisco

43.0

29.4

28.5

3.7

9.4

11.0

5.2

KU 107194c3

Jalisco

43.6

30.1

28.9

3.8

9.7

11.6

5.5

Glossophaga alticola

KU 70624 9

Nicaragua

38.3

21.0

19.4

9.6

4.5

8.9

7.0

5.6

KU 70628 9

Nicaragua

38.1

20.4

18.8

9.7

4.5

8.9

7.0

5.7

KU 105966 9

Nicaragua

37.3

21.2

19.6

9.8

4.6

8.8

7.1

5.4

KU 114819 9

Nicaragua

36.8

20.9

19.3

9.5

4.4

8.6

7.1

5.6

KU 105964(3

Nicaragua

34.0

20.0

18.7

9.4

4.3

8.5

6.8

5.4

KU 105967(5

Nicaragua

35.8

20.1

18.4

9.4

4.6

8.9

6.9

5.3

KU 114820(3

Nicaragua

36.6

19.8

18.3

9.3

4.4

8.7

6.7

5.2

KU 114822(3

Nicaragua

36.3

20.7

19.0

9.8

4.5

9.0

7.1

5.8

Glossophaga commissarisi

KU 105972 9

Nicaragua

32.4

19.8

18.4

9.3

4.4

8.3

6.8

5.3

KU 105975 9

Nicaragua

32.7

20.2

18.8

9.6

4.5

8.2

6.9

5.5

KU 110770 9

Nicaragua

33.3

20.3

18.8

9.6

4.5

8.4

6.9

5.4

KU 110775 9

Nicaragua

34.5

20.4

19.0

9.3

4.3

8.1

7.1

5.4

KU 110730(3

Nicaragua

33.9

20.8

19.3

9.8

4.7

8.4

7.0

5.6

KU 110733 (3

Nicaragua

31.1

20.6

18.8

9.9

4.7

8.9

6.9

5.3

KU 110734(3

Nicaragua

34.6

20.3

18.8

9.9

4.5

8.4

6.7

5.5

KU 110767(3

Nicaragua

35.6

20.7

19.1

9.4

4.4

8.5

6.7

5.1

Glossophaga longirostris

TTU 9338 9

Grenada

38.6

23.1

21.5

9.4

4.7

8.6

7.9

5.8

KU 118105 9

Venezuela

38.0

22.8

21.4

10.1

4.4

8.8

8.0

5.5

KU 118117 9

Venezuela

38.6

23.3

21.6

10.1

4.6

8.8

8.1

5.9

KU 118123 9

Venezuela

39.5

23.3

22.0

9.9

4.5

8.8

8.2

6.0

KU 110073(3

Grenada

37.5

23.1

21.5

10.2

4.5

8.6

7.9

5.7

KU 118114(3

Venezuela

36.4

22.2

21.1

9.8

4.5

8.8

7.6

5.9

KU 118115 <3

Venezuela

37.6

23.0

21.2

9.8

4.4

8.8

7.7

5.8

KU 118116(3

Venezuela

36.3

22.8

21.4

10.1

4.7

9.4

8.0

5.8

Glossophaga soricina

KU 106015 9

Nicaragua

36.5

21.0

19.7

9.1

4.6

8.2

7.3

5.4

KU 106018 9

Nicaragua

36.7

21.4

19.9

9.3

4.5

8.6

6.9

5.2

KU 106019 9

Nicaragua

36.5

21.5

19.9

9.6

4.6

8.5

7.2

5.3

KU 106020 9

Nicaragua

36.0

21.9

20.6

10.0

4.9

8.9

7.7

5.7

KU 106008(3

Nicaragua

36.7

21.4

19.7

9.7

4.6

8.8

7.0

5.5

KU 106016(3

Nicaragua

35.0

20.9

19.2

9.2

4.7

8.5

7.0

5.3

KU 106021 (3

Nicaragua

34.5

21.1

19.3

9.4

4.5

8.4

7.0

5.3

KU 106022 (3

Nicaragua

36.8

21.7

20.1

9.7

4.5

8.6

7.3

5.4

Hylonycleris underwood i

KU 108603 9

Jalisco

36.3

20.6

20.0

4.0

8.1

7.2

4.2

KU 108605 9

Jalisco

33.0

20.6

20.0

3.9

8.1

7.0

4.2

KU 98140 9

Oaxaca

33.9

23.0

22.0

4.5

8.8

8.2

4.9

TTU 13142 9

Costa Rica

32.2

21.9

21.2

4.0

8.1

7.6

4.5

KU 108604(3

Jalisco

31.6

20.0

19.0

4.1

8.1

6.7

4.0

KU 108606(3

Jalisco

32.5

20.3

19.5

3.8

8.0

7.0

4.2

KU 23709 <3

Veracruz

33.1

21.6

20.8

4.1

8.6

7.4

4.6

KU 98139(3

Oaxaca

33.5

21.5

20.8

4.2

8.2

7.5

4.6

Leplonycteris curasoae

USNM 444799 9

Venezuela

53.7

28.1

26.8

11.2

5.2

9.8

9.6

7.0

USNM 444800 9

Venezuela

53.2

27.9

26.5

10.9

5.0

10.2

9.4

7.2

USNM 444802 9

Venezuela

54.4

27.5

26.7

11.0

4.7

9.6

9.3

7.3

BIOLOGY OF THE PHYLLOSTOMATIDAE

Appendix 1. — Continued.

USNM 444803 9

Venezuela

54.0

27.8

26.9

10.8

4.8

9.9

9.5

6.9

USNM 444734 6

Venezuela

50.6

21 A

26.1

11.3

4.9

10.0

9.4

7.2

USNM 444736 <J

Venezuela

52.6

27.1

26.1

11.2

4.7

10.1

9.1

7.0

USNM 444739(3

Venezuela

53.3

27.4

26.5

11.1

5.1

9.9

9.1

7.0

USNM 444740cJ

Venezuela

53.8

27.9

26.6

11.2

5.2

10.3

9.5

7.1

Leptonycteris nivalis

TTU 6565 9

Texas

58.2

27.8

27.1

11.5

5.3

11.0

9.6

7.1

KU 33068 9

Coahuila

52.0

28.9

27.5

11.2

5.2

11.0

9.6

7.0

KU 33070 9

Coahuila

50.6

27.5

26.8

11.3

5.5

10.7

9.2

7.1

KU 33071 9

Coahuila

52.9

29.1

27.5

11.4

5.6

11.0

9.4

6.7

TTU 9208 (3

Texas

56.7

27.7

26.8

11.0

5.5

10.6

9.0

6.7

KU 98378(3

Nuevo Leon

56.3

28.1

27.1

11.4

5.3

10.7

9.0

6.6

KU 98379d

Nuevo Leon

54.8

28.4

26.8

11.4

5.5

10.9

9.0

7.0

KU 98413 d

Nuevo Leon

56.8

27.5

26.3

11.0

4.9

10.5

8.9

7.0

Leptonycteris

sanborni

TTU 6564 9

Sonora

53.4

27.1

25.9

10.6

4.8

10.0

8.9

6.9

TTU 10603 9

Sonora

54.8

27.5

26.6

10.6

4.8

9.9

9.1

6.8

TTU 10604 9

Sonora

50.9

26.7

25.6

10.4

4.7

9.8

9.0

6.6

TTU 10605 9

Sonora

50.0

26.1

25.5

10.3

4.6

9.8

8.4

6.5

KU 33349(3

Jalisco

51.3

25.9

25.0

10.7

4.4

9.5

8.3

6.1

KU 34148(3

Jalisco

53.1

26.4

25.3

10.6

4.3

9.6

9.0

6.6

KU 34149(3

Jalisco

51.6

26.4

25.8

11.0

4.7

9.9

8.7

6.5

KU 34222(3

Jalisco

51.8

27.1

26.0

10.8

5.0

9.9

9.0

6.6

Lichonycteris

degener

AMNH 95118 9

Brazil

18.4

17.9

4.3

8.4

6.0

4.4

AMNH 95485 9

Brazil

32.4

USNM 239520 9

Brazil

18.8

18.2

3.8

7.9

6.0

4.2

Lichonycteris ohscura

TTU 13124 9

Nicaragua

31.7

19.2

18.0

4.1

8.0

6.2

4.4

TTU 13125 9

Nicaragua

32.2

18.4

17.6

4.0

8.1

5.9

4.4

TTU 13126 9

Nicaragua

32.6

18.8

18.2

4.0

7.7

6.3

4.4

TTU 13128 9

Nicaragua

33.0

18.8

17.9

4.2

7.9

6.0

4.4

KU 110785 (3

Nicaragua

30.7

18.2

17.0

3.9

7.9

5.7

4.3

TTU 13117(3

Nicaragua

30.3

18.0

16.8

3.9

8.1

5.5

4.2

TTU 13127(3

Nicaragua

32.1

18.5

17.3

4.0

8.2

5.8

4.1

TTU 18967(3

Nicaragua

31.9

17.9

16.9

3.9

7.9

5.5

4.5

Lionycteris spurrelli

USNM 385702 9

Venezuela

37.1

19.5

17.7

3.8

7.9

5.9

5.0

USNM 385704 9

Venezuela

36.8

20.3

18.8

4.2

7.9

6.4

5.3

USNM 385705 9

Venezuela

35.3

20.7

19.0

4.0

8.1

6.3

5.1

USNM 385706 9

Venezuela

34.8

19.5

17.5

4.1

7.5

6.2

5.1

BMNH 13.8.10.1 (3

Colombia

32.5

18.9

17.1

3.7

8.0

5.9

4.6

USNM 385698(3

Venezuela

33.4

19.3

17.8

4.0

8.2

6.0

4.9

USNM 385699(3

Venezuela

35.2

19.5

18.0

4.0

7.9

6.0

4.7

USNM 239477 6

Brazil

35.2

19.5

18.0

4.2

8.0

6.0

5.1

Lonchophylla

concava

TCWC 9826 9

Costa Rica

33.7

23.0

21.5

4.4

8.7

7.4

5.1

TCWC 9827 9

Costa Rica

33.7

22.8

21.5

4.6

9.1

7.6

5.3

TCWC 22528 9

Costa Rica

33.5

22.5

20.9

4.3

8.8

7.7

5.0

USNM 309389 9

Panama

34.4

23.6

22.0

4.5

8.9

7.9

5.4

TCWC 9828 (3

Costa Rica

34.0

22.8

21.5

4.4

8.9

7.4

5.2

TCWC 22526 c3

Costa Rica

34.4

23.3

21.6

4.4

8.8

7.6

5.2

TCWC 22527 6

Costa Rica

33.7

23.1

21.7

4.3

8.8

7.5

5.0

USNM 179621 6

Panama

33.5

23.5

22.1

4.5

9.0

7.9

5.5

Lonchophylla

hesperia

TCWC 11899 9

Peru

38.4

27.4

26.1

4.6

9.2

8.9

5.6

TCWC 23274 9

Peru

38.7

26.0

24.5

4.8

9.0

8.3

5.4

USNM 283177(3

Peru

36.0

25.5

24.5

4.8

9.1

8.6

5.8

SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

Appendix 1. — Continued.

Lonchophylla mordax

BMNH 3.9.5.32 3

Brazil

34.6

23.1

21.5

4.3

8.3

7.7

5.1

BMNH 3.9.5.33 3

Brazil

34.6

23.7

22.2

4.3

8.5

8.3

5.3

BMNH 3.9.5.34 <3

Brazil

34.3

23.8

21.7

4.3

9.1

8.0

5.3

USNM 283008 <3

Brazil

33.7

22.7

20.4

4.0

8.2

7.6

4.8

Lonchophylla rohiislu

TCWC 18945 9

Nicaragua

41.8

26.4

24.9

5.4

10.2

9.7

6.5

USNM 305237 9

Panama

42.4

26.9

25.1

5.4

10.5

9.7

6.9

USNM 483361 9

Colombia

44.3

26.9

24.8

5.1

10.1

9.4

7.0

TCWC 1 1879 9

Peru

45.0

27.4

25.6

5.1

10.4

9.9

6.3

TCWC 18944c3

Nicaragua

41.0

26.5

24.8

5.4

10.2

10.0

6.7

TTU 13137(3

Costa Rica

43.0

27.4

25.8

5.2

10.3

9.8

6.5

TTU 13138c3

Costa Rica

45.1

27.1

25.4

5.3

10.3

9.8

7.0

AMNH 230214(3

Peru

45.2

27.0

25.9

5.0

9.8

10.1

6.4

Lonchophylla thomasi

USNM 335180 9

Panama

32.0

21.7

20.3

4.1

8.0

7.0

5.1

USNM 483363 9

Colombia

31.4

21.3

19.8

4.2

8.3

6.7

5.3

ROM 33112 9

Guyana

32.4

21.2

19.4

4.2

8.3

6.7

5.2

AMNH 210688 9

Bolivia

31.8

21.8

20.2

4.2

8.0

6.8

5.2

USNM 483359(3

Colombia

31.0

21.7

19.7

4.2

8.6

6.9

5.4

AMNH 16120(3

Venezuela

31.2

20.8

19.1

4.2

8.5

6.5

5.1

ROM 31607(3

Guyana

31.9

20.2

18.7

4.2

8.3

6.2

5.0

ROM 33986(3

Guyana

33.2

20.4

18.9

4.2

8.3

6.4

5.0

Monophyllus plelhodon

TTU 20798 9

Guadeloupe

41.2

23.5

22.0

10.0

4.5

9.5

7.9

5.4

TTU 20799 9

Guadeloupe

41.7

23.5

21.6

10.0

4.6

9.5

8.2

5.6

KU 104771 9

Dominica

40.2

22.8

21.2

9.6

4.4

9.2

7.8

5.2

KU 110088 9

St. Vincent

41.4

23.0

21.5

9.4

4.6

9.3

8.0

5.4

TTU 20795 3

Guadeloupe

40.1

23.5

21.4

10.3

4.5

9.5

7.8

5.5

TTU 20796(3

Guadeloupe

42.8

23.7

21.9

10.4

4.8

9.6

8.0

5.5

TTU 20800 d

Guadeloupe

41.8

23.3

21.7

10.2

4.6

9.3

7.9

5.6

TTU 9337(3

Dominica

40.9

23.3

21.5

10.4

4.5

9.7

7.7

5.6

Monophyllus redmani

TTU 22544 9

Haiti

39.6

22.0

20.7

8.8

4.2

8.8

7.9

5.1

TTU 22545 9

Haiti

40.0

21.7

20.2

9.1

4.3

9.0

7.8

5.0

TTU 22546 9

Haiti

39.6

21.3

19.8

9.0

4.3

8.9

7.8

5.0

TTU 22547 9

Haiti

39.6

21.5

20.0

9.1

4.2

9.1

7.8

4.8

TTU 22537(3

Haiti

39.8

21.2

20.0

9.2

3.9

8.7

7.8

4.9

TTU 22548(3

Haiti

41.4

21.8

20.4

9.2

4.1

9.1

7.9

5.0

TTU 22549 (3

Haiti

40.8

22.0

20.6

9.4

4.2

9.2

7.8

4.9

TTU 22552 (3

Haiti

41.0

22.3

20.7

9.3

4.3

9.0

7.8

5.1

Musonycteris harrisoni

LACM 1 1487 9

Colima

41.8

32.0

30.8

4.4

9.2

12.5

4.9

LACM 11488 9

Colima

41.5

31.7

30.5

4.6

9.2

11.6

4.7

USNM 314689 9

Colima

42.7

32.2

31.0

4.0

9.0

12.2

4.8

USNM 324971 9

Colima

42.4

31.5

30.5

4.2

9.0

11.7

4.8

AMNH 235179(3

Colima

42.4

34.4

32.9

4.0

9.1

12.3

4.5

BMNH61. 1612(3

Colima

42.3

34.4

33.1

4.4

9.0

13.2

4.9

KU 98874 3

Colima

40.8

34.5

33.3

4.1

9.1

13.6

5.0

TTU 9307 6

Colima

42.2

33.3

32.2

4.0

8.2

12.8

4.8

Plalalina genovensium

USNM 268765 9

Peru

32.7

30.2

5.1

10.3

5.5

BMNH 27.11.19.38(3

Peru

46.1

32.7

30.3

4.9

10.3

10.7

5.3

FMNH 24336 6

Peru

48.5

31.1

29.4

4.6

9.6

10.2

5.5

MCZ 34843 3

Peru

49.6

31.9

29.9

4.7

9.5

10.7

5.8

MCZ 32948 3

Peru

50.0

32.6

30.0

4.8

9.5

11.2

5.7

BIOLOGY OF THE PHYLLOSTOMATIDAE

Appendix 1. — Continued.

Scleronycleris ega

BMNH 7.1.1.671 9

Brazil

34.7

4.3

8.7

7.5

4.8

USNM 407889 c}

Venezuela

35.0

22.0

21.2

4.5

8.8

7.7

5.0

Carolliinae

Carollia brevicauda

KU 110866 9

Nicaragua

38.9

22.0

19.4

5.2

9.2

7.0

7.6

KU 1 10870 9

Nicaragua

41.9

22.3

19.7

5.5

9.4

7.0

8.0

KU 1 10875 9

Nicaragua

39.8

22.5

19.5

5.1

9.1

6.7

7.2

KU 110878 9

Nicaragua

39.7

22.6

19.6

5.5

9.3

7.2

7.7

KU 1 10873 c}

Nicaragua

39.9

22.5

19.9

5.6

9.6

6.7

7.6

KU 110874 c}

Nicaragua

41.3

23.4

20.4

5.2

9.5

7.7

8.1

KU 110876 c}

Nicaragua

39.0

22.7

20.2

5.7

9.5

7.1

8.1

KU 110877 c}

Nicaragua

38.6

21.6

18.9

5.2

9.5

6.8

7.5

Carollia custanea

KU 110890 9

Nicaragua

36.5

19.0

17.0

5.2

8.5

6.0

6.9

KU 114871 9

Nicaragua

35.8

19.4

17.1

5.2

8.6

6.3

7.1

KU 114873 9

Nicaragua

35.8

19.4

17.0

5.2

8.9

6.3

6.9

KU 114880 9

Nicaragua

35.2

19.5

17.0

5.1

9.0

6.1

6.7

KU 110889 c}

Nicaragua

35.2

19.4

17.0

5.2

8.9

6.0

6.5

KU 110892 c}

Nicaragua

35.5

19.7

17.2

5.1

8.6

6.0

6.9

KU 114872 c}

Nicaragua

35.9

19.9

17.4

5.1

8.8

6.3

6.8

KU 114881 c }

Nicaragua

36.3

19.7

17.0

5.2

8.6

6.3

7.0

Carollia perspicillata

KU 97645 9

Nicaragua

42.3

23.7

20.7

5.4

9.6

7.7

7.8

KU 110791 9

Nicaragua

44.2

24.3

21.3

5.6

9.7

7.8

KU 114895 9

Nicaragua

42.7

23.4

20.5

5.3

9.5

7.5

7.7

KU 114896 9

Nicaragua

44.8

23.6

21.3

5.5

9.6

7.8

8.2

KU 110793 c }

Nicaragua

44.8

24.4

21.4

5.7

9.7

8.0

8.2

KU 1 10805 rf

Nicaragua

43.8

24.0

21.2

6.0

10.1

7.6

7.8

KU 110806 c}

Nicaragua

43.0

23.9

20.7

5.3

9.6

7.8

KU 114897 c}

Nicaragua

44.2

24.4

21.5

5.8

9.9

7.7

8.5

Carollia subrufa

KU 114906 9

N icaragua

37.1

21.3

18.9

5.3

9.2

6.7

7.7

KU 114908 9

Nicaragua

39.5

21.1

18.8

5.1

9.0

6.7

7.5

KU 114915 9

Nicaragua

38.4

21.0

18.5

5.0

8.9

6.6

7.5

KU 114916 9

Nicaragua

38.9

20.8

18.4

5.2

9.0

6.6

7.4

KU 1 14905 6

Nicaragua

39.5

21.5

19.0

5.3

9.4

6.9

7.5

KU 114912c }

Nicaragua

38.1

21.5

19.2

5.3

9.3

6.9

7.7

KU 1 14913 6

Nicaragua

38.0

21.7

19.3

5.3

9.2

6.7

7.7

KU 114914c }

Nicaragua

38.7

21.6

19.1

5.3

9.0

6.6

7.5

Rliinophylla alethina

USNM 483445 9

Colombia

36.1

20.4

17.8

5.4

8.8

5.2

7.1

USNM 483446 9

Colombia

35.4

20.4

17.9

5.5

8.9

4.9

6.8

USNM 483447 9

Colombia

33.5

19.0

16.7

5.3

8.8

4.7

6.8

USNM 483449 9

Colombia

37.5

21.3

18.4

5.4

9.0

5.1

6.5

USNM 324988 c}

Colombia

35.7

19.9

17.3

5.3

8.9

4.9

6.7

USNM 483448<}

Colombia

34.5

20.0

17.4

5.4

9.1

4.8

6.5

Rb inophylla fisch

terae

AMNH 94557 9

Brazil

30.5

16.8

14.6

4.8

7.8

4.5

5.9

TCWC 12102 9

Peru

30.0

17.0

14.7

5.1

7.9

4.3

6.1

USNM 364385 9

Peru

30.5

17.0

14.8

5.3

7.9

4.2

6.3

USNM 364386 9

Peru

30.0

17.0

14.7

5.1

7.6

4.3

6.2

AMNH 94555 c }

Brazil

30.6

16.8

14.7

4.7

7.4

4.4

6.1

TCWC 12096 c}

Peru

29.0

16.2

14.1

4.8

7.9

4.2

5.7

TCWC 12097 c }

Peru

29.8

16.8

14.5

5.0

8.1

4.3

5.9

SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

Appendix 1. — Continued.

Rhinophylla pnmilio

USNM 386528 5

Venezuela

34.0

18.7

16.5

5.7

8.3

4.9

6.4

USNM 386530 9

Venezuela

34.5

19.2

17.1

5.5

8.5

5.2

6.5

USNM 386531 9

Venezuela

34.8

19.4

17.4

5.5

8.2

5.4

6.6

USNM 386532 9

Venezuela

34.4

19.8

17.6

5.6

8.7

5.3

6.8

USNM 386539.3

Venezuela

34.3

19.4

17.5

5.6

8.2

5.2

6.3

USNM 386551 c5

Venezuela

32.4

19.1

16.8

5.5

8.4

5.1

6.5

USNM 393674(3

Brazil

32.3

19.3

16.9

5.5

8.2

5.1

6.5

USNM 393676(3

Brazil

33.6

18.9

16.9

5.4

8.2

4.8

6.3

Stenoderminae

Ametrida centurio

TTU 8814 9

Trinidad

32.9

16.4

13.7

11.4

4.1

8.7

4.9

8.1

TTU 8815 9

Trinidad

31.1

16.0

13.5

10.8

4.0

8.4

4.7

7.7

TTU 8816 9

Trinidad

31.7

16.5

13.6

11.2

4.4

8.2

4.9

7.9

TTU 8817 9

Trinidad

33.1

16.7

13.8

1 1.4

4.4

8.7

4.8

8.1

TTU 5215c3

Trinidad

25.2

15.1

12.2

10.3

4.5

8.3

4.0

7.0

TTU 8888(3

Trinidad

25.5

15.4

12.1

10.6

4.0

8.4

4.1

7.3

TTU 9545 (3

Trinidad

26.0

14.9

11.7

10.4

4.0

8.4

4.2

7.3

TTU 9548 (3

Trinidad

24.7

14.9

11.3

10.7

3.8

8.6

4.0

7.1

Ardops rtichollsi

TTU 20802 9

Guadeloupe

49.3

23.5

19.9

15.0

5.6

10.2

7.4

10.0

TTU 20820 9

Guadeloupe

48.8

23.2

20.2

15.0

5.8

10.5

7.5

10.1

TTU 20821 9

Guadeloupe

50.8

23.4

20.2

15.3

5.8

10.7

7.5

10.3

TTU 20822 9

Guadeloupe

51.4

24.4

20.8

15.8

5.7

10.6

7.8

10.4

TTU 20806 (3

Guadeloupe

47.9

22.3

18.7

14.9

5.9

10.6

6.8

9.7

TTU 20808 (3

Guadeloupe

47.3

22.6

19.3

15.0

5.7

10.7

7.1

9.8

TTU 20809 J

Guadeloupe

47.4

22.3

19.4

15.0

5.8

10.4

7.0

9.6

TTU 20824 6

Guadeloupe

49.6

22.4

19.4

14.7

5.6

10.4

7.1

9.8

Ariteus flavescens

TTU 21721 9

Jamaica

42.7

20.6

17.3

14.2

4.9

9.8

5.9

9.1

TTU 21773 9

Jamaica

41.3

19.8

17.1

13.9

4.7

9.6

5.9

8.9

TTU 21777 9

Jamaica

43.0

21.3

17.9

14.5

5.2

10.3

6.2

9.3

TTU 21782 9

Jamaica

43.1

20.4

17.4

14.4

4.7

9.8

5.9

8.9

TTU 21763 (3

Jamaica

37.8

18.5

15.2

12.9

4.5

9.4

5.4

8.2

TTU 21769(3

Jamaica

38.7

19.3

15.5

13.2

4.7

9.7

5.5

8.5

TTU 21774(3

Jamaica

39.8

18.6

15.7

13.0

4.6

9.2

5.3

8.2

TTU 21781 (3

Jamaica

38.1

19.2

16.0

13.6

4.7

9.5

5.4

8.4

Artibeus aztecus

TTU 12907 9

Costa Rica

48.0

23.2

20.6

13.8

5.5

10.3

7.5

10.6

TTU 12911 9

Costa Rica

46.6

22.9

20.3

13.8

5.5

10.0

7.5

10.5

TTU 12913 9

Costa Rica

44.6

22.3

19.7

12.9

5.1

9.8

7.3

10.1

TTU 12914 9

Costa Rica

45.3

23.1

20.7

13.8

5.3

10.4

7.6

10.6

KU 94141 (3

Sinaloa

42.9

22.0

19.6

13.3

5.9

9.8

7.0

9.3

KU 94142 (3

Sinaloa

44.2

22.0

19.7

12.7

5.4

9.8

7.0

9.0

TTU 12908(3

Costa Rica

46.5

22.6

20.1

13.3

5.4

10.2

7.3

10.7

TTU 12910(3

Costa Rica

42.1

21.8

19.0

12.8

5.0

9.7

7.2

9.8

Artibeus cinereus

TTU 5335 9

Trinidad

37.6

20.3

18.2

11.3

4.6

9.0

6.5

8.2

TTU 5352 9

Trinidad

39.2

20.0

18.1

12.2

5.0

8.6

6.3

8.4

TTU 5769 9

Trinidad

40.6

21.2

18.8

12.2

4.7

9.2

6.8

8.5

TTU 5859 9

Trinidad

40.2

20.5

18.3

12.1

4.9

8.5

6.5

8.6

TTU 5229 (3

Trinidad

39.4

20.4

18.5

11.2

4.8

8.9

6.6

8.8

TTU 5230(3

Trinidad

38.2

20.9

18.5

11.8

4.9

9.0

6.8

8.6

TTU 5541 (5

Trinidad

41.8

21.1

19.2

12.3

5.0

9.0

6.8

9.0

TTU 9015 (3

Trinidad

40.4

20.8

18.5

11.6

5.1

8.9

6.5

8.7

Artibeus concolor

ROM 36827 9

Guyana

48.2

21.7

18.9

13.5

5.6

9.9

6.7

9.5

ROM 36830 9

Guyana

47.4

22.0

19.5

13.1

5.1

10.0

7.3

9.4

BIOLOGY OF THE PHYLLOSTOMATIDAE

Appendix 1. — Continued.

ROM 36847 9

Guyana

46.3

22.5

19.7

13.6

5.4

10.0

7.0

9.5

ROM 60446 9

Guyana

48.8

22.4

19.8

13.0

5.3

9.4

7.0

9.4

ROM 57444 6

Guyana

46.1

20.6

17.8

12.6

5.5

9.4

6.8

9.1

ROM 59925 6

Guyana

48.4

21.5

18.8

13.1

5.3

10.0

6.8

9.1

ROM 66581 &

Guyana

49.4

21.5

19.0

13.0

5.4

9.1

7.2

9.5

ROM 67478 &

Guyana

45.0

21.3

18.4

12.8

5.3

9.6

6.8

9.2

Artibeus glaucus

AMNH 2I436I 9

Peru

38.1

20.0

17.9

11.6

5.0

9.1

6.2

8.4

AMNH 233750 9

Peru

40.8

20.6

18.4

11.5

5.4

9.1

6.5

8.4

AMNH 233751 9

Peru

41.5

20.1

17.7

11.5

5.4

8.9

6.3

8.1

AMNH 233775 9

Peru

40.1

19.6

17.2

11.2

4.7

8.9

6.1

8.1

AMNH 214363 &

Peru

37.5

19.0

17.4

10.8

4.7

8.6

6.2

8.2

AMNH 233755 <3

Peru

41.1

20.5

18.1

11.7

5.2

9.3

6.5

8.4

AMNH 233763 <3

Peru

40.7

20.2

17.7

11.7

5.0

9.1

6.3

8.6

AMNH 233771 c 3

Peru

41.0

20.3

17.9

11.7

5.0

9.3

6.3

8.5

Artibeus hirsutus

TTU 8700 9

Jalisco

56.0

27.8

24.5

16.8

6.8

11.7

10.0

12.2

TTU 8701 9

Jalisco

55.0

26.8

23.4

16.8

6.5

11.9

9.4

11.6

TTU 8703 9

Jalisco

55.2

27.6

24.4

17.3

6.8

12.3

9.7

11.9

TTU 8704 9

Jalisco

56.9

27.3

23.9

16.8

6.7

11.8

9.6

11.6

TTU 8702 (3

Jalisco

56.0

27.1

23.6

17.0

6.9

12.2

9.5

11.8

TTU 10592c3

Jalisco

55.2

26.7

23.7

16.5

6.8

12.3

9.8

11.4

TTU 10593 6

Jalisco

53.0

27.0

23.8

16.4

6.9

12.0

9.7

11.6

TTU 10596d

Jalisco

57.3

26.3

23.0

15.7

6.7

12.0

9.8

11.3

Artibeus inopinatus

TCWC9517 9

Honduras

52.8

26.1

22.2

16.2

5.6

11.6

9.0

10.9

TTU 7685 9

Honduras

50.3

25.7

21.9

15.8

5.4

11.2

8.6

10.4

TTU 7686 9

Honduras

52.0

25.8

22.2

15.6

5.4

1 1.4

8.8

10.7

TTU 12915 9

Nicaragua

51.1

25.3

21.7

15.4

5.4

11.2

8.6

10.6

TTU 7688 (3

Honduras

50.0

25.9

22.4

15.6

5.4

11.7

8.9

10.7

TTU 7689 (3

Honduras

50.0

25.2

21.5

15.7

5.3

11.3

8.7

10.6

TTU 7690 <3

Honduras

50.2

25.2

21.8

15.6

5.6

1 1.4

8.8

10.7

TTU 129163

Nicaragua

50.0

25.6

21.7

15.5

5.4

1 1.4

8.6

10.6

Artibeus jamaicensis

AS 5234 9

Jamaica

61.4

29.5

26.1

17.1

7.2

12.8

10.4

13.0

AS 5236 9

Jamaica

57.0

28.3

24.7

17.0

7.1

12.1

9.5

12.4

KU 97801 9

Nicaragua

60.1

29.3

25.7

17.4

6.9

12.7

9.8

12.0

KU 97802 9

Nicaragua

56.4

27.9

24.3

17.0

7.2

12.2

9.5

12.1

COLU 316 3

Jamaica

59.2

28.7

24.8

17.3

7.2

12.6

10.0

12.8

COLU 323 3

Jamaica

57.3

27.8

24.5

16.8

6.7

12.0

9.6

12.1

AMNH 28335 3

Nicaragua

56.4

29.4

25.7

16.9

7.0

12.4

10.4

12.5

KU 115030 3

Nicaragua

58.8

28.8

24.9

17.7

7.3

12.9

9.7

12.9

Artibeus lituratus

KU 115967 9

Nicaragua

67.3

31.7

27.7

19.2

6.4

13.2

10.4

12.9

KU 115068 9

Nicaragua

72.6

31.9

28.8

19.1

6.6

13.7

11.1

13.6

KU 115069 9

Nicaragua

70.5

31.1

27.3

18.9

6.3

13.9

10.4

13.5

KU 115072 9

Nicaragua

71.1

32.1

28.3

19.9

6.5

14.3

1 1.2

13.8

KU 115062 3

Nicaragua

69.3

31.7

27.8

19.5

7.0

14.2

10.9

13.0

KU 115065 3

Nicaragua

72.8

31.1

27.1

19.0

6.7

14.0

10.1

12.9

KU 115070 3

Nicaragua

73.0

31.9

27.9

19.6

6.4

14.0

11.1

13.6

KU 115071 3

Nicaragua

69.3

31.0

27.2

18.9

6.6

13.6

11.0

13.0

Artibeus phaeotis

KU 106145 9

Nicaragua

35.1

18.3

15.9

10.6

4.5

8.6

5.6

7.7

KU 106146 9

Nicaragua

34.2

18.3

16.0

11.3

4.8

8.5

5.7

7.5

KU 106153 9

Nicaragua

35.7

19.4

17.2

11.8

4.7

9.0

5.8

7.7

KU 106155 9

Nicaragua

34.9

18.5

16.1

11.1

4.8

8.9

5.6

7.7

KU 106147 3

Nicaragua

34.9

18.3

15.8

11.1

4.8

9.0

5.6

7.5

KU 106148 3

Nicaragua

37.2

18.1

15.7

10.9

4.5

8.4

5.6

7.7

SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

Appendix 1. — Continued.

KU 105149 <3

N icaragua

36.9

18.3

16.0

10.8

4.3

8.8

5.7

7.4

KU 106150 6

Nicaragua

36.3

19.0

17.0

11.5

4.5

8.7

5.8

7.7

Artibeus toltecus

TTU 73519

Tamaulipas

39.4

20.5

17.9

12.2

4.7

9.2

6.6

8.9

TTU 7354 9

Tamaulipas

37.5

20.8

18.1

12.5

5.1

9.3

6.6

8.9

TTU 7355 9

Tamaulipas

39.7

21.5

18.7

12.5

4.9

9.4

6.7

9.0

TTU 12930 9

Honduras

40.3

21.4

19.4

12.7

5.7

9.7

6.9

8.7

TTU 8163 i

San Luis Polosi

36.9

19.5

17.2

12.6

5.0

9.4

6.4

8.9

TTU 12929 6

Honduras

40.6

21.0

18.7

12.0

5.6

9.3

6.9

8.8

TTU 12931 6

El Salvador

39.0

20.3

17.9

11.7

5.0

9.2

6.7

9.0

TTU 12932 6

El Salvador

40.2

19.7

17.4

11.5

4.8

9.2

6.2

8.7

Artibeus walsoni

KU 82102 9

Guatemala

37.8

19.7

17.8

11.2

4.6

8.8

6.4

8.2

TTU 12964 9

Honduras

36.3

19.0

16.3

11.2

4.7

8.7

5.8

8.4

TTU 12967 9

Honduras

38.8

19.8

17.3

12.1

5.0

8.9

6.1

8.6

KU 111171 9

Nicaragua

38.5

19.9

17.6

11.5

4.9

8.7

6.6

8.4

TTU 12962 6

Honduras

37.5

19.1

16.6

11.7

4.7

9.0

6.0

8.5

TTU 12963 <3

Honduras

37.6

19.8

17.1

11.8

4.8

8.8

6.0

8.5

TTU 12934(3

Nicaragua

39.3

20.0

17.7

11.3

4.9

8.5

6.2

8.0

TTU 12948(3

Nicaragua

37.9

19.5

17.4

11.4

4.8

8.6

6.2

8.3

Centurio sene x

FHKSC 9813 9

Chiapas

45.7

18.5

15.1

15.0

5.8

10.4

4.8

10.7

TTU 13076 9

Honduras

42.6

18.9

14.5

14.9

5.7

9.3

4.8

10.6

KU 115113 9

Nicaragua

42.6

19.0

14.8

14.9

5.9

9.8

5.0

10.6

KU 115114 9

Nicaragua

43.5

18.9

15.0

6.0

10.0

4.7

10.6

FHKSC 9812(3

Chiapas

42.0

18.7

14.5

14.8

5.5

10.0

4.7

10.5

KU 115108(3

Nicaragua

41.6

18.7

14.6

14.5

5.6

10.6

4.6

10.3

KU 115111 (3

Nicaragua

42.7

19.0

14.5

15.0

5.7

10.8

4.6

10.4

TTU 5221 (3

Trinidad

44.1

19.8

15.2

15.8

6.1

10.5

5.1

11.1

Chiroderma

doriae

BMNH 9.1 1.19.15 9

Brazil

53.7

28.0

25.9

17.8

6.1

11.2

10.3

13.6

TTU 30707 (3

Brazil

28.1

25.8

17.6

6.4

12.0

10.0

13.0

TTU 30708 c3

Brazil

28.8

26.3

17.9

6.2

12.0

10.2

13.4

TTU 30709 9

Brazil

29.0

26.4

18.1

6.3

12.5

10.2

13.5

Chiroderma improvisum

TTU 19900(3

Guadeloupe

57.5

29.9

27.7

18.9

6.5

12.2

10.7

7.2

Chiroderma salvini

USNM 33871 1 9

Colima

46.1

24.2

22.0

15.2

6.2

10.6

8.6

1 1.4

TCWC 17499 9

Guatemala

47.8

26.4

23.8

16.1

6.0

11.0

9.2

11.5

TTU 12809 9

Honduras

51.8

27.6

24.8

16.9

6.1

11.2

9.5

12.1

AMNH 142484 9

Costa Rica

51.5

27.6

24.8

17.5

6.3

11.6

10.1

13.0

TTU 6123 (3

Colima

43.6

24.5

21.9

15.0

5.8

10.5

8.4

11.1

TTU 12800(3

Honduras

48.0

26.6

24.1

16.2

6.2

11.0

9.4

11.7

TTU 12801 c3

Honduras

45.6

26.0

23.6

16.0

5.7

11.0

9.1

11.8

TTU 12802c3

Honduras

49.4

26.6

24.2

16.6

6.2

11.2

9.3

12.2

Chiroderma trinitatum

TTU 5223 9

Trinidad

41.5

22.7

19.8

13.4

5.4

9.5

7.8

10.6

TTU 5224 9

Trinidad

38.0

22.5

20.0

13.3

5.3

9.4

7.5

10.1

TTU 5336 9

Trinidad

40.3

22.4

19.7

13.8

5.4

9.5

7.5

10.1

TTU 5382 9

Trinidad

38.8

22.5

19.6

13.7

5.4

9.6

7.4

10.2

TTU 5487 (3

Trinidad

39.0

22.4

19.8

13.5

5.5

9.7

7.4

10.3

TTU 5675 (3

Trinidad

38.7

22.5

19.8

13.8

5.2

9.5

7.4

9.6

TTU 8989 (3

Trinidad

39.5

22.3

19.8

13.6

5.3

9.6

7.5

9.7

TTU 9014 (3

Trinidad

39.1

22.2

19.5

13.5

5.5

9.4

7.4

10.0

Chiroderma villosum

TTU 5289 9

Trinidad

46.5

26.0

23.4

16.4

5.7

10.8

9.1

11.6

TTU 5321 9

Trinidad

45.3

25.0

22.4

16.0

5.6

11.0

8.7

11.5

BIOLOGY OF THE PHYLLOSTOMATIDAE

Appendix 1. — Continued.

TTU 5353 9

Trinidad

47.9

26.6

23.6

16.5

5.8

11.0

9.1

12.0

TTU 5354 9

Trinidad

47.2

26.2

23.4

17.0

6.2

10.4

9.0

12.0

TTU 5262 c?

Trinidad

45.9

26.4

23.3

16.4

6.2

11.3

9.1

11.5

TTU 5276 c?

Trinidad

46.0

25.3

22.4

15.7

5.9

10.5

8.5

11.4

TTU 5668 <5

Trinidad

44.3

26.0

22.9

15.7

6.1

10.8

9.0

11.6

TTU 9016 c?

Trinidad

46.8

26.5

22.8

15.1

5.8

10.6

8.6

10.9

Ectophylla

alba

KU 88025 9

Costa Rica

28.1

16.4

15.1

9.8

4.0

7.3

6.0

7.4

USNM 335318 9

Panama

29.1

16.4

15.5

10.0

4.2

7.5

6.0

7.2

USNM 335320 9

Panama

28.9

16.7

15.5

10.0

4.0

7.7

6.0

7.3

USNM 335322 9

Panama

29.4

16.3

15.2

10.3

4.3

8.0

6.0

7.6

TCWC 19372<?

Honduras

28.4

17.1

15.7

10.1

4.2

7.9

6.1

7.3

TCWC 19373 c?

Honduras

28.5

17.1

15.7

10.3

4.2

7.8

6.3

7.4

USNM 315563 c?

Panama

28.4

16.9

15.4

10.3

4.3

7.8

6.1

7.5

USNM 319426c?

Panama

28.7

16.5

15.4

9.9

4.2

7.5

6.0

7.2

Enchisthenes

hurtii

AMNH 206872 9

Oaxaca

40.1

21.1

19.1

13.0

6.1

9.5

6.8

9.0

KU 102600 9

Chiapas

39.5

20.7

18.7

12.2

5.9

9.3

6.6

8.5

TTU 5371 9

Trinidad

38.6

20.5

18.3

12.0

5.7

9.4

6.8

8.7

AMNH 233798 9

Peru

36.7

20.3

18.6

12.4

5.9

9.6

7.0

8.6

KU 97039 c?

Jalisco

39.8

21.0

18.9

12.9

5.9

9.8

6.7

8.9

AMNH 126239 c?

Honduras

36.5

20.9

18.6

11.3

5.4

9.4

7.1

8.6

BMNH 92.9.7.8 c?

Trinidad

37.1

20.4

18.5

12.0

6.1

9.1

7.2

8.4

AMNH 233599 c?

Peru

39.6

20.9

18.9

12.1

5.7

9.6

6.7

8.0

Mesophylta macconnelli

TTU 5359 9

Trinidad

32.6

18.6

16.6

10.7

4.6

8.2

6.2

7.6

TTU 5475 9

Trinidad

31.5

18.2

16.3

10.4

4.5

7.8

6.3

7.4

TTU 9786 9

Trinidad

33.5

19.0

16.7

10.8

4.8

8.3

6.5

7.7

BMNH 1.6.4.64 9

Guyana

30.0

17.7

15.5

10.1

4.4

7.8

6.0

7.0

TTU 5211 c?

Trinidad

32.0

18.5

16.5

10.6

4.7

8.2

6.1

7.5

TTU 5212 c?

Trinidad

31.5

18.5

16.3

10.8

4.5

8.3

6.2

7.5

TTU 5213 c?

Trinidad

32.3

18.7

16.6

11.0

4.6

8.2

6.2

7.7

BMNH 70.1008 c?

Brazil

29.5

17.7

15.4

9.8

4.2

7.7

5.8

7.0

Phyllops falcatus

AMNH 176190 9

Cuba

44.0

20.9

18.9

14.2

5.6

10.0

6.0

8.7

USNM 143844 9

Cuba

43.3

20.8

18.7

14.1

5.3

10.0

6.2

8.5

BMNH c?

Cuba

42.9

5.3

5.8

8.1

Phyllops halt

iensis

TTU 22675 9

Haiti

41.8

20.3

18.3

13.7

5.7

10.0

5.9

8.2

TTU 22676 9

Haiti

43.8

20.7

18.3

13.6

5.4

10.1

6.2

8.5

TTU 22677 9

Haiti

44.0

20.4

18.4

13.8

5.7

10.3

6.1

8.4

TTU 22678 9

Haiti

42.8

20.5

18.3

13.2

5.5

9.9

6.1

8.3

TTU 22697 c?

Haiti

39.0

19.4

17.2

12.5

5.2

9.6

5.5

7.8

TTU 22698 c?

Haiti

40.2

19.4

17.5

12.9

5.3

9.6

5.6

7.9

TTU 22699 c?

Haiti

40.9

19.5

17.4

13.2

5.7

9.9

5.8

7.9

TTU 22700 c?

Haiti

42.1

19.7

17.4

13.3

5.4

9.9

5.6

8.1

Phyllops veins

AMNH 41001 ?

Cuba

18.1

13.5

5.4

5.9

7.9

AMNH 41002 ?

Cuba

19.5

17.3

13.0

5.2

9.7

5.4

7.3

AMNH 41003?

Cuba

20.1

18.0

5.3

10.0

5.5

7.5

AMNH 41005 ?

Cuba

17.0

5.0

10.0

5.3

7.3

Pygoderma hilabialum

AMNH 234288 9

Paraguay

38.9

20.5

17.5

14.0

7.4

9.9

6.0

8.0

AMNH 234290 9

Paraguay

37.6

20.9

17.5

14.3

8.0

10.1

5.7

8.0

AMNH 234292 9

Paraguay

39.8

21.0

17.9

14.7

7.7

10.3

6.1

8.4

KU 92656 9

Paraguay

39.5

20.2

17.4

14.1

7.4

10.1

6.0

7.9

AMNH 234291 c?

Paraguay

36.4

20.1

16.5

13.2

7.2

10.0

5.4

7.1

100

SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

Appendix 1. — Continued.

AMNH 234294 c?

Paraguay

36.6

20.0

16.8

13.4

7.3

10.0

5.3

7.2

AMNH 234297 c?

Paraguay

36.2

20.5

17.2

13.7

7.7

10.4

5.5

7.5

AMNH 234298 c?

Paraguay

37.0

19.9

17.0

13.7

7.5

10.3

5.4

7.3

Sphaeronycleris

toxophyllum

TCWC 28252 9

Venezuela

39.5

17.2

14.2

12.1

5.6

9.5

4.7

7.9

USNM 370848 9

Venezuela

40.0

17.4

14.5

12.2

5.7

9.4

4.6

7.9

USNM 370849 9

Venezuela

40.1

17.2

14.5

12.3

5.7

9.2

4.7

7.8

AMNH 209704 9

Bolivia

39.6

17.5

14.6

12.1

5.6

9.0

4.4

8.0

TTU 10227 c?

Colombia

36.6

16.1

13.8

11.7

5.5

8.9

4.4

7.2

USNM 405688 c?

Venezuela

37.0

16.8

13.9

12.2

5.6

9.5

4.3

7.3

USNM 409233 c?

Venezuela

37.3

16.5

13.4

11.9

5.6

8.9

4.4

7.4

AMNH 209741 c?

Bolivia

38.7

16.9

13.8

12.4

5.7

9.0

4.2

7.6

Stenoderma rufum

TTU 8876 9

Puerto Rico

49.0

23.0

19.4

15.5

5.7

11.4

7.2

10.1

TTU 8879 9

Puerto Rico

49.0

22.5

19.1

15.2

5.7

10.6

6.8

9.7

TTU 8880 9

Puerto Rico

51.2

23.5

19.8

15.8

5.7

11.4

7.0

10.2

TTU 8884 9

Puerto Rico

50.3

22.9

19.4

15.3

5.7

10.7

7.0

10.0

TTU 8860<?

Puerto Rico

46.5

22.2

18.5

15.0

5.5

10.5

6.6

9.7

TTU 8861 <J

Puerto Rico

47.1

22.5

19.0

14.9

5.6

10.6

6.6

9.7

TTU 8864 $

Puerto Rico

46.1

22.0

18.0

14.4

5.2

10.2

6.2

9.7

TTU 8865 <?

Puerto Rico

48.5

22.5

18.7

14.9

5.4

10.7

6.3

9.5

Sturnira aratathomasi

ROM 70874 9

Colombia

58.0

29.1

25.4

17.2

7.5

12.9

7.6

10.2

USNM 501064 9

Colombia

57.5

28.5

25.5

16.9

6.9

12.5

7.7

10.1

USNM 501066 9

Colombia

56.8

28.8

25.0

16.7

7.2

12.5

7.4

9.7

ROM 46349 9

Ecuador

60.5

29.7

26.2

17.5

7.2

12.8

8.1

10.5

ROM 70875 <?

Colombia

57.7

29.4

26.5

16.7

7.2

12.8

7.8

10.4

ROM 70876 e?

Colombia

54.8

28.8

25.2

16.8

7.0

12.7

7.6

10.2

USNM 395158 c?

Colombia

57.1

29.4

26.5

17.5

7.3

13.0

7.9

10.1

USNM 501065 c?

Colombia

57.5

28.8

25.9

16.5

6.9

12.3

7.7

10.0

Sturnira i

bide ns

USNM 386557 9

Venezuela

39.3

21.2

18.9

11.7

5.5

9.4

6.0

6.8

USNM 386558 9

Venezuela

40.2

21.6

19.7

11.7

5.3

9.7

6.0

6.8

USNM 386560 9

Venezuela

39.7

22.1

19.6

12.0

5.5

9.8

6.1

7.1

USNM 386562 9

Venezuela

40.8

21.7

19.5

12.0

5.5

9.6

6.0

6.9

USNM 386559 c?

Venezuela

39.7

21.2

19.0

11.9

5.4

9.6

5.7

6.9

USNM 386567 c?

Venezuela

39.7

21.3

18.7

11.7

5.4

9.5

5.9

7.0

USNM 386570 c?

Venezuela

39.5

21.0

18.7

11.7

5.3

9.6

5.9

6.6

AMNH 214349 c?

Peru

41.2

21.3

18.7

11.7

5.4

9.7

5.9

6.7

Sturnira erythromos

ROM 67254 9

Colombia

39.3

21.3

18.6

12.7

5.7

10.0

6.0

8.0

ROM 67267 9

Colombia

41.1

20.7

18.3

12.0

5.3

9.5

5.9

7.5

USNM 483451 9

Colombia

40.6

21.5

19.0

12.7

6.0

9.9

5.8

7.5

USNM 483452 9

Colombia

40.6

21.0

18.9

12.5

6.1

9.7

6.0

7.4

ROM 67270 c?

Colombia

41.6

21.4

19.2

12.1

6.0

9.6

5.9

7.4

BMNH 15.7.1 1.13 c?

Ecuador

40.8

22.0

19.4

12.9

6.0

10.4

6.3

8.0

Sturnira

1 ilium

TTU 5367 9

Trinidad

42.5

23.4

20.4

13.6

6.0

10.3

6.4

8.4

TTU 5407 9

Trinidad

43.9

22.9

20.2

13.7

5.8

10.5

6.4

8.2

TTU 5669 9

Trinidad

42.4

22.7

19.9

13.4

5.6

10.1

6.5

8.0

TTU 5670 9

Trinidad

41.4

22.8

19.9

13.6

6.2

10.4

6.3

8.0

TTU 5408 c?

Trinidad

43.2

23.1

20.4

13.8

6.0

10.4

6.6

8.2

TTU 5415 c?

Trinidad

41.9

23.2

20.3

13.7

6.3

10.5

6.3

7.9

TTU 5775 c?

Trinidad

41.3

22.9

20.4

13.6

6.4

10.5

6.8

8.5

TTU 5776 c?

Trinidad

42.7

22.4

19.6

13.4

5.9

10.5

6.4

8.0

BIOLOGY OF THE PHYLLOSTOMATIDAE

101

TTU 15543 9
TTU 15546 9
TCWC 14359 9
TCWC 14360 9
TTU 7341 £
TTU 6124 £
TTU 6125 £
KU 97689 £

AMNH 214347 9
TCWC 27474 9
LSU 16518 9
LSU 19031 9
LSU 16517(3
LSU 19027 £
LSU 19028<J

BMNH 69.1263 9
TCWC 10034(3
TCWC 10035c3
TCWC 10041 £
TCWC 10042 £

AMNH 219138 9
LSU 16521 9
LSU 16522 9
LSU 16524 9
AMNH 219171 £
AMNH 219172 (3
AMNH 219173 £
TCWC 28071 £

TTU 19904 9
TTU 19905 9
TTU 19906 9
TTU 19907 9
AMNH 234950(5
USNM 361883 (3

TTU 5406 9
TTU 5667 9
TTU 5786 9
TTU 5791 9
TTU 5337 £
TTU 5372 £
TTU 5402 £
TTU 5454(3

KU 114985 9
TTU 5327 9
TTU 5485 9
TTU 5813 9
KU 114986(5
TTU 5254(3
TTU 5300(3
TTU 5301 £

Appendix 1. — Continued.

Sturnira ludovici

Hidalgo

44.1

24.0

20.8

13.0

6.1

10.3

6.5

8.0

Hidalgo

43.2

24.0

20.8

13.5

6.1

10.2

6.5

8.0

Guatemala

45.1

23.2

20.1

13.2

5.6

10.3

6.3

8.0

Guatemala

46.9

24.4

21.6

13.9

6.1

10.6

6.6

8.3

Tamaulipas

42.5

23.9

21.0

13.6

6.0

10.6

6.4

8.3

Jalisco

44.3

23.4

20.3

13.9

6.1

10.0

6.2

7.9

Jalisco

43.2

23.2

20.1

12.7

6.0

10.1

6.2

7.8

Nicaragua

45.1

24.0

21.4

14.2

6.2

10.4

6.5

8.3

Sturnira magna

Peru

59.2

29.0

25.3

16.7

6.9

12.0

7.1

9.0

Peru

57.7

27.9

24.7

16.4

7.0

11.8

7.2

8.8

Peru

57.7

29.1

25.6

17.2

7.0

12.5

7.4

9.9

Peru

57.4

28.5

24.4

16.0

6.8

11.5

7.3

8.8

Peru

57.0

29.5

25.6

17.2

7.0

11.9

7.4

9.3

Peru

55.4

28.5

24.7

17.0

7.0

12.1

7.1

9.1

Peru

56.0

28.8

24.9

16.9

6.9

12.2

7.5

9.3

Sturnira mordax

Costa Rica

46.2

25.8

22.4

13.1

5.9

10.6

6.7

7.8

Costa Rica

48.3

26.1

22.9

13.8

6.1

10.9

6.9

8.2

Costa Rica

46.1

25.5

22.0

13.3

5.9

11.0

6.7

7.9

Costa Rica

47.7

25.7

22.4

13.3

6.0

10.7

6.7

7.8

Costa Rica

48.3

26.3

23.1

13.7

6.2

10.9

6.9

8.0

Sturnira

nana

Peru

34.7

18.8

16.6

10.2

4.6

8.2

4.8

5.8

Peru

34.8

18.5

16.5

10.0

4.7

8.1

4.7

5.6

Peru

33.7

18.9

16.6

9.8

4.8

8.3

4.8

5.7

Peru

34.1

19.0

16.8

10.1

4.6

8.5

4.9

6.0

Peru

34.5

18.7

16.5

10.1

4.7

8.5

4.7

5.5

Peru

35.4

18.8

16.5

10.1

4.6

8.5

4.9

5.8

Peru

35.0

18.5

16.3

9.7

4.7

8.2

4.7

5.6

Peru

32.6

18.4

16.1

9.9

4.7

8.1

4.8

5.7

Sturnira thomasi

Guadeloupe

45.9

25.3

23.3

12.1

5.7

9.8

7.0

8.1

Guadeloupe

46.4

24.4

22.4

11.9

5.6

9.5

6.7

7.7

Guadeloupe

46.1

24.9

22.9

12.2

5.5

9.8

6.9

8.0

Guadeloupe

47.7

25.1

23.6

12.5

5.9

9.6

6.9

8.0

Guadeloupe

46.5

25.1

23.7

12.2

5.7

9.5

6.7

8.2

Guadeloupe

48.1

26.2

24.7

12.7

6.0

9.9

7.7

8.2

Sturnira tildae

Trinidad

44.0

23.6

21.1

14.6

6.0

10.7

6.8

8.1

Trinidad

44.1

23.9

21.5

14.3

6.1

10.8

6.9

8.5

Trinidad

44.7

24.3

21.8

14.3

5.7

10.6

7.1

8.3

Trinidad

43.4

22.8

20.2

13.7

6.1

10.4

6.6

7.8

Trinidad

44.7

23.9

21.1

14.2

6.3

10.7

7.1

8.5

Trinidad

44.5

23.1

20.5

14.0

5.9

10.6

6.9

8.2

Trinidad

46.3

24.4

22.2

14.7

6.6

10.7

7.4

8.6

Trinidad

44.2

23.7

21.5

14.6

6.5

11.0

6.8

8.8

Uroderma bilobatum

Nicaragua

41.6

22.8

20.2

13.0

5.4

9.5

7.9

9.3

Trinidad

42.1

23.6

20.8

12.8

4.6

9.5

8.2

9.4

Trinidad

39.6

23.0

20.6

12.5

5.3

9.3

7.9

9.3

Trinidad

42.4

23.6

21.4

13.0

5.6

9.4

8.5

9.9

Nicaragua

43.0

22.6

19.9

12.7

5.3

9.9

7.8

9.1

Trinidad

43.1

24.7

22.1

13.4

5.7

9.9

8.6

9.9

Trinidad

40.5

23.7

21.6

13.1

5.4

9.7

8.3

9.5

Trinidad

41.4

24.0

21.0

12.8

5.5

9.4

7.9

8.9

102

SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

Appendix 1. — Continued.

Uroderma magnirostrum

TTU 17111 9

El Salvador

42.3

23.0

20.8

12.7

5.7

9.7

7.6

8.9

KU 114987 9

Nicaragua

45.1

23.9

21.8

13.1

5.8

9.3

8.3

9.1

TTU 9080 9

Colombia

42.6

23.1

21.0

12.6

5.6

9.3

8.1

9.2

TTU 9517 9

Colombia

41.8

22.4

20.3

12.2

5.6

9.3

7.8

8.9

TCWC 17189 6

Honduras

41.0

22.5

20.5

12.6

5.6

9.5

7.6

8.6

KU 106109 c?

Nicaragua

41.6

23.6

21.7

12.9

5.5

9.5

7.7

8.7

TTU 9054 c?

Colombia

43.6

24.0

21.7

13.5

6.0

10.3

8.1

9.5

TTU 9056 c?

Colombia

43.4

23.5

21.3

12.9

5.3

9.7

8.0

9.1

Vampyressa bidens

ROM 59895 9

Guyana

36.0

20.2

17.2

11.8

5.4

9.1

6.3

8.3

ROM 66587 9

Guyana

38.2

20.5

17.8

12.3

5.3

9.0

6.4

8.9

AMNH 208072 9

Peru

36.6

20.4

17.8

12.1

5.1

8.8

6.5

8.9

TCWC 27508 9

Peru

36.4

19.5

16.8

11.4

4.8

8.6

5.9

8.1

AMNH 98780.?

Peru

39.1

20.0

17.1

12.2

5.2

9.1

6.1

8.6

TCWC 27503 c?

Peru

35.6

20.0

17.3

11.7

5.2

9.3

6.2

8.5

TCWC 27505 <?

Peru

35.3

19.8

17.1

11.2

5.2

8.4

6.3

8.2

TCWC 27506 c?

Peru

35.5

20.2

17.3

12.2

5.4

9.1

6.3

8.3

Vampyressa brocki

TTU 8827 9

Colombia

35.4

18.4

16.0

10.9

4.9

8.4

5.7

7.9

TTU 8832 9

Colombia

32.1

18.3

15.8

10.8

4.7

8.4

5.7

7.6

TTU 9047 9

Colombia

33.2

18.4

16.2

10.7

5.1

8.0

5.7

7.8

ROM 38515 9

Guyana

33.0

17.7

15.5

10.4

4.7

7.8

5.6

7.6

Vampyressa melissa

BMNH 26.5.3.4 9

Peru

37.1

21.5

19.6

12.8

5.0

9.0

6.8

9.6

LSU 16580 9

Peru

39.2

22.2

20.4

13.2

5.2

9.5

7.1

9.5

LSU 16583 9

Peru

38.2

21.8

20.1

12.9

5.1

8.9

6.7

9.2

LSU 19100 9

Peru

37.3

21.3

19.8

13.1

5.1

8.9

6.6

9.1

USNM 319283 c?

Panama

37.9

22.8

21.2

12.0

5.1

8.8

7.6

9.1

USNM 319284 c?

Panama

36.5

22.8

21.3

12.1

5.2

9.1

7.5

9.2

USNM 319285 c?

Panama

37.8

22.8

21.3

12.3

5.1

9.0

7.7

9.3

AMNH 233769 c?

Peru

36.5

21.9

20.0

13.1

5.2

9.4

6.9

9.4

Vampyressa nymphaea

KU 115005 9

N icaragua

36.2

21.1

18.4

12.3

4.7

9.2

7.0

8.6

TCWC 19368 9

Nicaragua

35.7

21.2

18.0

12.1

4.9

9.2

7.1

8.7

TTU 12611 9

Nicaragua

37.9

21.6

19.1

13.0

4.6

9.4

7.5

9.4

USNM 483687 9

Colombia

39.0

21.6

18.7

12.2

4.8

9.3

7.0

8.8

TCWC 19367 c?

Nicaragua

38.2

21.7

18.7

12.4

4.5

9.3

7.1

8.8

TTU 12612 c?

Nicaragua

37.4

22.0

19.0

12.8

5.0

9.5

7.5

9.1

AMNH 233189 c?

Colombia

37.0

21.2

18.4

11.9

4.9

9.3

6.8

8.9

BMNH 9.7.17.40 <?

Colombia

34.9

21.0

18.3

12.2

4.7

9.2

7.2

8.8

Vampyressa pusilla

KU 114082 9

Nicaragua

31.2

18.2

16.5

10.3

4.7

8.1

5.7

7.7

KU 114084 9

Nicaragua

30.0

17.9

16.2

10.2

4.7

8.5

5.5

7.8

KU 114085 9

Nicaragua

31.1

18.7

16.7

10.8

4.7

8.3

6.0

7.9

KU 114086 9

Nicaragua

30.1

18.2

16.5

10.6

4.6

7.8

6.0

7.8

TTU 12894 c?

Honduras

29.9

18.0

15.9

10.5

4.9

8.2

5.8

8.0

KU 114083 c?

Nicaragua

31.9

18.6

16.0

10.5

4.7

8.5

6.0

7.9

TTU 9431 c?

Colombia

31.9

18.1

16.0

10.5

4.5

7.7

5.5

7.3

TTU 9480 <?

Colombia

32.5

18.6

16.7

10.5

4.6

8.1

5.7

7.5

Vampyrodes caraccioli

KU 11 1033 9

Nicaragua

53.7

28.1

24.3

17.0

6.6

11.6

9.6

12.2

KU 111035 9

Nicaragua

53.9

28.3

24.8

17.8

6.9

11.8

9.9

12.4

TTU 5288 9

Trinidad

49.5

25.7

22.7

16.1

6.5

10.8

9.1

11.4

TTU 5355 9

Trinidad

49.5

26.3

22.6

16.2

6.1

11.0

9.1

11.4

KU 111034 c?

Nicaragua

53.3

28.3

24.9

17.8

6.7

11.9

9.7

12.7

TTU 5366 c?

Trinidad

46.8

25.9

22.5

16.0

6.2

10.9

9.0

11.2

BIOLOGY OF THE PHYLLOSTOMATIDAE

103

Appendix 1. — Continued.

TTU 5373 &

Trinidad

47.2

25.8

22.4

16.2

6.2

10.8

8.6

11.3

TTU 5509 6

Trinidad

47.4

26.0

22.5

16.1

6.0

11.2

9.0

11.5

Vampyrops

aurarius

USNM 387157 9

Venezuela

52.0

28.2

25.2

16.0

6.5

11.3

10.6

12.5

USNM 387159 9

Venezuela

51.9

28.8

25.5

17.0

6.6

11.6

10.8

13.0

USNM 387171 9

Venezuela

52.5

29.1

26.0

16.8

6.6

11.8

10.5

12.1

USNM 387172 9

Venezuela

53.4

29.9

27.0

17.8

6.6

11.9

11.0

13.0

USNM 387153 6

Venezuela

52.4

28.8

25.9

16.5

6.6

11.8

10.6

12.3

USNM 387154 <3

Venezuela

51.1

29.5

26.9

16.8

6.8

11.8

11.0

12.8

USNM 387155 c?

Venezuela

51.4

29.3

25.9

16.6

6.7

12.2

10.9

12.6

USNM 387161 c?

Venezuela

50.4

28.4

25.9

16.8

6.6

11.4

10.5

12.4

Vampyrops brachycephalus

TCWC 29658 9

Brazil

37.2

20.8

18.1

11.8

5.1

9.0

6.8

8.2

AMNH 230639 9

Peru

36.7

21.2

18.5

12.5

5.4

9.3

7.0

8.8

TCWC 12184 9

Peru

37.8

20.8

18.4

12.2

5.1

9.2

7.0

8.6

TCWC 12185 9

Peru

36.9

20.1

18.4

12.5

5.4

9.5

7.0

8.9

TCWC 29657 c?

Brazil

37.5

20.9

18.3

12.5

5.5

9.3

7.1

8.8

TCWC 12177 6

Peru

37.5

20.5

18.1

11.8

5.3

9.2

6.9

8.2

TCWC 12178 6

Peru

36.8

21.0

18.3

12.3

5.3

8.9

7.0

8.6

TCWC 12193 c?

Peru

40.7

21.9

19.2

13.4

5.7

9.6

7.8

9.9

Vampyrops dorsalis

AMNH 235778 9

Colombia

49.1

28.3

25.2

16.8

6.5

11.7

10.6

11.5

AMNH 235779 9

Colombia

47.2

27.2

24.0

16.1

6.8

11.7

10.0

11.4

AMNH 233614 9

Peru

48.0

26.7

24.0

15.4

6.0

11.1

10.4

11.8

AMNH 233615 9

Peru

50.5

26.7

24.2

16.0

6.1

10.8

10.3

12.1

AMNH 233186 6

Colombia

50.7

29.2

25.9

17.6

6.5

12.3

11.6

11.9

AMNH 233187 c?

Colombia

49.6

28.8

25.6

17.0

6.4

11.7

10.4

11.5

BMNH 99.12.5.1 c?

Ecuador

48.2

27.4

25.2

15.3

6.3

10.8

10.6

11.5

AMNH 214356c?

Peru

49.3

28.0

25.0

17.0

7.0

11.4

10.5

13.0

Vampyrops helleri

KU 106131 9

Nicaragua

38.0

22.3

19.8

11.9

5.2

8.8

7.5

8.7

KU 106133 9

Nicaragua

36.3

21.8

19.5

12.1

5.2

9.2

7.9

9.0

KU 106134 9

Nicaragua

37.1

21.4

19.0

11.8

5.2

8.9

7.3

8.4

FHKSC 8839 9

Colombia

37.0

22.5

20.2

12.7

5.3

9.0

7.8

9.0

FHKSC 9734 c?

Chiapas

38.6

22.8

20.5

13.0

5.5

9.3

7.9

9.3

KU 106129 c?

Nicaragua

37.6

21.7

19.3

12.2

5.6

10.0

7.5

8.8

KU 106130 c?

Nicaragua

38.5

21.2

18.8

12.1

5.2

8.9

7.4

8.4

KU 106131 c?

Nicaragua

38.7

22.3

20.2

12.3

5.5

9.2

7.7

8.9

Vumpyrops infuscus

AMNH 67661 9

Ecuador

55.9

30.6

27.4

18.3

6.3

12.4

10.9

13.8

AMNH 67664 9

Ecuador

55.0

29.9

26.2

17.6

6.5

12.0

11.1

13.4

AMNH 236131 9

Peru

54.9

30.5

27.3

17.8

6.5

12.3

11.5

13.1

AMNH 236132 9

Peru

55.0

30.1

26.9

17.6

6.4

12.2

11.3

12.8

TTU 9494 c?

Colombia

53.4

29.5

26.7

17.5

6.8

11.7

11.6

13.8

AMNH 67662 c?

Ecuador

55.5

30.5

27.3

17.8

7.0

12.2

11.3

13.8

AMNH 67663 c?

Ecuador

54.6

30.9

27.2

18.1

6.8

12.1

12.0

13.7

AMNH 233729 c ?

Peru

56.9

30.5

27.3

18.6

6.8

12.4

11.6

13.8

Vampyrops lineutus

AMNH 37013 9

Brazil

48.5

25.5

22.5

14.3

6.3

10.7

9.1

10.5

AMNH 37015 9

Brazil

47.6

25.0

22.4

14.6

6.2

10.2

8.8

10.3

AMNH 37016 9

Brazil

46.2

24.3

21.8

14.1

6.3

10.4

8.3

10.2

AMNH 205185 9

Paraguay

47.0

25.4

22.5

14.3

6.4

10.4

8.8

10.2

AMNH 36995 c?

Brazil

46.6

24.5

22.0

13.7

5.9

10.3

9.0

9.8

AMNH 148666 c?

Paraguay

50.1

25.0

22.1

14.0

6.3

10.5

8.6

10.2

AMNH 205184 c?

Paraguay

44.6

24.6

21.9

14.1

6.4

10.6

8.7

10.1

AMNH 234285 <3

Paraguay

45.8

25.2

22.2

14.4

6.4

10.7

8.8

10.1

104

SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

Appendix 1. — Continued.

Vampyrops nigettus

AMNH 233686 9

Peru

43.9

25.2

22.8

14.2

6.0

10.3

9.2

10.6

AMNH 233710 9

Peru

44.1

24.6

22.0

13.9

5.9

10.4

8.9

10.1

AMNH 233716 9

Peru

43.1

25.0

22.3

14.8

6.0

10.6

9.3

10.7

AMNH 236106 9

Peru

44.4

25.2

22.2

14.4

6.0

10.8

9.0

10.8

AMNH 214353 c?

Peru

43.2

24.4

22.2

13.9

5.9

10.3

9.0

10.3

AMNH 233644 c?

Peru

41.1

24.4

21.8

13.5

5.6

10.3

9.0

10.2

AMNH 233646 c?

Peru

43.5

25.2

22.8

14.0

5.8

10.4

9.2

10.3

AMNH 2361 1 1 c?

Peru

44.3

25.3

22.9

14.4

5.9

10.6

8.8

10.1

Vampyrops recifinus

BMNH 93.1.9.15 9

Brazil

42.1

23.7

21.5

14.0

5.7

10.2

8.7

10.2

BMNH 81.3.16.4c?

Brazil

40.6

24.0

21.3

14.0

5.7

10.2

8.7

10.4

Vampyrops vil talus

TTU 12891 9

Costa Rica

63.7

34.0

30.6

19.8

7.7

13.4

13.0

14.6

K.U 93925 9

Panama

60.1

32.7

29.6

19.1

7.6

13.5

12.7

14.0

TTU 9439 9

Colombia

64.3

33.4

31.1

19.9

7.6

12.8

13.6

15.2

AMNH 233725 9

Peru

57.1

32.9

30.1

20.0

7.4

13.5

13.2

14.8

TCWC 10051 c?

Costa Rica

61.5

32.8

29.6

19.5

7.2

13.2

12.5

14.5

KU 99355 c?

Panama

57.8

32.4

28.6

18.8

7.4

13.0

12.5

14.3

AMNH 233718 c?

Peru

59.5

32.8

29.6

20.7

7.3

13.3

12.7

15.5

AMNH 233728 c?

Peru

57.6

31.6

28.6

19.0

6.6

12.9

14.7

Brachyphyllinae

Brachyphylla cavernarum

TTU 20972 9

Guadeloupe

66.4

32.1

28.7

17.9

6.3

13.1

1 1.2

12.3

TTU 20989 9

Guadeloupe

63.3

30.9

27.4

16.6

6.2

12.7

10.7

11.6

TTU 20991 9

Guadeloupe

64.2

32.3

28.8

17.3

6.3

13.0

11.0

12.1

TTU 20995 9

Guadeloupe

66.0

31.0

27.7

17.2

6.5

12.6

10.7

11.6

TTU 20970 c?

Guadeloupe

63.5

31.1

27.9

17.2

6.5

12.6

11.0

11.7

TTU 20977 c?

Guadeloupe

68.7

32.6

29.0

16.9

6.3

12.8

11.0

11.6

TTU 20980 c?

Guadeloupe

66.4

31.8

28.2

17.6

6.5

12.4

10.7

12.1

TTU 20985 <?

Guadeloupe

65.3

31.1

27.1

16.6

6.6

12.5

11.2

11.7

Brachyphylla nana

AMNH 19085 9

Cuba

58.1

28.0

24.7

15.0

6.0

11.5

9.0

10.0

AMNH 19090 9

Cuba

59.1

28.9

25.8

14.7

6.2

11.4

9.8

10.5

TTU 22762 9

Haiti

58.8

28.1

25.3

14.9

6.3

11.7

9.4

9.9

TTU 22764 9

Haiti

58.3

28.1

24.8

14.6

6.4

11.9

9.5

10.1

AMNH 214390 c?

Dominican Republic

56.7

28.2

25.2

14.6

6.5

11.3

9.5

9.8

AMNH 214393 c?

Dominican Republic

57.9

28.6

25.1

14.5

6.5

11.8

9.4

9.6

TTU 22760 c?

Haiti

58.0

28.4

25.1

15.5

6.3

11.7

9.5

10.1

TTU 22761 c?

Haiti

58.8

28.2

25.0

14.7

6.2

11.2

9.5

9.4

Erophylla bombifrons

AMNH 97591 9

Dominican Republic

47.7

23.8

21.7

11.3

4.6

10.0

7.7

6.4

AMNH 212998 9

Dominican Republic

46.6

23.6

21.5

11.1

4.5

9.7

7.5

6.4

ROM 45709 9

Dominican Republic

47.1

24.0

22.1

10.8

4.5

9.6

7.6

6.1

TTU 22767 9

Haiti

46.8

24.4

22.5

1 1.7

4.5

10.2

8.1

6.7

ROM 45710 c?

Dominican Republic

45.9

24.3

21.9

11.5

4.5

10.0

7.5

6.7

ROM 72710c?

Dominican Republic

49.7

24.5

22.3

11.2

4.6

10.1

8.1

6.4

AMNH 39339c?

Puerto Rico

48.8

24.7

22.4

11.6

4.6

10.2

7.8

6.5

AMNH 39340 c?

Puerto Rico

48.8

24.8

22.5

11.8

4.5

10.3

7.9

6.7

Erophylla sezekorni

AS 5814 9

Jamaica

47.9

24.5

22.6

11.5

4.7

9.7

8.0

6.6

AS 5815 9

Jamaica

47.9

24.7

22.5

11.0

4.1

9.7

8.2

6.3

AS 5816 9

Jamaica

46.5

24.0

22.1

11.0

4.4

9.5

8.0

6.5

AS 5817 9

Jamaica

49.1

24.8

22.7

11.0

4.7

9.7

8.1

6.6

AMNH 45178 c?

Jamaica

45.4

25.7

23.3

11.8

4.7

10.1

8.2

6.8

AMNH 45179 c?

Jamaica

47.8

25.5

22.6

11.3

4.5

10.0

8.0

6.6

BIOLOGY OF THE PHYLLOSTOMATIDAE

105

Appendix 1. — Continued.

AMNH 45181 6

Jamaica

45.5

25.3

22.9

11.4

4.5

10.0

8.2

6.6

AMNH 45 1 82 d

Jamaica

45.4

24.1

22.9

11.2

4.5

9.7

7.8

6.2

Pliyllonycteris aphylla

TTU 21907 9

Jamaica

46.0

24.6

22.4

4.9

9.7

7.5

6.7

TTU 21908 9

Jamaica

45.3

24.9

22.6

5.0

9.9

7.8

6.8

TTU 21913 9

Jamaica

45.4

24.4

22.4

5.1

9.8

7.8

6.8

TTU 21914 9

Jamaica

44.8

23.9

21.8

5.0

9.5

7.6

6.8

TTU 21905 6

Jamaica

48.3

25.8

23.5

4.7

10.0

8.0

7.0

TTU 2 1 906 6

Jamaica

44.3

25.2

22.8

4.8

10.2

7.9

6.9

TTU 21909 d

Jamaica

47.6

24.7

22.7

5.2

9.9

7.9

7.0

TTU 21915 <3

Jamaica

46.0

25.1

23.1

5.2

10.1

7.9

6.9

Pliyllonycteris major

AMNH 40925 ?

Puerto Rico

26.3

24.6

5.7

11.4

8.4

8.1

AMNH 40926 ?

Puerto Rico

26.8

25.1

5.9

1 1.1

8.7

7.9

AMNH 40927 ?

Puerto Rico

27.0

25.2

5.6

11.0

8.5

7.8

AMNH 40928 ?

Puerto Rico

25.9

5.8

8.8

8.3

Pliyllonycteris poeyi

USNM 103548 9

Cuba

47.6

24.5

22.5

5.4

10.0

7.7

6.9

USNM 103588 9

Cuba

46.5

23.9

21.5

5.2

10.5

7.4

6.8

USNM 103589 9

Cuba

46.2

23.7

21.6

5.3

10.7

7.0

6.8

USNM 103592 9

Cuba

46.6

24.3

22.2

5.3

10.0

7.5

6.9

USNM 103537 d

Cuba

46.9

25.3

23.0

5.7

10.6

7.8

7.3

USNM 103586 d

Cuba

46.5

24.8

22.5

5.4

10.8

7.3

6.8

USNM 103597 d

Cuba

46.9

25.7

23.9

5.3

10.3

7.9

7.4

USNM 103600.3

Cuba

47.1

24.8

22.6

5.4

10.4

7.6

7.2

Pliyllonycteris poey

i obtusa

TTU 22783 9

Haiti

49.8

24.2

22.1

5.5

10.2

7.1

TTU 22792 9

Haiti

46.4

23.9

21.6

5,7

10.9

7.4

6.9

TTU 22793 9

Haiti

47.2

24.0

22.1

5.5

10.8

7.2

7.0

TTU 22794 9

Haiti

47.7

23.7

22.0

5.6

10.0

7.4

7.2

TTU 22772 3

Haiti

47.8

25.2

22.6

5.5

10.5

7.5

7.2

TTU 22773 3

Haiti

47.5

24.8

22.4

5.5

10.4

7.6

6.7

TTU 22782 3

Haiti

48.7

25.4

22.9

5.4

10.5

7.4

TTU 22783 3

Haiti

48.7

24.7

22.4

5.5

10.2

7.3

6.9

Desmodontinae

Desmodus rotund us

TTU 8228 9

Tamaulipas

60.2

24.8

21.4

11.7

5.1

11.6

3.5

6.1

TTU 8170 9

San Luis Potosi

60.1

25.0

21.5

11.9

5.2

11.9

3.3

6.0

KU 111209 9

Nicaragua

62.2

25.2

21.3

12.4

5.6

12.3

3.1

6.1

KU 111210 9

Nicaragua

60.1

25.0

21.0

12.1

5.4

11.7

3.4

6.5

TTU 9927 3

San Luis Potosi

56.9

24.2

20.8

11.4

5.5

11.8

3.4

5.7

KU 111204 3

Nicaragua

58.6

24.2

20.8

11.8

5.3

12.0

3.4

6.2

TTU 5426 3

Trinidad

55.3

23.9

20.3

11.5

5.2

11.9

3.5

5.8

TTU 5894 3

Trinidad

55.5

23.5

20.3

11.7

5.2

11.8

3.5

5.7

Diaemus youngii

USNM 409368 9

Venezuela

53.4

25.3

21.7

13.8

6.1

13.2

3.4

6.0

USNM 409374 9

Venezuela

54.5

26.0

21.7

14.1

6.1

13.2

3.4

6.2

USNM 409375 9

Venezuela

53.5

24.8

21.1

14.1

6.5

12.6

3.8

6.4

TTU 5232 9

Trinidad

51.0

24.1

20.4

13.6

6.2

13.0

3.2

6.0

USNM 405767 3

Venezuela

51.0

24.3

20.2

13.5

6.4

12.9

3.4

5.8

TTU 5233 3

Trinidad

51.3

25.4

21.5

14.3

6.1

13.1

3.3

5.8

TTU 5411 3

Trinidad

49.5

24.7

20.7

13.4

6.0

13.0

3.1

5.9

TTU 5428 3

Trinidad

50.1

25.1

21.2

14.4

6.0

13.5

3.3

6.0

Diphylla ecaudata

TTU 5658 9

Texas

53.7

23.5

20.5

13.0

7.6

11.8

3.3

5.9

TTU 10171 9

Tamaulipas

55.3

23.5

19.6

13.0

7.2

11.7

3.4

6.2

106

SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

Appendix 1. — Continued.

TTU 10157 9

Veracruz

55.8

24.0

20.1

12.9

7.6

11.6

3.6

6.2

KU 115131 9

Nicaragua

56.1

23.0

20.0

12.6

7.0

11.1

3.6

5.9

TTU lOOOOrf

Veracruz

54.0

23.5

19.9

12.8

7.3

11.8

3.5

5.7

KU 97854 d

Nicaragua

56.1

23.8

20.6

13.0

7.4

11.6

3.6

6.1

KU 1 15129 <3

Nicaragua

55.4

23.1

19.7

12.7

7.2

11.2

3.5

5.8

KU 115132 c?

Nicaragua

54.7

23.1

20.5

13.1

7.4

11.4

3.5

6.1

* Measurements as given by Gardner and Patton (1972).
** Measurements as given by Starrett (1969).

KARYOLOGY

Robert J. Baker

This chapter is in memory of Dr. Claude M. Ward, who introduced me to
the world of bats and whose premature death robbed me of a good friend
and the world of a dedicated educator.

The systematics of the New World leaf-nosed bats are based primarily on
classical morphological features such as shoulder articulation, dentition, and
other cranial features. The available fossil record is inadequate and probably
will always be too poor to determine much about the evolutionary relationships
of subfamilies and genera (Smith, 1976). As an adjunct to the data based on
classical morphological features, data from chromosomal and electrophoretic
studies are being generated (see also Straney et al., this volume). Hopefully, a
synthesis of the data from these and other works will result in a reasonably
complete understanding of the systematics and genetic strategies of members of
the family Phyllostomatidae. Data derived from bat chromosomes also serve to
verify, refute or modify the proposed models of chromosomal evolution (Wilson
et al., 1975; Bush, 1975).

In 1966 when I first began working with the chromosomes of this family, I
assumed that chromosomal divergence in the standard karyotypes of species,
genera, subfamilies, and the like generally would reflect their taxonomic status
and the evolutionary time that any two lineages had been separated. However, some
taxa (for instance, Glossophaga and Erophylla ) that obviously have been sepa¬
rated long enough to evolve morphological distinctness deserving of generic and
subfamilial status had indistinguishable karyotypes, whereas other species (such
as Uroderma bilobatum and Choeroniscus intermedius, see also Rhogeessa,
Bickham and Baker, 1977) contained considerable intraspecific chromosomal
divergence. If evolutionary relationships were based solely on standard karyo¬
typic data, one would produce a considerably different classification than that
currently derived from classical osteological and exomorphological studies.
Therefore, I began to question the value of chromosomal divergence as a
taxonomic indicator. 1 presently am opposed to placing too much emphasis on
degree of gross karyotypic divergence as a justification for taxonomic status
(with the possible exception of specific distinctness). Of course, the longer two
lineages have been separated, the more probable it is that events have occurred
that result in karyotypic divergence. However, karyotypic changes become
established in a species at such irregular intervals that one cannot depend on the
rate of their establishment to indicate taxonomic position.

John Bickham and I are preparing a manuscript in which we propose that the
rate and magnitude of chromosomal change is primarily a function of the degree
to which the karyotype is adaptive to the adaptive zone occupied by the organism.

107

108

SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

If this model proves accurate then, at times, organisms would undergo relatively
rapid chromosomal evolution and at other times there would be long periods of
reduced rates of chromosomal change.

The fact that karyotypic changes do not evolve at a constant rate is not too
startling if one realizes it is a well documented fact that morphological features
also evolve at different rates. In a given taxon, some features can become highly
derived from the ancestral condition, whereas others remain indistinguishable
from the primitive condition. Meanwhile, in a closely related taxon, a different
suite of characters can become derived whereas all other characters remain near
the primitive. If greater emphasis is placed on the derived characters, the sys-
tematics would result in greater taxonomic distance than if the classification were
based only on the characteristics that remained in the primitive condition. A
similar case might be made for the degree of morphological divergence — it does
not necessarily reflect the evolutionary history. Certainly, parallelism and con¬
vergence can result in incorrect “lumping,” and yet, emphasis on most rapidly
evolving features may result in oversplitting. However, the fossil record reveals
that generally there is agreement between total morphological divergence and
evolutionary history. In light of data from the fossil record, I believe that in the
majority of cases an overview of classical morphological data gives a more
reasonable and accurate reflection of the evolutionary history than does degree
of chromosomal divergence.

On the other hand, there are cases where karyotypic data can be more valuable
than general morphology. To a much greater extent than general morphological
information, G-band chromosomal data are applicable to the cladistic methodol¬
ogies of Hennig (1966). The likelihood of extensive convergence of G-banding
patterns is sufficiently low to warrant placing considerable confidence in the
data. The typical mammalian genome is arranged in such a manner that there are
enough chromosomal arms (linkage groups) to provide an adequate number of
data points to determine the relationships within complex taxa. Additionally,
G-band chromosomal characteristics are independent of exomorphological,
cranial, or osteological features and, therefore, serve as an independent data
source. A synthesis of findings from all of the aforementioned, plus those of a
biochemical nature (such as electrophoretic, immunological, and DNA hy¬
bridization), should give the most accurate interpretation of the phylogeny and
systematics of a taxon. Also, data from these three sources (general morphology,
karyology, and biochemical) will be necessary to understand the evolutionary
strategy of major taxa.

Of the 137 phyllostomatid species recognized by Jones and Carter (1976),
basic karyotypic data are available for 105 (Table 1). In addition, Gardner (1977)
reported karyotypic data for two additional taxa, Artibeus fuliginosus and A.
planirostris , which were not recognized by Jones and Carter (1976). Rep¬
resentative standard karyotypes for 60 species are presented in Plates 1 through
60, which follow the literature cited. I have attempted to illustrate the major
chromosomal complements found in the Phyllostomatidae. Plates are arranged
alphabetically by generic and species names within subfamilies: Phyllostomatinae,

BIOLOGY OF THE PHYLLOSTOMATIDAE

109

Table 1. — Chromosomal data for phyllostomatid hats. Subfamilies are arranged in the
order followed by Jones and Carter (1976). Genera and species are in alphabetical order.
Symbols are 2 n, diploid number; FN, Fundamental Number; M, metacentric; SM, sub-
metacentric; ST, subtelocentric; A, acrocentric. Two species names not recognized by Jones
and Carter (1976) are identified by an asterisk. Mesophylla is recognized as distinct from

Ectophylla.

Number of

Taxon

Y2 Authority

specimens

Phyllostomatinae

Chrotopterus auritus

Yonenaga, 1968;

Yonenaga et al., 1969

Lonchorhina aurita

Baker and Hsu, 1970

Baker, 1973

Lonchorhina orinocensis

No information

Macro phyll um macro phyllum

No information

Macrotus californicus

Kniazeff et al., 1967

Nelson-Rees et al., 1968

Davis and Baker, 1974

155

Greenbaum and Baker, 1976

100

Macrouts waierhousii

Baker, 1967; Hsu et al., 1968

Nelson-Rees et al., 1968

Davis and Baker, 1974

Nagorsen and Peterson, 1975

Greenbaum and Baker, 1976

118

Patton, 1976

Micronycteris behni

No information

Micronycteris brachyotis

Patton, 1976

Micronycteris daviesi

No information

Micronycteris hirsuta

Baker, 1973

Baker et al., 1973

Micronycteris megalot is

Baker, 1967; Hsu et al., 1968

Patton, 1976

Micronycteris minuta

Baker, 1973

Patton, 1976

Micronycteris nicefori

Baker and Hsu, 1970

Patton, 1976

Micronycteris pits ilia

No information

Micronycteris schmidtorum

Baker, 1973

Micronycteris sylvestris

No information

Mimon bennettii

No information

Million cozumelae

Patton, 1976

Mimon crenulatum

Baker and Hsu, 1970

Baker et al., 1 972 /z

Hsu and Benirschke, 1974

Gardner, 1977

Patton, 1976

Mimon koepckeae

Gardner, 1977

Pbylloderma stenops

Baker and Hsu, 1970

Baker, 1973

Phyllostom us d i scalar

Baker, 1967; Hsu et al., 1968

Yonenaga, 1968

Kiblisky, 1969

Yonenaga et al., 1969

Baker and Hsu, 1970

Baker, 1970

Patton, 1976

Phyllostomus elongatus

Baker, 1973

Phyllostomus hastatus

Yonenaga, 1968

Yonenaga et al., 1969

Kiblisky, 1969

Baker and Hsu, 1970

Patton, 1976

110

SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

Table 1. — Continued.

Phyllostomus lutifolius

No information

Tonatia bidens

Baker and Hsu, 1970

Patton, 1976

Tonatia brasiliensis

Gardner, 1977

Tonatia carrikeri

Gardner, 1977

Tonatia minuta

Baker and Hsu, 1970

Baker, 1973

Patton, 1976

Tonatia silvicola

Gardner, 1977

Tonatia venezuelae

This paper

Trachops cirrhosus

Baker, 1967; Hsu et al., 1968

Vampyrum spectrum

Baker and Hsu, 1970

Baker, 1973

Glossophaginae

Anoura brevirostrum

No information

Anoura caudifer

Yonenaga, 1968

Baker, 1973

Anoura cultrata

This paper

Anoura geoffroyi

Baker, 1967; Hsu et al., 1968

Baker and Hsu, 1970

Pathak and Stock, 1974

Anoura werckleae

No information

Choeroniscus godmani

A Baker, 1967

Hsu et al., 1968

A Baker, 1970a

Patton and Gardner, 1971

This paper

Choeroniscus inca

No information

Choeroniscus intermedius

Baker, 1970a

Baker, 1973

Pathak and Stock, 1974

Stock, 1975

Choeroniscus minor

No information

Choeroniscus periosus

No information

Choeronycteris mexicana

24-26

Baker, 1967; Hsu et al., 1968

Baker, 1973

Glossophaga ulticola

Baker, 1967

Glossophaga commissarisi

Baker, 1967; Hsu et al., 1968

Glossophaga longirostris

This paper

Glossophaga soricina

Baker, 1967; Hsu et al., 1968

Baker and Hsu, 1970

Baker, 1970a

Hylonycteris underwoodi

Baker, 1973

Leptonycteris curasoae

information

Leptonycteris sanborni

Baker, 1967; Hsu et al., 1968

Leptonycteris nivalis

Baker, 1973

Lichonycteris degener

No information

Lichonycteris obscura

Baker, 1973 (data incorrect)

This paper

Lionycteris spurred i

This paper

Lonchophylla concava

No information

Lonchophylla hesperia

No information

Lonchophylla mordax

No information

Lonchophylla robusta

Baker, 1973

Lonchophylla thomasi

Baker, 1973

Gardner, 1977

Monophyllus plethodon

This paper

Monophyllus redmani

Baker and Lopez, 19706

Musonycteris harrisoni

No information

Platalina genovensium

No information

Scleronycteris ega

No information

BIOLOGY OF THE PHYLLOSTOMATIDAE

Ill

Table 1. — Continued.

Carolliinae

CaroIIia brevicauda

20-21

Patton and Gardner, 1971

20-21

Stock, 1975

CaroIIia casianea

20-21

Baker and Bleier, 1971

Patton and Gardner, 1971 (Peru)

20-21

Patton and Gardner, 1971

(Costa Rica)

Hsu and Bernischke, 1973

20-21

Pathak and Stock, 1974

Stock, 1975

Hsu et al., 1975

CaroIIia perspicillata

20-21

Baker, 1967

Hsu et al., 1968

20-21

Yonenaga, 1968

20-21

Kiblisky, 1969

20-21

Yonenaga et al., 1969

Baker, 1970 a, 19706

20-21

Baker and Hsu, 1970

20-21

Baker and Bleier, 1971

20-21

Patton and Gardner, 1971

20-21

Pathak and Stock, 1974

20-21

Hsu et al., 1975

20-21

Stock, 1975

CaroIIia subrufa

20-21

Baker, 1967

20-21

Hsu et al., 1968

Baker, 1970a, 19706

20-21

Baker and Bleier, 1971

Rhinophylla alethina

No information

RJi inophylla fischerae

Baker and Bleier, 1971

RJi inophylla pumilio

Baker and Bleier, 1971

Hsu and Benirschke, 1973

Stenoderminae

Ametrida centurio

30-31

Baker and Hsu, 1970

Ardops nichollsi

30-31

Greenbaum et al., 1975

Ariteus flavescens

30-31

Greenbaum et al., 1975

Artibeus aztecus

30-31

Baker, 1973

Artibeus cinereus

30-31

Baker and Hsu, 1970

Arlibeus concolor

No information

Arlibeus ful ig ino su s*

30-31

Gardner, 1977

Artibeus glaucus

30-31

Gardner, 1977

Artibeus hirsutus

30-31

Baker, 1973

Artibeus inopinatus

30-31

This paper

Artibeus jamaicensis

30-31

Baker, 1967

30-31

Hsu et al., 1968

30-31

Kiblisky, 1969

30-31

Baker and Hsu, 1970

30-31

Baker and Lopez, 19706

Artibeus lituratus

30-31

Baker, 1967

30-31

Hsu et al., 1968

30-31

Yonenaga, 1968

30-31

Becak et al., 1969

30-31

Kiblisky, 1969

30-31

Yonenaga et al., 1969

30-31

Baker and Hsu, 1970

Pathak and Stock, 1974

Artibeus phaeotis

Baker, 1967

Hsu et al., 1968

Artibeus planirostris*

30-31

Gardner, 1977

Artibeus toltecus

30-31

Baker, 1967

30-31

Hsu et al., 1968

Artibeus watsoni

Baker, 1973

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SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

Table 1. — Continued.

Centurio sen ex

Baker, 1967; Hsu et al., 1968

Baker and Hsu, 1970

Chiroderma doriae

No information

Chiroderma im provision

Baker and Genoways, 1976

Chiroderma salvini

Baker, 1973

Chiroderma trinit atom

Baker and Hsu, 1970

Baker and Genoways, 1976

Gardner, 1977

Chiroderma villosum

Baker, 1967; Hsu et al., 1968

Baker and Hsu, 1970

Pathak and Stock, 1974

Gardner, 1977

Ectophylla alba

Greenbaum et al., 1975

This paper

Enchisthenes hartii

Baker, 1967; Hsu et al., 1968

30-31

Baker and Hsu, 1970

Me sophy lla macconnelli

21-22

Baker and Hsu, 1970

21-22

Hsu and Benirschke, 1971

21-22

Baker et al., 1973

Phyllops falcatus

No information

Phyllops haitiensis

30-31

Greenbaum et al., 1975

Nagorsen and Peterson, 1975

Pyyoderma bilab iatum

No information

Sphaeronvcteris toxophyllum

Baker, 1973

Stenoderma rufum

30-31

Baker and Lopez, 1 970/z

30-31

Genoways and Baker, 1972

Sturnira aratathomasi

No information

Sturnira bidens

Gardner and O’Neill, 1969

Sturnira erythromos

Gardner and O’Neill, 1969

Sturnira lilium

Baker, 1967; Hsu et al., 1968

Kiblisky, 1969

Baker and Hsu, 1970

Sturnira ludovici

Baker, 1967; Hsu et al., 1968

Kiblisky, 1969

Sturnira mayna

Gardner, 1977

Sturnira mordax

Baker, 1973

Sturnira nana

Gardner, 1977

Sturnira thomasi

This paper

Sturnira tildae

Baker and Hsu, 1970

Uroderma b Hob atom

Baker, 1967; Hsu et al., 1968

Baker and Hsu, 1970

Baker and Lopez, 1970a

Hsu and Benirschke, 1971

44 or 43

Baker and McDaniel, 1972

122

Baker et al., 1972

total of 144

44 or 43

Baker et al., 1972

Baker et al., 1975

Uroderma magnirostrum

Baker and Lopez, 1970a

Hsu and Benirschke, 1971

Vampyressa bidens

Gardner, 1977

Vampyressa brocki

Baker and Genoways, 1972

Baker et al., 1973

Gardner, 1977

BIOLOGY OF THE PHYLLOSTOMATIDAE

113

Table 1. — Continued.

Vampyressa melissa

Gardner, 1977

Vampyressa nymphuea

Baker, 1973

Baker et al. , 1973

Gardner, 1977

Vampyressa pusilla

23-24

Baker, 1973

Baker et al., 1973

24-23

Baker et al., 1973

22-23

ST-A

Gardner, 1977

Vam pyrod es caraceiol i

Baker and Hsu, 1970

Baker, 1973

Vampyrops aurarius

information

Vampyrops brachycephalus

Baker, 1973

Vampyrops dorsalis

Baker, 1973

Vampyrops helleri

Baker, 1967; Hsu et al., 1968

Baker and Hsu, 1970

Vampyrops infuscus

Gardner, 1977

Vampyrops lineatus

information

Vampyrops nigellus

Gardner, 1977

Vampyrops recifinus

information

Vampyrops vittatus

Baker, 1973

Brachyphyllinae

Brachyphylla cavernarum

Baker and Lopez, 19706

Brachyphylla nana

This paper

Brachyphylla pumila

Nagorsen and Peterson, 1975

Erophylla sezekorni

Baker and Lopez, 19706

Nagorsen and Peterson, 1975

Phyllonycteris aphylla

This paper

Phyllonycteris major

information

Phyllonycteris ohtusa

Nagorsen and Peterson, 1975

This paper

Phyllonycteris poeyi

information

Desmodontinae

Desmodus rotundas

SM A

& ST

Forman et al., 1968

Yonenaga et al., 1969

Cadena and Baker, 1976

Diaemus younyii

Forman et al., 1968

Cadena and Baker, 1976

Diphylla ecaudata

Baker, 1973 (data incorrect)

Cadena and Baker, 1976

Gardner, 1977

Plates 1 to 17; Glossophaginae, 18 to 29; Carolliinae, 30 to 32; Stenoderminae,
33 to 52; Phyllonycterinae, 53 to 57; Desmodontinae, 58 to 60.

Determination of Primitive Karyotype

One very important point of information relative to determining evolutionary
events and their systematic implications is an understanding of the primitive
versus the derived condition. Because there is no fossil record for karyotypes,
primitive cytogenetic aspects are difficult to ascertain.

Prior to the availability of G-band data, two theories were developed as to the
diploid and fundamental characteristics of the primitive karyotype for the
family Phyllostomatidae. Baker (1967, 1973) proposed that the primitive karyo¬
type for the Phyllostomatidae consisted of a diploid number (2 n) of 30 or 32,
with a fundamental number (FN) of 56 to 60. This theory was based on the

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SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

widespread occurrence of the 2/7 = 30 or 32, FN = 56-60 karyotype among species
from the different subfamilies; the alternative explanation was to assume the
condition arose through convergent evolution. Gardner (1977), on the other hand,
proposed that the primitive karyotype was 2/7 = 36 to 40 with an FN near the
minimum for this diploid number, 38 or slightly higher. The significant difference
between the two theories centers around the types of chromosomal rearrange¬
ments required to derive the karyotypes found in extant species. The 2/7 = 30 or 32,
FN = 60, would require terminalization of centromeres by pericentric inversion or
centric transpositions in addition to translocations (especially centric fusions) as
theprimary rearrangements, whereasthe2/7 = 36-40, FN = 38,wouldrequirecentral-
ization of centromeres (by pericentric inversion or centric transpositions) in addi¬
tion to some fusions.

It is of interest to note that when Gardner (1977:314-315) interpreted phy¬
logenetic relationships within the family based on chromosomal evolution from
a primitive karyotypic condition of a higher diploid number (about 40) and a
lower fundamental number (about 38), his three “major deviations from more
classical portrayals” were essentially those proposed earlier based on a primitive
2/j = 32, FN = 60, karyotype. Relative to Gardner’s case 1, Baker and Lopez
(19706:471) pointed out the “possibility of a close phylogenetic relationship”
of the phyllonycterine genera to Monophyllus. In Gardner’s case 2, Baker (1967:
423), basing his remarks on karyotypes, not only suggested that Sturnira “must
have evolved from the Stenoderminae complex,” he also regarded the two sub¬
families as synonymous, which is the systematic relationship followed by Jones
and Carter (1976:20). In case 3, Greenbaum etal. (1975) suggested the recognition
of Mesophylla as generically distinct from Ectophylla. The point is that even
though a 2/7 = 40, FN = 38, primitive karyotype theory might be a viable alternative
to the 2/7 = 32, FN = 60, theory in several examples, the systematic implications
of the chromosomal data are the same.

With data from G-bands, it became possible to identify homologous segments
between variant karyotypes even at the subfamilial level (Mascarello et al., 1974),
and G-band studies became the means for testing these two theories. It could be
predicted that if the theory of 2/7 = 30 or 32, FN = 56-60, were true, there should be
considerable homology of banding patterns between the two arms of the supposed
homologous elements of the 2/7 = 30 or 32, FN =56 or 60 karyotypes within the
family, and although some elements in each karyotype may have been rear¬
ranged, the same pairs should not always be affected. On the other hand, if the
2/7 = 40, FN = 38 (Gardner, 1977) karyotype proved primitive, G-banding pat¬
terns of biarmed elements of the 2/7=32 karyotypes from separate subfamilies
should show little homology between the subfamilies. Therefore, G-banding
homology among these karyotypes with lower fundamental numbers from
different subfamilies would be strong proof in favor of Gardner’s theory.

Patton (1976) examined G-banded chromosomes of five genera (involving
10 species) of the subfamily Phyllostomatinae as well as one species from the
families Mormoopidae ( Pteronotus parnellii) and Noctilionidae ( Noctilio
albiventris). His results indicated that the FN = 60 was primitive for the Phyl-

BIOLOGY OF THE PHYLLOSTOMATIDAE

115

lostomatinae as well as for the Mormoopidae and Noctilionidae. Macrotus (as
well as several other species of the Phyllostomatinae), Pteronotus, and Noctilio
all have 30 pairs of autosomal arms. When the G-banded karyotypes of these three
genera are compared, the thirty homologous arms found in the karyotype of each
genus have a distinguishable counterpart in the karyotypes of the other two genera.
The most logical interpretation of these data is that the number of autosomal
arms in the karyotype of the common ancestor of Macrotus, Pteronotus, and
Noctilio was 30 pairs (FN = 60), which have retained their respective G-banding
patterns since their separation from a common ancestor. The alternative ex¬
planation, that the G-band similarity between the representatives of these three
families is the result of the evolution of convergent G-banding patterns in the exact
same number of pairs (30) of autosomal arms, is less plausible (Patton, 1976).
Additionally, data from the G-banded karyotypes of other taxa thus far studied
(by Patton, 1976, and unpublished data including representatives of the Des-
modontinae, Glossophaginae, and Stenoderminae) support the conclusion that
the FN = 60 was primitive for the Phyllostomatidae. Derivation of the various
karyotypes of the taxa studied from any of the karyotypes with the more aberrant
fundamental numbers (such as Tonatia bidens FN = 20 or Micronycteris megalotis
FN = 68) would require many convergent chromosomal rearrangements in order
to avoid concluding that Macrotus was more closely related to the mormoopids
and noctilionids than to the other phyllostomatids.

The primitive diploid number for the Phyllostomatidae was believed to be
2u = 46 (Patton, 1976). The following discussion, modified from Patton’s thesis,
points out the reasons for this conclusion.

A diploid number of 46 (with 16 biarmed autosomes, 28 acrocentric autosomes,
plus two sex elements) is most probably like the primitive condition (Patton,
1976). Essentially, this is the karyotype of Macrotus waterhousii (Fig. 1). Data
supporting this conclusion are the eight pairs of biarmed elements found in the
karyotype of Macrotus that have corresponding biarmed elements in the karyo¬
type of Noctilio. Seven of these eight pairs are present also in Pteronotus, Tonatia
minuta, Mimon crenulatum, Phyllostomus discolor, and Phyllostomus hastatus.
The majority of these eight pairs are identifiable in most of the karyotypes of other
phyllostomatine species studied. Therefore, it is likely that these eight biarmed
pairs were primitive for the phyllostomatoid karyotype. In addition to the eight
biarmed pairs described as common for Noctilio, Pteronotus, and Macrotus,
the karyotypes of most species examined include several other biarmed elements,
the banding patterns of which suggest independent fusions of acrocentric ele¬
ments.

An alternative hypothesis would be to propose a noctilionid-mormoopid-
like karyotype as primitive. Such a primitive karyotype would, however, require
additional events — fission would have to precede several independent fusions.
As demonstrated by Mascarello et al. (1974) for rodents, the establishment of
fission rearrangements is quite rare, whereas Robertsonian fusion products are
the most common type of euchromatic variation observed between closely related
taxa. Therefore, a fission-fusion mode not only would require additional events,
it would also be less probable from a cytogenetic standpoint.

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SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

BIOLOGY OF THE PHYLLOSTOMATIDAE

117

In the following paragraphs on the evolutionary relationships indicated by
karyotypic data, it is assumed that the primitive karyotype for the Phyllosto-
matoidea was 2 n = 46, FN = 60 and with a morphology similar to that of Macrotus
waterhousii. The discussion is essentially limited to G-band data because all
other would be too speculative and G-band studies of most subfamilies will un¬
doubtedly appear shortly. Proposed karyotypic relationships for some phyl-
lostomatid taxa, based on standard karyotypes, are presented in Baker (1973),
Greenbaum et al. (1975), and Gardner (1977).

Systematic Affinities
Familial Affinities

The first instance where the members of the Mormoopidae, Noctilionidae,
and Phyllostomatidae were classified together, but distinct from all other bats, was
Winge (1941). Smith (1972) drew similar conclusions — the Phyllostomatoidea
consisted of the families Mormoopidae, Noctilionidae, and Phyllostomatidae.
G-band chromosomal data strongly support this classification and suggest that
Pteronotus and Noctilio shared a common evolutionary ancestor in which five
Robertsonian fusions became established (Patton, 1976). These data indicate that
the Noctilionidae and Mormoopidae are more closely related to each other than
either is to the Phyllostomatidae. Smith (1972) came to the same conclusions
based on morphological data. The most recent common ancestor of Pteronotus
and Noctilio probably had a 2n = 36 condition.

The degree of chromosomal divergence distinguishing Noctilio from Pteronotus
is the least known to separate two mammalian families. Before someone
jumps to the conclusion that the families Mormoopidae and Noctilionidae are
confamilial, I would point out that prior to the study by Patton (1976), there had
been considerable disagreement as to the evolutionary affinities of both families
(Smith, 1972). In fact, there would be little agreement as to what family
Noctilio should be placed in if it were not awarded familial status. Some clas¬
sifications have included the mormoopids as a subfamily of the Phyllostomatidae
(Miller, 1907; see also the review by Smith, 1972), and the chromosomal data
merely indicate that if all lineages evolved from the most recent ancestor of the
mormoopid-phyllostomatid line are to be included in the family Phyllostomatidae,
then the Noctilionidae should also be reduced to a subfamily.

Chromosomal data from Noctilio and Mormoops further document the fact
that karyotypic change is not a requirement for the evolution of a magnitude of
morphological difference worth of recognition of a higher taxonomic category
(Patton, 1976). It has been suggested by Wilson et al. (1975) that the large degree
of morphological evolution in mammals is due to regulator gene alterations by

Fig. 1. — A composite of two G-banded karyotypes of Macrotus waterhousii prepared for
use as standard reference in describing chromosomal events in the family Phyllostomatidae
as proposed by Patton (1976). Both homologs from the two spreads are presented in order
that minor variation can be observed. Figure courtesy of Rebecca A. Bass.

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SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

chromosomal mutations. However, few changes in primitive linkage groups are
often characteristic of rather divergent taxa of bats and rodents (see also Mascarello
et al., 1974; Stock and Hsu, 1973), which leads one to conclude that at least some
mammalian taxa have evolved primarily via point mutations and have conserved
the primitive gene arrangements. A similar conclusion can be drawn for reptiles,
based on chromosomal banding analysis of turtles (Stock, 1972; Bickham and
Baker, 1976), and for birds (Stock et al., 1974).

Subfamilial Affinities

There has been only one paper in which G and C-band data have been used to
relate species from different subfamilies (Stock, 1975) and this work found
essentially no G-band autosomal homologies between Carollia (subfamily
Carolliinae) and Choeroniscus (subfamily Glossophaginae). From standard
karyotypes, a close relationship between these two genera had been proposed
(Baker, 1967). Stock noted that the X elements were essentially the same between
the two genera but concluded that there were no data supporting a close common
ancestor for Carollia and Choeroniscus and suggested that these two genera
be placed in separate subfamilies. I have little doubt that a complete G-band study
of the genera within all subfamilies will reveal the evolutionary relationships of
most subfamilies. G and C-band studies on the Brachyphyllinae and Des-
modontinae (by Rebecca A. Bass) and Stenoderminae (by Anette Johnson) are
presently being conducted in my laboratory.

Relationships Within Subfamilies

Phyllostomatinae. — Relationships within the subfamily Phyllostomatinae
were studied by Patton (1976), but his results were somewhat incomplete because
only five of 11 genera (involving 10 of 33 species) were studied; these were
arranged into three groups: 1) Micronycteris, 2) Tonatia, Mimon , and Phyl-
lostomus, and 3) Macrotus.

The Macrotus group could have evolved from any lineage just as long as it
became separated from the other stocks prior to the establishment of any chro¬
mosomal rearrangements. The karyotype of Macrotus waterhousii has been
proposed as like that which was primitive for the family (see above). The karyo¬
type of M. californicus (2n — 40, FN = 60) would then be derived by three centric
fusions (Davis and Baker, 1974), which would have been independent events
from fusions established in the other two lines discussed below.

Patton’s (1976) Micronycteris group is characterized by the sharing of two
derived arrangements. One is a terminal translocation of chromosome 13 onto
pair 26/25 and the other is a Robertsonian fusion between acrocentric pairs
1 8 and 2 1 . All other rearrangements within the Micronycteris cluster appear to
have been achieved through independent events within the three subgenera
( Trinycteris, Micronycteris, and Lampronycteris) studied by Patton. The
hypothesized primitive karyotype for the subgenera Trinycteris and Micronycteris
would be 2n = 42, FN = 58. The fact that these species ( minuta , nicefori, and

BIOLOGY OF THE PHYLLOSTOMATIDAE

119

brachyotis) representing three subgenera, can be chromosomally related, strongly
reinforces the natural status of at least portions of the genus. I have heard several
people propose that this genus is a catchall with several species of questionable
generic affinity. One species that cannot be related chromosomally to the other
representatives of the genus thus far studied is M. megalotis, the type species of
the genus.

The Tonatia- Mimon- Phyllostomus group is identified by five shared derived
chromosomal events: four Robertsonian fusions (22/3, 8/9, 17/12, 29/27) and
one inversion (4/5). These chromosomal characteristics are shared by Tonatia
minuta, Phyllostomus discolor, P. hastatus, and Mimon crenulatum. The ancestral
karyotype for the common ancestor probably had a 2 n = 38, FN = 60. A common
ancestor for Phyllostomus hastatus, P. discolor, and Mimon crenulatum is
suggested by three shared fusion events (18/13, 1 4a/2 1 , 30/28). This would
mean that the common ancestor for these three species had a karyotype with a
2n = 32 or 34. As Robertsonian fusion products occurring independently in forms
containing only two acrocentric linkage groups could only lead to the same
fusion product, a 2/? = 34 divergence cannot be totally discounted (Patton, 1976).
The possibility of a 2/7=34 divergence is strengthened by Mimon cozumelae
having a 2n = 34, FN = 60 karyotype.

The karyotype of Tonatia bidens (2/i = 16) is so derived from the Macrotus and
Tonatia minuta karyotypes that it could not be related to those of other members
of the subfamily. Again, this points out a case where most chromosome divergence
has been limited to changes that can be traced by homology of G-bands, but
during the evolution of T. bidens numerous chromosomal changes became
established. If systematic position were based solely on chromosomal divergence,
one would have to recognize T. bidens as generically distinct from other phyl-
lostomatines possibly with subfamilial status, a ridiculous conclusion in my
opinion.

Glossophaginae. — There are no G-band studies on the generic relationships
within the Glossophaginae. The only published G-banded karyotype is of
Choeroniscus intermedius (Stock, 1975), which is discussed above under sub-
familial relationships.

Gardner (1977) presented a phylogeny of the Glossophaginae based on
standard karyotypes and in most cases has followed the most parsimonious routes.
However, I cannot accept that the similar karyotypes of Choeronycteris and
Hylonycteris are the result of parallelism. This 2/7=16 karyotype is undoubtedly
derived, and I feel that it is explained best as being due to their common ancestor
having a diploid number of 16. G-banding should be valuable in settling this
difference in interpretation.

Carolliinae. — G-band data (Stock, 1975) have been published for one (Carollia,
three species studied) of the two genera of the Carolliinae. Carollia brevicauda
and C. perspicillata share two chromosomal features (an X-autosomal trans¬
location and similar heterochromatin patterns) that distinguish these two species
from at least some individuals of C. castanea. Pine (1972), in a study based on
classical morphological features, concluded that C. brevicauda and C. perspicillata

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SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

Fig. 2. — G-banded karyotype of Artibeus jamaicensis. Figure courtesy of M. Anette
Johnson.

were more closely related to each other than either is to C. castanea. Carollia
castanea has two chromosomal races that are based on the presence of the X-
autosomal translocation in specimens from Central America (Patton and Gardner,
1971) and Colombia (Baker and Bleier, 1 970) and the absence of this translocation
in Peruvian specimens (Patton and Gardner, 1971).

Patton and Gardner (1971) argued that the absence of the X-autosomal trans¬
location in some populations of C. castanea is the result of the primitive condition
being maintained. This would best explain the current taxonomic distribution
of the X-autosomal translocation if the ancestor of all Carollia species was
polymorphic for this translocation. In C. perspicillata , C. subrufa, and C. brevi-
cauda, this translocation became fixed and characteristic of the species, whereas

BIOLOGY OF THE PHYLLOSTOMATIDAE

121

Chromosome groups
B C D E F

* %

n» *■*

«•«

* M

<u> £&

2N=44

c a

£ J

A a

«•

fittt

- _■

»u

»er

ii ii m i« si

tZ s;

r»

*** **

for each karyotype: 2 n of 44 is A A BB
40 is Aa Bb cc ; and 2// = 39 is aa Bh cc.

Fig. 3. — G-banded karyotype and partial karyo¬
types of specimens of Uroderma bilobatum with
various diploid numbers. Top row. Complete
karyotype of a specimen with 2/7 = 44. Rows 2-4
are group A, B, and C chromosomes of specimens
with a 2/7 of 43, 42, 40, and 39, respectively. Chro¬
mosomes in groups A, B, and C account for all the
variation in diploid number. D chromosomes are
the two sex elements. E chromosomes are the large
biarmed elements and F chromosomes are the
acrocentric elements that are not involved in the
variation. Group A chromosomes appear as either
a small pair of biarmed elements plus a pair of
acrocentrics (designated as the A A morph), a
small biarmed element plus an acrocentric and a
subtelocentric element (Aa morph) or as two sub-
telocentric elements (aa). Group B chromosomes
appear as two pairs of acrocentric elements (BB),
a pair of acrocentrics unequal in size and a sub-
metacentric element (Bb) or two submetacentric
elements (bb). Group C appears as four acrocentric
elements (CC), three acrocentric elements (Cc) or
two acrocentric elements (cc). Genetic designation
CC; 2/7 = 43 is AA Bb CC: 2/7 = 42 is Aa BB Cc;2n —

in C. castanea some populations became fixed for both conditions. This explana¬
tion might be correct and I agree that it is the first choice; however, based on the
limited data now available, an alternative explanation cannot be ruled out. It
is possible that the absence of the X-autosomal translocation in some C. castanea
is due to a fission of these elements and represents a condition more derived than
that characteristic of C. perspicillata, C. subrufa, and C. brevicauda.

Stenoderminae. — G and C-band chromosomal data for Sturnira lilium,
Artibeus jamaicensis (Fig. 2), Enchisthenes harti, and Uroderma bilobatum
(2n = 44 cytotype, Fig. 3) are described by Baker et al. (1979). The G-band-
ing pattern for Artibeus and Sturnira revealed that the similarity in the gross
karyotypes reflected homology with only one autosomal change (a pericentric

122

SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

inversion) distinguishing their respective karyotypes. The karyotype of Enchist-
henes harti could be derived from the Artibeus karyotype by a reciprocal trans¬
location involving two autosomes. This translocation changes two submetacentric
chromosomes in Artibeus to two subtelocentric chromosomes in Enchisthenes.

It was more difficult to show homology between Artibeus and the Uroderma
2/i = 44 karyotype. Two pairs of Artibeus autosomes were homologous with
elements in Uroderma ; the other 1 2 pairs of Artibeus (85 per cent of the autosomal
pairs) autosomes required rearrangement to derive the Uroderma karyotype.
For some chromosomal segments, homologous elements could not be determined
between the two karyotypes.

Artibeus has five (Baker et al., 1979) pairs of biarmed chromosomes that are
homologous with pairs found in Macrotus. The biarmed pairs homologous between
Artibeus and Macrotus are thought to be primitive for the family (Patton, 1976).
Uroderma and Macrotus have no homologous biarmed chromosomes; however,
they share acrocentric chromosomal homologies.

Only two pairs of chromosomes (both biarmed) were shared by all four
stenodermine genera studied. These two pairs are not found in any of the other
subfamilies studied (Phyllostomatinae, Patton, 1976; Glossophaginae and
Carolliinae, Stock, 1975) and are, therefore, potentially valuable indicators of
a common ancestry for these and other stenodermine genera. Such marker elements
should prove valuable in determing if Brachyphylla has evolutionary affinities
with the Stenoderminae. The G-band data for Sturnira are interpreted as
additional documentation that the genus Sturnira has a common ancestry with
the Stenoderminae and should be recognized as a member of that subfamily
(Baker et al., 1978). G-banded karyotypes for Uroderma bilobatum are shown in
Fig. 3 and are discussed below in the following section.

Desmodontinae and Brachyphyllinae. — No G-banded karyotypes have been
published for the subfamilies Desmodontinae and Phyllonycteririae.

[Note added in galley. — G-band data are now available for several additional
species so that the following important conclusions can be drawn. The glos-
sophagine genera Glossophaga and Monophyllus have identical G-band chromo¬
somal homologies with species of Phyllonycteris, Erophylla, and Brachyphylla.
These data indicate that these five genera shared a common ancestor after sep¬
arating from the other subfamilial lineages (with the possible exception of the
Carolliinae) and that Brachyphylla is properly associated with the genera
Phyllonycteris and Erophylla (Baker and Bass, 1979), as was suggested by Silva
Taboada and Pine (1969). However, when the genus Brachyphylla is placed in
this subfamily, Brachyphyllina Gray, 1866, becomes the oldest available
family-group name for the subfamily (Phyllonycterinae was first proposed by
Miller, 1907). The proper name of the subfamily then would be Brachyphyllinae.

In a manuscript recently submitted for publication by Rebecca A. Bass and the
author, it was shown that the vampire bats (Desmodontinae) shared a common
ancestry with the glossophagines and brachyphyllines, after this lineage separated
from the remainder of the family. ]

BIOLOGY OF THE PHYLLOSTOMATIDAE

123

Variation within Species

From the standpoint of population biology, this is the level where chromosomal
variation can be used to make the most significant studies. The role of various
types of mechanisms of chromosomal evolution can be studied as an isolating
mechanism, effective means of producing heterosis, supergenes, etc. Variation at
this level can be due to populational polymorphisms or chromosomal races.

Polymorph isms

A widely distributed polymorphism has been described (Baker et al., 1912b)
for Mimon crenulatum. The polymorphism was found in samples from Peru,
Trinidad, and Colombia and is believed to be restricted to the fifth largest pair of
autosomes. Three morphs of this chromosome were identified from each of the
three localities. For polymorphism to be maintained over such a wide geographic
range, it must offer a selective advantage to the species greater than the expense
of its maintenance.

Baker and Lopez, (1970a) demonstrated a polymorphism also for Uroderma
magnirostrum. Eleven of thirteen specimens examined from Colombia had a
diploid number of 36, whereas two had a diploid number of 35. Because the size
of the additional biarmed element was greater than a fusion between any two
acrocentrics, the polymorphic system may not be the result of a simple centric
fusion.

Other cases of chromosomal variation at a single locality are based on the
discovery of a single aberrant individual, which may represent a balanced poly¬
morphic system or variation that originated within that individual.

A centric fusion was reported in a female Mesophylla macconnelli from
Trinidad; nine other specimens from this locality did not possess the chromosome.
An Artibeus toltecus from San Luis Potosi, Mexico, had a 2n = 32 with what
appeared to be a trisomy for a small autosome and one other male from this locality
had a 2n = 31, which is normal for the species.

In a sample of 78 Uroderma bilobatum from near Choluteca, Honduras, one
individual had a 2n — 31, which resulted from a fusion of two acrocentrics
into a metacentric of the same general size range as the subtelocentric autosomes.
Chromosomal variation at this locality is common as a result of hybridization
between two cytotypes (see the discussion on Uroderma below); however, this
centric fusion is easily identifiable from those events that separate the two
cytotypes because the fusion product is a metacentric, and such an element has
not been observed in 332 other specimens of Uroderma bilobatum from Central
America.

Chromosomal Races

Chromosomal races are known for three species of phyllostomatid bats. What
originally was reported as chromosomal races in Macrotus waterhousii proved
to be specific differences characteristic of two species: M. waterhousii , with

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SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

2n — 46, and M. californicus, with 2n — 40 (Davis and Baker, 1974; Greenbaum
and Baker, 1976). Two races are known for Micronycteris hirsuta (Baker et al.,
1973). One is a 2n= 30 cytotype from Middle America characterized by a single
pair of submetacentric autosomes. Specimens from Trinidad, on the other hand,
have a karyotype with a 2n = 28, FN = 32 and show two pairs of submetacentric
autosomes and two less pairs of acrocentrics. The degree of divergence in cranial
and forearm measurements in the specimens karyotyped is too low to suggest
that the two chromosomal races represent distinct species (Baker et al., 1973).

Two races of Vampyressa pusilla were described by Baker et al. (1973). One
race has a 2n=18, FN = 20 with two pairs of submetacentric autosomes and
six acrocentric pairs. The X is a subtelocentric, and the Y is a small distinctly
biarmed element. This race is known from Honduras, Nicaragua, and Costa
Rica. The second race, found in Colombia, has a karyotype that consists of a
2n — 24 in females and a 2n = 23 in males, with an FN of 22. There are no sub¬
metacentric autosomes. To explain divergence between the two races requires at
least three events. Even though the magnitude of variation is greater than that
characteristic of most congeneric species of phyllostomatids, no exomorphological
or cranial differences were found that could distinguish the races (Baker et al.,
1973). Data from V. pusilla documents another case of discordant rates of
evolution between classical and karyotypic morphology.

Uroderma bilobatum, Peters’ tent-making bat, is the third species of phyl-
lostomatid bat known to have chromosomal races. The three chromosomal races
reported for this species have been the object of considerable study (Baker and
Lopez, 1970a; Baker et al., 1972a; Baker and McDaniel, 1972; Baker et al.,
1975); one zone of contact between two races has been located. Information on
the nature and dynamics of this zone could be valuable in understanding some
aspects of the speciation process.

Elucidation of the processes by which one species becomes transformed into
two or more is the key to understanding evolution. The genetic interactions
involved between two diverging populations within a species dictate the evolu¬
tionary future of these populations. Although several theories have been
postulated for such genetic interactions and their relationships to the process of
speciation, actual measurements of the interaction are difficult to make and
definitive data are lacking.

An important aspect involved in speciation is the chromosomal com¬
patibility between diverging populations. One proposed model of speciation
(stasipatric speciation by White, 1968) is based entirely on chromosomal di¬
vergence. The situation with JJroderma bilobatum (see details below) does not
exactly fit the stasipatric model put forth by White; however, Uroderma offers
a unique opportunity to examine the role of karyotypic diversity and the resulting
interaction between two interbreeding populations. A detailed understanding of
the mechanisms and events occurring at the contact zone between two chro-
mosomally characterized populations of Uroderma bilobatum is important
because we will be able to observe a stage of evolution that could result in the

BIOLOGY OF THE PHYLLOSTOMATIDAE

125

formation of two species. It could provide insight into how chromosomal changes
become fixed within a population.

The paucity of measurements on the genetic interactions resulting in speciation
(especially in mammals) can be attributed to both the difficulties in obtaining such
measurements and the inability of available techniques to identify appropriate
biological situations for study. In order to attempt to measure the genic interactions
that might produce speciation, it is first essential to locate a situation where popu¬
lations have diverged. In addition, it is necessary to be able to identify within
the population first generation crosses between the types (referred to as Fis,
although this does not imply specific status of the types) and backcross individ¬
uals.

Measurements of degree of exomorphological and cranial divergence have
proven inadequate for such studies. By the point in time when organisms are
sufficiently diverged to enable the recognition of Fi and backcross individuals by
these techniques, the stage at which the most significant interactions occur has
passed. Numerous studies can be cited to document this problem (see Lidicker,
1962, for a review of the problems of subspecific evolution in mammals). Eveji
when interpreted with the use of the most sophisticated multivariate techniques,
measurements of exomorphological and cranial features cannot identify with any
certainty the Fi and backcross individuals or measure genetic interaction (see
Baker et al. , 1 975). The extent to which gene flow has been reduced when alloptric
populations reestablish contact simply cannot be ascertained with any degree of
accuracy from measurements of exomorphological and cranial features.

In cases where two interacting populations are characterized by chromosomal
differences, Ft individuals will have a predictable karyotype unique from that of
both parental types. If the chromosomal differences are of sufficient magnitude,
the first generation backcross individuals will have karyotypes distinguishable
from the Fi and parental karyotypes. Such biological situations provide an ex¬
cellent case for detailed investigations into the genetic interactions of divergent
populations and the process of speciation.

It should be pointed out, however, that anytime karyotypes are used to identify
diverging populations, one is studying a special case because chromosomes are
involved and chromosomes could be the primary isolating mechanism. There are
many isolating mechanisms known, and it is possible that each represents a
special case. It is also probable that no single isolating mechanism is involved in
all cases of speciation. The aim of the detailed study of Uroderma in my laboratory
is to investigate the role of chromosomal divergence in the evolutionary process
as exemplified by these bats.

The classical systematics and distribution of U. bilobatum are as follows:
Uroderma bilobatum occurs at lower elevations from southern Mexico southward
through parts of tropical South America. Based on variations in external and
cranial measurements, karyology, and pelage color, six subspecies ( bilobatum ,
molaris, convexum, trinitatum, davisi, and thomasi ) are recognized (Davis,
1968; Baker and McDaniel, 1972). Extensive chromosomal investigations of the

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SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

Fig. 4. — Geographic distribution of samples of Uroderma bilobatum within contact zone
(see Table 2). Specific localities are 1) El Salvador: La Paz: 3.0 mi. NW La Herradura; 2)
Usulatan: 3.0 mi. E Usulatan; 3) Honduras: Valle: 9 mi. SE Nacaome; 4) Valle: 10.0 mi.
SSW Nacaome, 5) Choluteca: 10.2 mi. NW Choluteca, 6) Choluteca: 11.5 mi. SW Cholu-
teca; 7) Nicaragua: Chinandega: 3.5 mi. NW and 1.5 mi. S Chinandega. The 2/i = 44 parental
cytotype occurs at localities 1-3, both parental cytotypes are present at locality 4, and the
2/; = 38 parental cytotype occurs at localities 5-7.

Uroderma bilobatum complex have revealed chromosomal divergence greater
than that reported for any other species of bat (Baker, 1967, 1970a, 1970ft;
Hsu et al., 1968; Baker and Hsu, 1970; Capanna and Cibitelli, 1970).

Karyotypically, the U. bilobatum complex can be divided into the following
groups: 2n = 44, davisi (central Honduras north to southern Mexico; (Baker and
McDaniel, 1972); 2« = 38, including convexum (central Honduras south to
northern South America on the Pacific versant), and molar is (Mexico to Nicaragua
on Atlantic versant; as suggested by Davis, 1968); and 2n = 42, consisting of the
nominal subspecies trinitatum and bilobatum (South American mainland).
Uroderma ft. thomasi, which has not been karyotyped, is known from western
South America. Uroderma ft. convexum (2 n = 38) and U. ft. davisi (2n = 44) have
been found to form a contact zone over 200 kilometers in length (Fig. 4) that
extends from southern El Salvador, across the Pacific coast of Honduras and
northwestern Nicaragua (Baker et al., 1975).

BIOLOGY OF THE PHYLLOSTOMATIDAE

127

Conclusions concerning the nature of chromosomal variation in Uroderma
between the 2/i = 38 and 2/7 = 44 forms are based on G-band data (Fig. 3). The
diploid number at the contact zone in Central America ranges from 38 to 44,
with individuals of all intermediate diploid numbers being represented. North¬
west of the contact zone the diploid number is 44 and to the southeast is 38
(Fig. 4). Intermediate individuals are not known from outside of the zone of
contact. The differences between the two parental types (38 to 44) result from
three separate events, each involving a translocation or fission, depending on
direction of the event. The first change to be discussed (designated the A chro¬
mosomes) is shown in column A of Fig. 3. One morph (represented by a capital
A) has a small biarmed element and an acrocentric element; the other morph
(represented by a lower case a) has these two elements fused to form a subtelo-
centric chromosome. Where only a standard karyotype is available, the number
of large A' s in the karyotype will be reflected by the number of small biarmed
autosomes present in the complement.

The second event (designated the B chromosomes) involves a centric fusion-
fission in which morph B (column B, Fig. 3) appears as two acrocentrics (the
smallest acrocentric in the 2/7 = 44 karyotype and one of the medium-sized
acrocentric elements). Morph b is a subtelocentric element representing a fusion
of the two acrocentric elements in B. This variation can be recognized in a standard
karyotype because each b is reflected by a decrease of one in the diploid number
and an increase of one in the number of large biarmed elements, without effecting
a decrease in the number of small biarmed autosomes.

The third change (designated the C chromosomes), as shown in column C of
Fig. 3, is a terminal translocation in which a small acrocentric element is trans¬
located to the end of the long arm of the longest acrocentric element in the karyo¬
type. For each morph C, there will be two acrocentrics in the karyotype, whereas
each morph c is a single large acrocentric in which the segments homologous to
the two C acrocentrics are fused. Production of the c morph reduces the diploid
number by one and reduces the number of acrocentrics by one but does not alter
the number of biarmed elements (either small or large) in the karyotype.

Although the exact nature of these changes can be identified only by the G-band
patterns, the three changes produce distinct morphological differences in the
chromosomes that allows one to determine the chromosomal phenotype from a
standard karyotype for the A, B, and C chromosomes of any individual. Using
the ABC designation for the chromosomal variation enables the characterization
of all of the individuals involved in the contact zone. An animal with AABBCC
would be a 2n = 44 parental type and an animal with aabbcc would be a 2n — 38
parental type. Each capital letter in the phenotype will raise the diploid number
above 38 by one. For example, an animal with a phenotype of aaBbCCo r AaBbCc
would have a diploid number of 41 and an animal with AABbCC would have a
diploid number of 43. I have determined the chromosomal phenotype for 333
specimens from the zone of contact.

C-banding patterns are important because they identify segments of the
chromosomes that are believed to be heterochromatic in nature. Variation in

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SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

the amount of C-band material between karyotypes is not thought to interfere
with meiosis as does variation in euchromatin. It is important, therefore, to know
the amount and placement of C-band positive material within the three chro¬
mosomal variants. The karyotype of Uroderma bilobatum contains very little
C-band positive material. Most biarmed elements have a small amount near the
centromere and one medium-sized acrocentric (not one of the A, B, or C chro¬
mosomes) has a C-band proximal to the centromere. Although all of the A, B, and
C chromosomes have a small amount of centromeric C-band material, none of the
major segments involved in the variation is C-band positive. The small biarmed
pair involved in the Aa variation, however, has heterochromatin incorporated
into part of one arm. All of the C-bands identified in this small biarmed element
(of the A morph) are present in the subtelocentric a morph fusion product and
the a morph subtelocentric has about as much C-band material as do the two
elements of the A morph.

Although the break and alteration may have occurred in this C-band positive
area, no major addition or deletion of C-band material seems to have occurred.
Variations in the C-banding patterns do not seem to be involved in the genetic
strategy of Uroderma. This constitutes a major difference between chromosomal
evolution in this species and that seen in some rodents, for example, Peromyscus
(Duffey, 1972; Pathak et al., 1973).

The zone of contact between the cytotypes of Uroderma is about 200 kilometers
in length but its width is not known. Because Uroderma is ecologically restricted
to the relatively low lands of the coast, the zone cannot be over 100 kilometers
wide at many places and must be considerably narrower at some. The two parental
cytotypes occur sympatrically at a single locality in my sample (Fig. 4), and the
area of overlap of parental cytotypes is probably not much longer than 30 kilo¬
meters. At the locality where the two parental cytotypes occur sympatrically, most
individuals have a hybrid karyotype (for instance, within a sample size of 15,
one bat had 2n — 38, one had 2n=39, two had 2/7 = 40, five had 2n = 4l, one had
2u = 41, one had 2/7 = 42, three had 2/7 = 43, and two had 2/7 = 44). Intensive
hybridization occurs in the central part of the zone between Nacome and Choluteca,
Honduras. Away from this area, parental cytotypes probably do not come into
direct contact, and hybrid karyotypes are found much less frequently; I suggest
that these are primarily the result of the survival and successful reproduction of
backcross individuals.

Different types of chromosomal rearrangements produce different meiotic
aberrations and, therefore, the percentage of sterile gametes in a heterozygote
will be a function of the nature of the rearrangement. If the rate of production of
sterile gametes is the only factor regulating the penetration of a chromosomal
morph of one parental type into a population of the other parental form, an increase
of sterile gametes should result in a decrease in successful penetration into the
other cytotype. Furthermore, across the zone of contact the frequency of the
penetrating chromosomal morphs should produce a symmetrical bell-shaped
curve reflecting the greater number of Fi backcross individuals near the zone and
the decrease in such individuals with distance away from the area of primary

BIOLOGY OF THE PHYLLOSTOMATIDAE

129

contact. The width of this symmetrical curve for a given chromosomal aberration
would be a function of the severity of meiotic selection against heterozygotes of
that type of aberration.

If factors other than meiotic mechanisms play a role in the penetration of one
chromosomal morph into populations of the other, there is no reason to assume
that selection on both sides of the zone should be the same and the frequency of Fi
and heterozygous individuals across the zone would not be symmetrical.

The frequencies ( p and ^-values) of the various chromosomal morphs from
333 specimens of Uroderma bilobatum from the contact zone are shown in Table
2. The two northernmost localities (La Herradura and Usulatan) have similar
chromosomal frequencies. Notably, the b morph of the B chromosomes has been
the most successful in surviving in these populations, whereas the c morph was
not found to be present in any of the 133 specimens from these two localities.
This might be predicted based on the type of segregation that would result in a
heterozygote for the respective B and C chromosomes. Centric fusions and
fissions (origin of the B chromosomal system) are not believed to interfere greatly
with the meiotic process, especially if preferential segregation occurs. Proper
segregation probably would not be affected, and therefore natural selection at the
meiotic level would be ineffective in eliminating such variation from the popu¬
lation. On the other hand, such translocations as might have given rise to the
C chromosomal variation should result in loss of about 25 per cent of the gametes
in the heterozygote if there has been crossing over in the portion homologous to
the large acrocentric. It would appear that, in the absence of other factors, the
variation in the B chromosomes would be more common in all populations than
would variation in the C chromosome. In samples from the southeastern part
of the contact zone, survival of the B chromosome is less frequent than C; C
actually accounts for about 4.5 per cent of the C chromosomes at the Choluteca
locality (Table 2, Fig. 4). Two of 86 individuals were heterozygous (Cc) at the
Chinandega locality. The per cent variation resulting from each chromosomal
change is not the same northwest and southeast of the central part of the zone (Table
2), which suggests that successful reproduction of hybrid and backcross individuals
is not explained totally by meiotic problems, but that possibly fitness of the adult
varies as well.

It also should be noted that although Chinandega is closer to the central part
of the contact zone than is La Herradura, less total chromosomal variation is
found at Chinandega (4.6 per cent of the individuals had hybrid karyotypes) than
at La Herradura (14 per cent had hybrid karyotypes).

Baker et al. (1975) concluded that the chromosomal data pointed to con¬
siderable chromosomal flow between the cytotypes. At that time it was not
possible to identify patterns in exchange and survival of the different morphs.
From the above data (Table 2), there is clearly a pattern of selective chromosomal
flow between cytotypes. If the variation in the C chromosomes is used to estimate
chromosomal flow (and implied gene exchange) of the 2n = 38 chromosomes into
the 2n = 44 populations, the data strongly suggest no exchange (the one individual
at Nacome that was heterozygous, Cc, was a presumed Fi). On the other hand, if

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SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

Table 2. — Frequency of the chromosomal morphs at localities in the zone of hybridization.
Numbers preceding localities identify geographic samples in Fig. 4, where exact localities
are given. The 2n = 44 parental type occurs at localities 1-3. Both parental cytotypes occur at
locality 4. At localities 5-7, the 2n = 38 parental type is present.

1 . La Herradura

2. Usulatan

3. Nacaome

4. Hybrid
locality

5. San
Lorenzo

6. Choluteca

7. Chinandega

Sample size

Chromosomal

morphs

p = 98; q = 02

p= 99; q = 01

p = 94 ; q = 06

p = 60; q = 40

p = 42; q — 58

p = 05; q = 95

p = 01; q= 99

p = 95; q — 05

p= 95; q = 05

p = 78; q = 22

p = 53; q = 47

p = 29; q = 71

p = 01; q = 99

p = 00; q = 100

p= 100; q = 00

p = 100; q = 00

p = 94; q = 06

p= 57; q = 43

p= 33; q = 67

p= 04; q = 96

p = 01; q= 99

the B chromosomes are used, the implications are different. Chromosomal data
fit the pattern of introgression in which some chromosomes are allowed to enter
the “chromosome pool” of another type by hybridization and backcrossing,
but other chromosomes are selected against.

The pattern of chromosomal morphs across this contact zone closely fits the
tension zone (White, 1973; Key, 1974) characteristic of stasipatric speciation.
In evaluating my data in light of White’s model, several points need to be made.
First, at this time it is impossible to determine if this zone is the result of primary
or secondary contact. White’s model requires that the zone be the product of
primary contact. Second, stasipatric tension zones have been described for several
species (Bush, 1975), and a suite of the biological characteristics of these species
do not fit those of Uroderma. In species with low vagility, the tension zone is
usually not more than a few hundred meters wide; in Uroderma , species with high
vagility, the zone is more than 200 kilometers in breadth. Third, Uroderma is
K-selected, whereas other species with tension zones are R-selected.

My data point out the fact that tension zones need not be composed of species
characterized by low vagility and R-selection. Although the zone of inter¬
action between the two Uroderma cytotypes might or might not be in equilibrium,
it will eventually proceed to one of several endpoints. One possibility is that the two
cytotypes could develop additional isolating mechanisms, such as behavioral or
postmating, and evolve into two species. Another possibility is the replacement of
one parental type by the other via the mechanisms of competition or genetic
swamping. A less likely outcome could be the survival of some intermediate
cytotype with, say, 42 chromosomes (for instance, AABBcc). At any
rate, this type of chromosomal variation undoubtedly offers a unique set of
possibilities on which evolution can act. The unique nature of these biological
circumstances certainly offers a rare chance to observe evolution in action.

Electrophoretic data would be extremely valuable in shedding some light on
the history of Uroderma populations that have produced this tension zone.
Electrophoretic data indicate that when two species have been derived by the
classical allopatric model, the level of similarity of allozymes is usually about 85
per cent or less (Avise, 1974). If these chromosomal differences accumulated
during a long allopatric period, it could be predicted that these two chro¬
mosomal races should have accumulated a significant number of fixed allelic

BIOLOGY OF THE PHYLLOSTOMATIDAE

131

I II ** I i ii

I! il

• •

I • * • *

* *

# 41

» §

Fig. 5. — Sample patterns of C-bands of phyllostomatid bats: A, Phyllostomus elongatus ;
B, Enchisthenes hcirtii.

differences; however, if the process that gave rise to the current condition has been
like that proposed by White (1968, 1973), very few electrophoretic differences
should be detectable. This situation is currently under study by Ira F. Greenbaum.

Miscellaneous Cytogenetic Studies

In addition to the more systematically oriented papers discussed above, there
have been a few more detailed studies on biochemical aspects involving karyo¬
typic data for phyllostomatid bats.

C-bands. — C-band material for phyllostomatid bats is described in enough
species that general trends can be predicted (Stock, 1975; Patton, 1976; Baker
et al., 1978). In general, phyllostomatid bats have C-band material restricted
to the centromeric region. The amount found is small, similar to that shown in
Fig. 5 for Phyllostomus and Enchisthenes , respectively. However, in some species
( Carollia perspicillata and Choeroniscus intermedius) there are additional portions
of the karyotype carrying C-band positive material (Stock, 1975). Also, see the dis¬
cussion on C-band material in Uroderma under the section on chromosomal races.

Nucleolar organizer regions. — Two papers, both dealing with Carollia
perspicillata and C. castenea, have reported studies of nucleolar organizer
regions (NOR) in phyllostomatids (Hsu et al., 1975; Goodpasture and Bloom,
1975). Hsu et al. (1975) used DNA/rRNA (ribosomal RNA) hybridization to
reveal NOR’s. In the karyotype of C. perspicillata, the only NOR was located on
s the X chromosome; their studies of C. castanea were made on a transformed
culture. Hsu et al. concluded that the origin of the NOR on the Carollia X/auto-

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SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

somal chromosome was from the X and not the translocated autosomal portion.
Because DNA-RNA hybridization is difficult and expensive, Goodpasture and
Bloom (1975) tested the feasibility of using ammoniacal silver to reveal NOR’s.
Their methods localized NOR’s at the same points as did the methods of Hsu
et al. The same individuals were studied from in vitro cultures. Goodpasture and
Bloom (1975) present theories on the cytological basis for silver NOR staining.

Cesium chloride buoyant densities. — Arrighi et al. (1968, 1972) reported on
cesium chloride buoyancy in phyllostomatid bats. Findings are summarized in
the latter paper. Ten species of phyllostomatid bats ( Anoura geoffroyi, Artibeus
fallax= A. lituratus in Jones and Carter, 1976, Artibeus lituratus, Carollia
perspicillata, Chiroderma villosum, Choeroniscus intermedius, Sturnira
erythromos, Sturnira lilium , Sturnira magna , and Uroderma bilobatum ) were
studied and values ranged from 1.6982 for Carollia perspicillata to 1.7005 for
Anoura geoffroyi and Sturnira erythromos. These values fall within those given
for other Microchiroptera (1.696 to 1.702) from the families Rhinolophidae,
Molossidae, and Vespertilionidae, but only slightly overlap the values reported for
Megachiroptera (1.694 to 1.697). Of the families of Microchiroptera, Phyl-
lostomatidae had values nearest those for the Megachiroptera. Although the
magnitude of difference between the suborders is small, it is the greatest found
between suborders of mammals and is interpreted as supporting relatively
ancient lineages for the two suborders (Arrighi et al., 1972).

X chromosomes. — G-banded X chromosomes for a variety of mammals
(including eight species of phyllostomatids) were studied by Pathak and Stock
(1974). They found that X chromosomes always have two dark staining, trypsin
resistant bands regardless of the centromere placement. They interpreted these
data as supporting Ohno’s (1967) hypothesis that the mammalian X chromosome
is extremely conservative in genetic constitution.

Acknowledgments

Laura Kyle and Anette Johnson assisted in the preparation of plates and Table
1 . I am especially grateful to my graduate students who have been of considerable
assistance in accumulating these data. I also thank my many colleagues who aided
in the collection of specimens, read drafts of the manuscript, and offered
criticisms. This work was made possible by several grants from the National
Science Foundation, of which the most recent was No. DEB-76-20580.

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- . 1977. Implications of chromosomal variation in Rhogeessa (Chiroptera:

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136

SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

XX XI! XX XX XX u k»

XK XX XX XX ft* «* *»

«« M •

Plate 1 . — Karyotype of a male Lonchorhina aurita from Trinidad.

8 n n *i « a «

4* **

Ah A* i* no AO At At

fit M »• •• »»* ••

Plate 2. — Karyotype of a female Macrotus waterhousii from Haiti.

nn U on XX IS XX

Xl{ JfX XX »» ** ** **

X3*

Plate 3. — Karyotype of a female Micronycteris brachyotis from Trinidad.

BIOLOGY OF THE PHYLLOSTOMATIDAE

137

x* OA

/Ml AO no (Ia 04 Ax on

^ 1^ AA •»* ” •* ft

Plate 4. — Karyotype of a male Micronycteris hirsuta from Nicaragua.

Aft HU IX *x Aft A» |«

•» mm

(ift XR ha ax

Plate 5. — Karyotype of a male Micronycteris megalot is from Trinidad.

K6 xx f\j» xx x» x a

KM AM »*

lift flft A/» — A

Plate 6. — Karyotype of a male Micronycteris minuta from Trinidad.

138

SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

U 88 88 88 ** XX

XX XX X»

f)» to to 8.

Plate 7. — Karyotype of a male Micronycteris nicefori from Trinidad.

88 xx x* ** *« *»

K K • « MM

ilArt/) Ml IfK M

AM A i| «A X*

Plate 8. — Karyotype of a male Micronycteris schmidtorum from Costa Rica.

M88K8 KUKxa xk

XX XX XX *X XX xa XX

Plate 9. — Karyotype of a male Mimon cremilatum from Colombia.

BIOLOGY OF THE PHYLLOSTOMATIDAE

139

KX KK XX XX xx »*

*>\ UK MX XX xk «« **

"* K .

Plate 10. — Karyotype of a male Phylloderma stenops from Colombia.

«!i M X» KJI MS K# l«

XX KX *X XX X* *«
** X.

Plate 11. — Karyotype of a male Phyllostomus discolor from Trinidad.

MM (tamii

XX XX XX XA XX XX »x

Plate 12. — Karyotype of a female Phyllostomus elongatus from Colombia.

140

SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

/WUX KUtlxnu*

X# ** >«* «* xn **

Plate 13. — Karyotype of a male Phyllostomus hastat us from Trinidad.

at «« is*

Oft 00 AO Aft ft*

Plate 14. — Karyotype of a female Tonal ia bidens from Trinidad.

Hi ft! XX IX Xt n» **

xx XX xx xx as »•

A A

Plate 15. — Karyotype of a male Tonatia minuta from Trinidad.

BIOLOGY OF THE PHYLLOSTOMATIDAE

141

K! U It h n m n

XK X« xk x* Xfl **

M (14

Plate 16. — Karyotype of a female Trachops cirrhosus from Trinidad.

% XX n xh xk n xx

fix XX XX XX XX XX XX

Plate 17. — Karyotype of a male Vampyrum spectrum from Trinidad.

22 it a ii M
X* »*

16 16 it M XX X.

Plate 18. — Karyotype of a male Anoura caudifer from Colombia.

142

SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

Oft X# M *tc ** XX

XX mx

it il iA n* H*

Plate 19.— Karyotype of a male Anoura cultrata from Costa Rica.

/t :« Mi

II II Aft A* Aft i< *u

Plate 20. — Karyotype of a female Choeroniscus godmani from Honduras.

AA **

ftfi (SK M AA AA AA

Plate 21. — Karyotype of a female Choeroniscus intermedins from Trinidad.

BIOLOGY OF THE PHYLLOSTOMATIDAE

143

Plate 22. — Karyotype of a female Choeronycteris mexicana from Tamaulipas.

n n n n n t* a

XX XX X| (A H *x »

” X.

Plate 23. — Karyotype of a male Glossophaga soricina from Colombia.

KK U „

nn An

x x

Plate 24. — Karyotype of a female Hylonycteris underwood i from Costa Rica.

144

SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

XX XX Al

U HD ill Afl »

Plate 25. — Karyotype of a female Lichonycteris obscura from Nicaragua.

il ii xx H ii » «i

>* i* »a

6A fifi .. t.

Plate 26. — Karyotype of a male Lionycteris spitrrelli from Colombia.

HH till KM

nr- m

Plate 27. — Karyotype of a male Lonchophylla robusto from Nicaragua.

BIOLOGY OF THE PHYLLOSTOMATIDAE

145

AA AA Aft

00 Oft ft# AA Aft Art Art

AO OA Art »•

Plate 28. — Karyotype of a female Lonchophylla thomasi from Colombia.

18 M XU Un xx ix

XX XX XX XX xr aa

«•

Plate 29. — Karyotype of a male Monophyllus redmani from Puerto Rico.

Ktt AX «s

SIX

ttft.

Plate 30. — Karyotype of a male Carollia perspicillata from Colombia.

146

SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

Hi! 88 88 xx «x xx xx

X8 XA xx

ns ««

<io no ao »•

Plate 31. — Karyotype of a male Rhinophylla fischerae from Colombia.

KK AH XX XX *Jt «x »4

Aft O A Xl

Plate 32. — Karyotype of a male Rhinophylla pumilio from Colombia.

<*8 M XX xx *x xx

XK XX K* «* »*

U XX AX li »*

Plate 33. — Karyotype of a male Ametrida centurio from Trinidad.

BIOLOGY OF THE PHYLLOSTOMATIDAE

147

K B & II 19 n u

SI xa

II n u u U

Plate 34. — Karyotype of a male Ardops nichollsi from Guadeloupe.

XX XX XX xx xx

XX «x **

Aft Aft xx u IU-

Plate 35. — Karyotype of a male Ariteus flavescens from Jamaica.

XK U 8R kx n xx

XX a* ah

ftft AA Aft KX A..

Plate 36. — Karyotype of a male Artibeus lituratus from Colombia.

148

SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

)'K U NK Kfc VJf >tx Jt*

x a ** **

Aft Aft Aft ft* O"

Plate 37. — Karyotype of a male Artibeus pluieotis from Colombia.

Cl U W UK M XX

XX *n

ill! y ~ I.

Plate 38. — Karyotype of a male Chiroderma improvisum from Guadeloupe.

lc t! I* XI »* ** «»

IX «*

it it i< 1*

Plate 39. — Karyotype of a male Chiroderma salvini from Honduras.

BIOLOGY OF THE PHYLLOSTOMATIDAE

149

H !

U si si st

ii M

Plate 40-

ii U 5*

-Karyotype of a male Ectophylla alba from Costa Rica.

n jx

Its ns

Plate 41.-

fill

—Karyotype of a

male Enchisthenes hartiii

Ai ft,.

from Colombia.

nn on

r* n

Plate 42. — Karyotype of a male Mesophylla macconnelti from Trinidad.

150

SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

HI Hn 8X XX U 9»

XU XX *»

M u n .1,1 8.»

Plate 43. — Karyotype of a male Phyllops haitiensis from Haiti.

U M Kt K.h u n

II U (A >1 » XI

if.

Plate 44. — Karyotype of a male Sphaeronycteris toxophyllum from Colombia.

KX XX XX XX XX XX

XX X* *K

(MS ftrt no xx A.

Plate 45. — Karyotype of a male Sturnira erythromos from Colombia.

BIOLOGY OF THE PHYLLOSTOMATIDAE

151

n 22 n xs « « «

S U X * X 21

U 6» io M

Plate 46. — Karyotype of a female Sturnira mordax from Costa Rica.

XX xx xx *>

Ad Aft MAX AX Aft AX

AA A* XX

6A A* X A MX ft.

Plate 47. — Karyotype of a male Uroderma magnirostrum from Colombia.

XX nX n XX XX XX XX

** aa

Kft Aft hf>

Plate 48. — Karyotype of a female Vampyressa brocki from Colombia.

152

SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

M XX xx xx xx ** «

«ft WoiiAM* A.

Plate 49. — Karyotype of a male Vampyressa nymphaea from Honduras.

U xx

Aft flA Oft oo aa **

Plate 50. — Karyotype of a male Vampyressa pusilla from Honduras.

ftR HO 00 lift Aft no ,<A

M tifi **

Plate 51. — Karyotype of a male Vampyressa pusilla from Colombia.

BIOLOGY OF THE PHYLLOSTOMATIDAE

153

U KX XX XX xx xx xx

XX Xtt MM

ftl UftAMA Aa

Plate 52. — Karyotype of a male Vampyrops vittatus from Colombia.

XI II Id IX 88 ti xx

tx XX XX a* ** ** **

Plate 53. — Karyotype of a female Brachyphylla caver narum from Puerto Rico.

81 88 XX 88 tf XX xx

IX *X xx *» *« ** "

Plate 54. — Karyotype of a male Brachyphylla nana from Haiti.

154

SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

ft U IK H n n

21 XX xx 14 *» *» tx

Plate 55. — Karyotype of a male Erophylla sezekorni from Puerto Rico.

XX XX XX XX x* XX

XA XX xx X* a* x* •«

A* K.

Plate 56. — Karyotype of a male Phyllonycteris aphylla from Jamaica.

;( n » si is xi ii

<1 XX « *» " “ !*

Plate 57. — Karyotype of a male Phyllonycteris poeyi from Haiti.

BIOLOGY OF THE PHYLLOSTOMATIDAE

155

XI lit Kt ss n g* xx

X. XX XX XX XX ft* X

Plate 58. — Karyotype of a male Desmodus rotundus from Veracruz.

M M U Xfi 8K xx xx

KK KX U XX A& Xft

i? «•

Plate 59. — Karyotype of a male Diaemus youngii from Nicaragua.

U IS n 88 n n it

88 kx x* « x* ** **

** s.

Plate 60. — Karyotype of a male Diphylla ecaudata from Veracruz.

BIOCHEMICAL GENETICS

Donald O. Straney, Michael H. Smith, Ira F. Greenbaum,

and Robert J. Baker

The current view of evolution is very much a genetic one. Theoretical develop¬
ments since the rediscovery of Mendel’s work have produced an intricate
mathematical theory, integrating genetic and ecologic characteristics, that
provides the basis for our understanding of the evolutionary process. Within this
theoretical framework are two genetic factors of critical importance: determination
of the genetic basis of fitness and the genetic structure of populations in space and
time. Unfortunately, information about these two factors is lacking for most
groups of organisms. The first is nearly impossible to establish (Lewontin, 1974),
and the second requires intensive breeding studies. Until recently, the spatial and
temporal genetic structure of natural populations had been described only for
Drosophila and a small number of other groups (Dobzhansky, 1970). In order to
apply theoretical evolutionary concepts to organisms such as bats, which are
difficult to breed in captivity, it has been necessary to assume that these organisms
behave genetically in a manner similar to that of Drosophila.

Most species of phyllostomatid bats are difficult or impossible to maintain in
captivity in the numbers required for genetic breeding studies (see Greenhall,
1976). In addition, lengthy gestation periods and low productivity make
chiropterans, in general, an inefficient group with which to work. Bats also exhibit
few clear-cut phenotypic variants within populations that could be exploited in
genetic studies, as has been done with Drosophila. Thus, the genetic properties
of chiropterans, in the classical sense, are unknown. It is not surprising that,
among mammals, easily tractable, prolific and variable groups, such as the rodent
genera Mus and Peromyscus, have been used to establish genetic baselines
(Rasmussen, 1968).

The development of biochemical techniques, such as electrophoresis, has
enabled genetic studies to be carried out at the protein level, thereby circumventing
many of the traditional problems mentioned above concerning maintaining and
breeding animals. Large numbers of individuals now can be assayed quickly, even
in species that cannot be bred in the laboratory, to give baseline data documenting
the spatial and temporal structure of natural populations. Breeding studies are
needed only to establish the inheritance of protein banding patterns, and for
most of the species studied so far, the inheritance of these banding patterns
appears to be the same (Selander et al., 1971, Straney et al., 1976a, 1976b). Al¬
though electrophoresis and other biochemical techniques do not provide a com¬
plete picture of evolutionary genetics, they can furnish information useful in
developing models of evolution and do have the potential for providing data that
can be used in testing phylogenetic hypotheses. Few families of eutherian mam¬
mals are as ecologically diverse as are phyllostomatid bats, but the genetic as-

157

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SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

pects related to this group’s adaptive radiation are poorly understood. It is from
studies of organisms such as the Phyllostomatidae that information on the rela¬
tionships of genetic, ecological, and morphological strategies can be obtained.
Study of phyllostomatid genetics though, has only begun and is limited to karyo¬
typic (Baker, this volume) and biochemical characters; available information
points more toward potential questions than to a unified picture of chiropteran
genetics. In this chapter, we review the published works on biochemical genetics
of phyllostomatid bats and present new data on several species from Trinidad.

Literature

Several methods have been used to study the biochemical genetics of phyl-
lostomatids. Most involve electrophoresis in some form of supporting medium,
such as cellulose acetate, polyacrilimide, or starch gel. Despite differences in
medium, the process is the same. Proteins in tissue extracts are placed in the
medium and an electric current applied through an electrode bridge. The pro¬
teins are ionized by the buffer used in the electrode bridge, migrate in the
electrical field in characteristic manners, and are identified by means of appro¬
priate histochemical stains. Differences in mobility between proteins are indicative
of variation in net electric charge on the molecules. Charge variation results
from changes in the amino acid composition of the proteins, which ultimately
reflect codon differences in the genes involved. Hence, differences in mobility
of proteins assayed under the same conditions are translatable into genetic dif¬
ferences.

The earliest examinations of chiropteran biochemical genetics focused on
vespertilionids and were conducted by Mitchell (1966), working with hemoglobin,
and Manwell and Kerst (1966), with hemoglobin, lactate dehydrogenases,
esterases, and general tissue proteins. Both papers established the multiple
component structure of chiropteran hemoglobin, and Manwell and Kerst (1966)
found genetic polymorphisms in several species that involved at least two alleles
at the lactate dehydrogenase- 1 locus and several alleles at the esterase and tissue
protein loci. Differences in protein mobility of several species and genera were
interpreted as genetic variation at loci encoding these proteins.

Variation in bat hemoglobins has been studied in some detail by Mitchell (1970),
Valdivieso et al. (1969), and Tamsitt and Valdivieso (1969). Differences in
hemoglobin molecules were found primarily at the familial level, although within
the vespertilionids examined there was a high degree of variation and poly¬
morphism; of the phyllostomatids studied, the same hemoglobin moiety was
present. Peptide mapping (Mitchell, 1970) confirmed the identity of the phyl¬
lostomatid hemoglobins. Desmodus hemoglobin (Tamsitt and Valdivieso, 1969)
was found to be the same as that for nine other species of phyllostomatids, whereas
hemoglobin from Pteronotus was unique, results consistent with current taxonomic
views (Smith, 1972; Jones and Carter, 1976). Our examination of samples of
phyllostomatids from Trinidad (see below) suggests that variation exists in
hemoglobins of some species of this family. The inheritance of this variation is

BIOLOGY OF THE PHYLLOSTOMATIDAE

159

not clear although banding patterns suggest allelic variation in a monomeric
protein, possibly in only one of the hemoglobin chains.

Valdivieso and Tamsitt (1974) examined serum proteins of 18 species from
four families of Neotropical bats and were able to isolate four to eight protein
fractions. Of the 14 species of phyllostomatids they examined, six exhibited
polymorphism in a -globulins; only Artibeus was polymorphic at both a - and />-
globulin loci. All species were monomorphic for a -globulin. Valdivieso and
Tamsitt found no polymorphism in phyllostomatid albumins; however, in our
samples from Trinidad, albumin is the single most variable protein locus (see
below). Although these authors noted differences in albumins between species,
genera, and families, their differences are not concordant with our data (Table 1).
Their finding that the albumin of Phyllostomus hastatus and P. discolor differ
from all other phyllostomatids appears to be a result of sampling error. Albumin
allozymes identical to those of Phyllostomus were present in other phyllostomatids
in our samples (Table 1). The fact that in their sample Molossus albumins were
indistinguishable from those of some phyllostomatids is probably due to the use
of cellulose acetate as an assay medium. Although cellulose acetate makes a
quick and effective medium for assaying serum protein profiles, the accompany¬
ing lack of resolution makes it a poor system for surveys of genetic variation.
Their (Valdivieso and Tamsitt, 1974) conclusion that serum protein electrophore¬
sis will be of little use in systematic work is a result of the assay medium em¬
ployed, the number of species examined, and sample size.

Straney et al. (1976 a) and Greenbaum and Baker (1976) used starch gel assay
systems to examine genetic variation at 17 and 21 loci, respectively, in popu¬
lations of Macrotus. In 45 individuals sampled from a population of M. californicus
in Pima County, Arizona, Straney et al. described six polymorphic loci, but the
level of polymorphism was low, with no locus segregating for more than two
alleles. Indeed, the proportion of loci in the heterozygous state in the average
individual (H) in this population was 0.03, a value low for mammals and much
less than that found in Myotis velifer{\\ =0.14; Straney et al., \916a). The authors
suggested that the low level of variation in Macrotus was consistent with the
niche width-variation hypothesis, as modified by Selander and Kaufman (1973).

Greenbaum and Baker (1976) examined genetic variation and intra and
interspecific similarity in Macrotus californicus and M. waterhousii from Arizona,
Mexico, and Jamaica. In addition to the polymorphisms mentioned above, they
described others at two gene loci in populations outside of Arizona. Average
population heterozygosity ranged from 0.030 to 0.041 in M. californicus and from
0.00 (for specimens from an interspecific contact locality) to 0.043 in M. water¬
housii.

Nei’s genetic distance (D; Nei and Roychoudhury, 1974) reflects the number
of net codon differences per locus between a pair of populations. Genetic distance
between populations of the same species of Macrotus are less than 0.07. Esti¬
mates of D among populations of Macrotus are within the range reported for
other mammals (Greenbaum and Baker, 1976). Jamaican M. waterhousii are

160

SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

Table 1. — Variation in albumin in Neotropical phyllostomatid bats. Listed are those species
examined both in this study and by Valdivieso and Tamsitt (1974). Entries are relative
mobility of albumin allozymes, Artibeus jamaicensis taken as 100. Where more than one
allele is present in a population, mobilities are listed in decreasing order of frequency.

Species

This study

Valdivieso and Tamsitt1

Artibeus jamaicensis

100, 101, 103

100, 106

Artibeus lituratus

103, 100

106

Carol lia perspicillata

105, 104.5, 100

106

Phyllostomus discolor

104.5

87.5

Glossophaga soricina

111, 127

Desmodus rotundas

127

100

Sturnira l ilium

127

100

Walues from measurements of mobility as indicated in fig. 3 of Valdivieso and Tamsitt (1974). Alb100
is taken as the most common allozyme in A. jamaicensis.

12 times as distant from mainland populations of this species as the latter are
among themselves (D = 0.065 and 0.005, respectively). Although this difference
involves very small D-values and is not statistically significant, it is consistent
with the view that Jamaican populations have been isolated from those on the
mainland for some time. This isolation might have resulted in genetic differenti¬
ation of Jamaican populations sufficient to warrant recognizing them as belong¬
ing to a separate subspecies, a conclusion reached by Anderson and Nelson
(1965) based on morphological analysis of M. waterhousii from Jamaica and
Mexico.

The genetic distance between species of Macrotus is substantial (D = 0.4); at
least 40 per cent of the loci in the two species having accumulated codon changes
since separation from a common ancestor. This value is high for congeneric
species of mammals and is near the value reported for intergeneric comparisons
of the vespertilionids Myotis and Pipistrellus (Straney et al., 1976/?). Indeed, this
value is nearly equal to that found separating Glossophaga and Desmodus (D —
0.35; see below), members of different phyllostomatid subfamilies. It was con¬
cluded that the large genetic difference between M. californicus and M. water¬
housii was a product of independent evolution during a long period of separa¬
tion — current parapatry represents secondary contact. Temporal calibration of
Nei’s D in phyllostomatids, discussed below, suggests that these species have
been separated for approximately 10 million years. Yet, during this time, al¬
though protein loci have diverged, morphological change has been slight (Ander¬
son and Nelson, 1965; Davis and Baker, 1974).

The electrophoretic analysis of Macrotus (Greenbaum and Baker, 1976) clearly
indicated that mainland Macrotus represent two species and that Antillean
populations are conspecific with Mexican M. waterhousii. Their study suggests
great potential for electrophoretic application to systematic problems on an
intrageneric level. Published information on biochemical genetics of phyllosto¬
matid bats establishes the presence of polymorphic and polytypic genetic variation
in members of the family. The results of Greenbaum and Baker (1976) and

BIOLOGY OF THE PHYLLOSTOMATIDAE

161

Table 2. — Gene loci and assay systems examined in Trinidad phyllostomatids.

Protein System

Buffer system'

Voltage

Time (hr.)

a-Glycerophosphate dehydrogenase (a-GPD)

Tris citrate

8.0

130

3.5

Albumin (ALB)

Lithium hydroxide

8.1

350

Alcohol dehydrogenase (ADH)

Phosphate

6.7

130

Glutamic oxaloacetic transaminase- 1 (GOT-1) Lithium hydroxide

8.1

350

Glutamic oxaloacetic transaminase-2 (GOT-2) Tris citrate

8.0

130

3.5

Isocitrate dehydrogenase (IDH-1, 2)

Tris citrate

6.7

150

Indophenol oxidase (IPO)

Lithium hydroxide

8.1

350

Lactate dehydrogenase- 1 (LDH-1)

Lithium hydroxide

8.1

350

Lactate dehydrogenase-2 (LDH-2)

Lithium hydroxide

8.1

350

Malate dehydrogenase- 1, -2 (MDH-1, -2)

Tris citrate

6.7

150

Phosphoglucomutase-1, -2 (PGM-1, -2)

Tris citrate

6.7

150

Phosphoglucose isomerase-1, -2 (PGI-1, -2)

Poulik

8.5

250

3.5

6-Phosphogluconate dehydrogenase (6-PGD)

Tris maleate

7.4

100

'Details of preparation in Selander et al., 1971.

Straney et al. (1976a) indicate a low level of genetic variation in the average
population of Macrotus. The level of divergence observed by Greenbaum and
Baker suggests that phyllostomatid taxa may be genetically quite distinct.
New data collected on the genetics of phyllostomatids from Trinidad, summarized
below, allow these points to be examined in more detail.

Implications of Genic Variation in Phyllostomatids from Trinidad

In August, 1974, we collected samples of 14 species of phyllostomatid bats at
six localities in Trinidad. Assay systems were similar to those described by
Straney et al. (1976 a) and Greenbaum and Baker (1976). Table 2 lists gene loci
examined and conditions of assays. Several proteins were examined but could
not be interpreted due to progressive denaturation (malic enzyme- 1, -2; hemo¬
globin). Esterases presented such a complex pattern that it was not possible to
establish locus homologies and these proteins have been disregarded.

Table 3 presents a summary of gene frequencies in the populations examined.
In many cases sample sizes are quite small and doubtless some polymorphic
loci were missed. Albumin was, as mentioned above, the most polymorphic
locus, segregating for two or three alleles in the three species of Artibeus sampled,
as well as in Chiroderma, Carollia, and Glossophaga. Other loci that show
relatively high levels of heterozygosity are 1DH-1 (A. jamaicensis and Anoura),
a-GPD (Carollia), and PGM-1 (Carollia). All other variable loci either are
present in samples too small to give fair estimates or show a proportion of
heterozygotes less than 0.10.

Genetics and Ecology

Table 4 summarizes heterozygosity values for all species of bats thus far
examined. Values from this study are restricted to populations with sufficient

162

SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

Table 3. — Alleles and frequencies (in parentheses) at 16 gene loci in

Species

Locality1

“ GPD

Alb

GOT-1

GOT-2

IDH-1

1DH-2

IPO

Phyllostomatidae

Ametrida centurio

75(1.00)

105.5(1.00)

83(1.00

-100(1.00)

800.00)

-67(1.00)

1000.00)

Artibeus cinereus

100(0.92)

62(0.08)

102(0.92)

101(0.08)

56( 1 .00)

-94(1.00)

87(1.00)

-67(1.00)

2000.00)

100(0.86)

62(0.14)

102(0.72)

101(0.21)

103(0.07)

56(1.00)

-94(1.00)

87(1.00)

-67(1.00)

200(0.79)

100(0.21)

Artibeus jamaicensis

100(1.00)

100(0.56)

103(0.27)

101(0.17)

100(1.00)

-1000.00)

100(0.50)

87(0.50)

-100(1.00)

1000.00)

100(0.98)

123(0.02)

100(0.48)

103(0.32)

101(0.20)

100(0.98)

56(0.02)

-1000.00)

87(0.57)

100(0.43)

- 100(1.00)

100(0.93)

50(0.07)

100(1.00)

100(0.50)

103(0.45)

101(0.05)

100(0.90)

56(0.10)

-100(1.00)

100(0.50)

87(0.50)

-100(1.00)

100(0.95)

50(0.05)

100(1.00)

100(0.67)

103(0.16)

101(0.16)

100(1.00)

-1000.00)

100(0.62)

87(0.38)

-100(1.00)

1000.00)

100(1.00)

100(0.50)

101(0.38)

103(0.12)

100(1.00)

-1000.00)

100(0.62)

87(0.38)

- 100(1.00)

1000.00)

Artibeus lituratus

1, 2, 3, 5

100(0.86

62(0.14)

103(0.93)

100(0.07)

100(0.93)

56(0.07)

-94(1.00)

87(1.00)

-67(1.00)

2000.00)

Chiroderma villosuni

100(0.67)

123(0.33)

106(1.00)

111(0.67)

61(0.33)

-94(1.00)

87(1.00)

-67(1.00)

1000.00)

Sturnira (Species “A”)

1, 3

108(1.00)

98(1.00)

56(1.00)

-1000.00)

50(1.00)

-100(1.00)

1000.00)

Uroderma bilob at um

3, 4, 5

123(1.00)

106(1.00)

56(1.00)

-500.00)

80(0.60)

100(0.40)

-67(1.00)

1000.00)

Vampryrops helleri

1, 3, 4

108(1.00)

100.5(1.00)

56(1.00)

-94(1.00)

87(1.00)

-67(1.00)

200(0.94)

100(0.06)

Carollia perspicillata

123(0.73)

146(0.24)

108(0.03)

105(0.98)

104.5(0.02)

136(0.97)

100(0.03)

-100(0.90)

-50(0.10)

67(1.00)

-67(1.00)

-30(1.00)

123(0.80)

146(0.20)

105(1.00)

136(0.90)

100(0.10)

- 100(0.60)

- 50(0.40)

67(1.00)

-67(1.00)

-30(1.00)

123(0.75)

108(0.15)

146(0.10)

105(0.95)

100(0.05)

136(0.85)

100(0.15)

-1000.00)

67(1.00)

-67(1.00)

-300.00)

123(0.88)

146(0.08)

108(0.04)

105(0.88)

104.5(0.08)

100(0.04)

136(1.00)

-100(0.96)

-50(0.04)

67(1.00)

-67(1.00)

-300.00)

Phyllostomus discolor

123(1.00)

104.5(1.00)

100(1.00)

-125(1.00)

77(0.50)

60(0.50)

-135(1.00)

900.00)

Phyllostomus hast at us

1, 3

123(1.00)

104(1.00)

17(1.00)

-125(1.00)

70(1.00)

-133(1.00)

900.00)

Glossophaga soricina

1, 4, 5

108(0.90)

123(0.10)

107(1.00)

56(1.00)

-50(1.00)

87(1.00)

-67(1.00)

-25(1.00)

108(0.65)

123(0.35)

107(1.00)

56(1.00)

-500.00)

87(1.00)

-67(1.00)

-25(1.00)

Anoura geoffroyi

169(1.00)

101(1.00)

66(1.00)

-94(1.00)

87(0.94)

90(0.03)

77(0.03)

-67(1.00)

150(0.97)

250(0.03)

Desmodus rotundus

2. 4

123(1.00)

107(1.00)

56(1.00)

-31(1.00)

60(1.00)

-67(1.00)

-25(1.00)

Molossidae

Molossus molossus

177(1.00)

104.1(1.00)

63(1.00)

-50(1.00)

73(1.00)

-133(1.00)

75(1.00)

Natal idae

Natalus

100(1.00)

99(0.52)

99.5(0.48)

5(1.00)

-97(1.00)

73(1.00)

-133(1.00)

-200(1.00)

1. Locality designations are:

Las Cuevas, St. George Co.;

2, Maracas

Valley, 2

mi. N (by

road) St.

Joseph, St. George Co.; 3, Guayaguayare, Mayaro Co.; 4, Maracas Valley, 12 mi. N (by road) St. Joseph,
St. George Co.; 5, 2 mi. E, 3 mi. S San Raphael, St. George Co.; 6, Tamana Cave, St. Andrew Co.

BIOLOGY OF THE PHYLLOSTOMATIDAE

163

bats from Trinidad. N is sample size; locus designations are as in Table 2.

LDH-l

LDH-2

MDH-1

MDH-2

PGM-1

PGM-2

PGM

PG1-2

6PGD

100(1.00)

-100(1.00)

100(1.00)

-100(1.00)

100(1.00)

-100(1.00)

175(1.00)

6000.00)

117(1.00)

100(1.00)

-100(1.00)

100(0.92)

60(0.08)

-100(1.00)

240(0.58)

100(0.42)

-100(1.00)

1000.00)

100(0.92)

40(0.08)

100(1.00)

-100(1.00)

100(1.00)

-100(1.00)

100(0.79)

240(0.21)

-100(1.00)

100(1.00)

1000.00)

100(0.97)

40(0.03)

100(1.00)

-100(1.00)

100(1.00)

-100(1.00)

100(1.00)

-100(1.00)

1000.00)

100(0.83)

166(0.17)

100(1.00)

-100(0.98)

-50(0.02)

100(1.00)

-100(1.00)

100(0.98)

240(0.02)

- 100(1.00)

100(1.00)

1000.00)

100(0.98)

166(0.02)

100(1.00)

- 100(1.00)

100(1.00)

-100(1.00)

100(1.00)

-100(1.00)

1000.00)

100(1.00)

- 100(1.00)

100(1.00)

-100(1.00)

100(1.00)

-100(1.00)

1000.00)

100(0.88)

166(0.02)

100(1.00)

-100(1.00)

100(1.00)

-100(1.00)

100(1.00)

-100(1.00)

1000.00)

100(1.00)

-100(1.00)

100(1.00)

-100(1.00)

100(1.00)

-100(1.00)

100(1.00)

1000.00)

97(1.00)

-100(1.00)

100(1.00)

-100(1.00)

100(1.00)

-1000.00)

1000.00)

84(1.00)

- 100(1.00)

100(1.00)

-100(1.00)

380(0.75)

640(0.25)

-183(1.00)

2000.00)

8000.00)

1000.00)

100(1.00)

- 100(1.00)

100(1.00)

-100(1.00)

100(1.00)

-1000.00)

1000.00)

100(1.00)

1000.00)

98(1.00)

-100(1.00)

100(1.00)

- 100(1.00)

100(1.00)

-1000.00)

100(1.00)

1000.00)

91(1.00)

-100(1.00)

100(1.00)

-100(1.00)

240(0.97)

380(0.03)

-1000.00)

62(1.00)

0(1.00)

1000.00)

91(1.00)

- 100(1.00)

100(1.00)

-100(1.00)

240(1.00)

-100(1.00)

62(1.00)

0(1.00)

1000.00)

91(1.00)

- 100(1.00)

100(1.00)

-100(1.00)

240(0.90)

380(0.10)

-1000.00)

62(1.00)

0(1.00)

100(1.00)

91(1.00)

-100(1.00)

100(1.00)

-100(1.00)

240(0.92)

380(0.08)

-1000.00)

62(1.00)

0(1.00)

1000.00)

100(1.00)

- 100(1.00)

100(1.00)

-100(1.00)

100(0.50)

240(0.50)

- 183(1.00)

160(1.00)

550(1.00)

100(1.00)

-100(1.00)

100(1.00)

-100(1.00)

240(0.75)

100(0.25)

-183(1.00)

1600.00)

5500.00)

1000.00)

84(1.00)

-100(1.00)

100(1.00)

-100(1.00)

225(1.00)

-1000.00)

206(1.00)

1000(1.00)

1660.00)

84(1.00)

-100(1.00)

100(1.00)

-100(1.00)

225(1.00)

-1000.00)

206( 1 .00)

1000(1.00)

166(1.00)

96(1.00)

-100(1.00)

80(1.00)

-100(1.00)

380(1.00)

-1000.00)

175(1.00)

7000.00)

90(1.00)

84(1.00)

+ 100(1.00)

100(1.00)

-100(1.00)

380(1.00)

-1000.00)

206(1.00)

10000.00)

1660.00)

68(1.00)

0(1.00)

40(0.98)

62(0.02)

-133(1.00)

38(1.00)

-183(1.00)

600(1.00)

170(0.98)

190(0.02)

73(1.00)

-200(1.00)

59(1.00)

-133(1.00)

400(1.00)

-2000.00)

238(1.00)

5000.00)

2000.00)

164

SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

sample size (N> 15) to permit relatively unbaised calculation of gene frequencies.
H-values listed are those expected under Hardy-Weinberg assumptions, from
which none of the phyllostomatid samples deviate significantly. Johnson (1974)
has suggested that enzymes in key regulatory positions in metabolic pathways are
more variable than those in nonregulatory positions, and that enzymes with
variable substrates show highest heterozygostities. There is no consistent agree¬
ment of the chiropteran data with this hypothesis (Table 4). Only six of the 11
species have higher heterozygosities in regulatory enzymes than in nonregulatory
ones. We agree with Selander (1976) that Johnson’s hypothesis will not in itself
account for heterozygosity differences seen between loci. Unfortunately, John¬
son’s hypothesis does not deal with general protein loci, which we found to be the
most variable in the phyllostomatids. General proteins usually have exhibited
low levels of polymorphism in other mammals (Selander, 1976).

The data presented in Table 4 suggest that phyllostomatids differ from species
of Myotis in having lower levels of genic heterozygosity. The frequency dis¬
tributions of per locus heterozygosity (h) differ between these groups (Fig. 1).
The average locus in the phyllostomatids examined has a heterozygosity of 0.036,
whereas for Myotis this value is 0.1 17. The pattern seen among phyllostomatids
is very near to that observed in a variety of rodents (Fig. 1 ; data for esterases are
excluded from this figure). When the h-\ alues for phyllostomatids and Myotis
are compared in an Analysis of Variance (after arcsine transformation), the
difference is highly significant (P< 0.001). Phyllostomatids possess more
monomorphic loci than do species of Myotis and do not show a second frequency
peak for loci with high heterozygosity. Most of the loci contributing to this second
peak in Myotis are not in Hardy-Weinberg equilibrium (Straney et al., 1976 a).

The patterns in Fig. 1 suggest that phyllostomatids might have levels of
heterozygosity equivalent to those observed in rodents. This is not apparent when
H-values presented here are compared (average H for rodents, 0.059), because
esterases account for 43 per cent of rodent H-values (Selander, 1976). Some
Myotis populations, however, are more similar in heterozygosity levels to in¬
vertebrates (average H = 0.12; Selander, 1976), but this is not true of vespertilio-
nids as a group. Pipistrellus populations exhibit low genetic variability, and it has
been suggested that this results from demographic factors (Table 4; Straney
et al., 19766). Preliminary data on California vespertilionids indicate that other
species also have low variability (J. L. Patton, personal communication).

Levels of genic variability in phyllostomatids, and at least some species of
Myotis, differ greatly, and it is likely that other evolutionary characteristics do
as well. A number of factors could produce the differences in heterozygosity ob¬
served between phyllostomatids and Myotis-. stochastic processes, gene flow,
adaptation to microgeographic conditions, and the grain of experienced environ¬
ments (Levins, 1968; Soule, 1976). Differences, on a much lower level, also are
apparent within the phyllostomatids examined (Table 4). Artibeus, Glossophaga,
and Carollia, the most common species in our collections, differ greatly in levels
of polymorphism. Population bottlenecks, inbreeding, and drift are not suffi-

BIOLOGY OF THE PHYLLOSTOMATIDAE

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Fig. 1. — Per cent occurrence of loci with different levels of heterozygosity in rodents
(summarized in Smith et al, 1978), phyllostomatid bats (this study), and bats of the genus
Myotis (Table 4). N is number of individuals.

cient to explain the low levels of heterozygosity in Glossophaga, Anoura, and
Carollia, compared to the relatively high levels in Artibeus, because all four
genera are widespread, highly vagile, and abundant. Isolation of the population
characteristics that might be responsible for differences in heterozygosity is not
possible using genetic data alone. Only genetic studies coupled with extensive
ecological investigations will provide the information needed to address this
point, and then only if temporal trends also are examined.

Differences in heterozygosity may index more subtle differences in population
characteristics. The data presented above suggest that different species of bats
have been exposed to different evolutionary forces, which are dictated by
differences in population structure. Although we are unable at this point to
determine why variation in population structure exists or what evolutionary

BIOLOGY OF THE PHYLLOSTOMATIDAE

167

forces effect these differences, it is clear that genetic models of chiropteran pop¬
ulations must account for several distinct patterns of variation.

Future studies of ecological genetics in bats should pay particular attention to
spatiotemporal structure of populations. With proper experimental design, it is
possible to estimate deme size, effective population size, and migration rate using,
for example, Kirby’s (1976) analysis of Wright’s F-statistics. More important
than estimates of these values, though, is an estimate of their variability through
time. Bat populations are conceivably temporally unstable in composition, due
in part to their vagility and roosting habits. Turner’s (1975) studies of Desmodus
in Costa Rica indicate that vampire populations can be either ephemeral or
relatively stable depending on where the bats roost. It is important to know on
what scale this temporal variability acts as well as which ecological factors, such
as roost site, can alter its periodicity. Species differences in these parameters are
to be expected in a group as diverse as the phyllostomatids, and comparative
studies will be necessary to indicate to what degree morphological and ecological
diversity is reflected in population structure. The evolutionary process proceeds
only within the limits set by the spatiotemporal structure of the populations
involved. Hence, a useful approach to understanding patterns of population
differentiation, speciation, and phyletic evolution in different lineages is to
determine to what extent structural differences in populations determine different
evolutionary strategies. Structural parameters of populations are major deter¬
minants of the fate of new mutants, the permanence of polymorphisms, and the
speed with which adaptive change can be effected.

Genetic Phyletics

An alternative to using traditional characteristics for reconstructing the
evolutionary history of a group is to employ measures of genetic comparisons
between taxa. Because evolution can be expressed as the change in genomes
through time, genetic comparisons can be used to estimate the degree of divergence
between taxa. With the advent of biochemical assay systems this has become
possible. As genetic comparisons dependent upon breeding studies cannot be
used to compare taxa above the species level in most mammals, the early interest
in electrophoresis of bat proteins was, in part, systematic.

Manwell and Kerst (1966), Valdivieso et al. (1969), Valdivieso and Tamsitt
(1974), Tamsitt and Valdivieso (1969), and Mitchell (1970), all working with
one or at most a few proteins, concluded that electrophoretic comparisons would
be of little use in chiropteran systematics below the family level. These studies
did, however, find confirming evidence for placing the mormoopids
( Pteronotus and Mormoops ) into a family separate from phyllostomatids and
for the inclusion of the vampires as a subfamily in the Phyllostomatidae. However,
phylogenetic conclusions based on a few biochemical characters cannot
be expected to be any more accurate than those based on a few morphological
characters (Avise et al., 1974). Biochemical data used to indicate phylogenetic
relationships are based on the assumption that the loci sampled are representative

168

SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

1001-
98 -
97

LDH-I

:§

Fig. 2. — Diagramatic representation of banding patterns of four protein gene loci in seven
species of stenodermine bats.

of the genome as a whole. The magnitude of sampling error, and the resolving
power of genetic divergence estimates, is a direct function of the number of loci
examined (Nei, 1976). Thus, electrophoretic comparisons utilizing only a few
loci provide data that must be approached with caution.

It is possible that, with a small group of closely related taxa, biochemical data
for a few loci will give quite useful information. The utility of this information,

BIOLOGY OF THE PHYLLOSTOMATIDAE

169

Anoura

Fig. 3. — Wagner tree calculated from Nei’s D (X 100). Numbers are the amount of diver¬
gence between branch points and represent the minimum number of net nucleotide changes
per 100 loci accumulated along the connecting branch.

however, will depend on the sample of loci examined. Fig. 2 illustrates banding
patterns of four gene products for the seven species of stenodermine bats we have
examined from Trinidad. Although these four loci are sufficient to identify all
seven species electrophoretically, they are insufficient for calculation of genetic
distance values, because D-v alues have large errors when based on only a few
loci (Nei and Roychoudhury, 1974).

In Fig. 3 we present a phylogenetic estimate of the relationships among 14
phyllostomatid species, based on the examination of 17 gene loci. The genetic
distances between taxa, upon which this tree is based, are summarized in Table 5.
Seventeen loci certainly are only a small fraction of the phyllostomatid genome.
The sampling error associated with these divergence values is not small (Nei,
1 976), and the tree in Fig. 3 must be evaluated in this light. It also should be pointed
out that this technique overestimates similarity, and additional refinement and
the inclusion of loci such as esterases should reveal further separation of taxa.
We present these preliminary data as a starting point for additional work.

Farris’ (1972) modified Wagner algorithm for Nei’s distance was used to
construct the tree in Fig. 3. This method does not assume that evolutionary rates
are the same in all lines of descent, as does the use of an unweighted pair-group
method for constructing phenograms. The modified Wagner method partitions
the genetic distance between taxa into branch lengths of the paths connecting
them. This is done in such a way that the resulting estimates of branch lengths are

170

SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

minimum estimates of the amount of change between cladistic events. Because the
tree is based on Nei’s D, the branch lengths can also be interpreted as the minimum
number of net codon changes per locus since a particular cladistic event. Thus,
according to Fig. 3, Artibeus cinereus and Vampyrops share an immediate
common ancestor. Since the cladistic splitting of the two, we estimate that
Vampyrops has accumulated 22 net codon changes per 100 loci, whereas A.
cinereus has accumulated a minimum of three. Because these taxa share a common
ancestor, the difference in divergence is also a difference in evolutionary rate
along the two branches. One of the striking characteristics of the tree in Fig. 3 is
that the branch lengths are unequal, implying that the rates of evolution have not
been the same in all lines of descent. This is consistent with the argument given
above that differences in levels of genetic variability within phyllostomatid species
mirror underlying differences in population structure, thereby differentially
affecting evolutionary potential.

The root in Fig. 3 has been placed using Farris’ (1972) minimum variance
criterion. This is an iterative procedure whereby the root is placed in the position
that minimizes the variance in divergences of terminal taxa from the hypothetical
ancestor of the group as a whole. There are three major lineages apparent when
the root is placed: 1) stenodermines, 2) Phyllostomus and Carollia, and 3) glos-
sophagines, Desmodus, and Sturnira. The average divergence of these three
lineages from the ancestor is similar (mean, 86, 76, and 85 codon changes per
100 loci, respectively). An analysis of variance of within and between lineage
effects on divergence indicates that 100 per cent of the variance in divergence
present in Fig. 3 is within lineages. As we can demonstrate no differences in
evolutionary rate between lineages, we can use the average divergence of the
lineages (82 codon changes per 100 loci) to estimate the age of the family. Nei’s
D is a linear function of time (Nei, 1976), and studies by Avise and Ayala
(1975, 1976) indicate that genetic distance is by and large independent of
cladistic history. Sarich (1977) has calibrated Nei’s D against his albumin clock
estimate of divergence time and has provided us with the conversion equation
1.0 D— 28 million years (for branch length, 1.00 = 56 million years). Using this
conversion, we estimate that the diversification of the family occurred 40 mil¬
lion years ago during the early Oligocene. Because this is a minimum estimate of
age, the age estimated is of diversification not origin, and the estimate is not
without sampling error, we feel that these data are comparable with Koopman’s
(1976) and Smith’s (1976) conclusion that the late Oligocene is the latest that
the family could have arisen.

Within the error of our estimates, the lineages represented in Fig. 3 appear
to have arisen at the same time. These lineages are not well defined, except for the
relatively compact stenodermine lineage, and there is no evidence of a “ Macrotus-
like” and “ Phy l lostom ws-like” (Smith, 1976) dichotomy within our sample.
Genera hypothesized as belonging to one lineage or the other are intermixed in
Fig. 3 (compare Smith, 1976, fig. 2). Even though our inability to distinguish
this dichotomy may be an artifact of sampling, we think it best to assume that the
major adaptive trends within the family are of coeval origin.

BIOLOGY OF THE PHYLLOSTOMATIDAE

171

Stenodermines

The discreteness of the stenodermine lineage in Fig. 3 probably results from
more extensive sampling of members of this subfamily. The radiation of this
group appears to be an early one, the line leading to Ametrida diverged perhaps
20 million years ago in the late Miocene. Artibeus is a basal taxon for the rest
of the subfamily represented here and two separate lineages derive from it. The
three species of Artibeus have undergone little divergence from their respective
common ancestors whereas the two lineages involving Vampyrops and Uroderma-
Chiroderma have evolved at a much faster rate. These results suggest that
Artibeus is a paraphyletic taxon.

With effort, it is possible to identify Smith’s (1976) “short-faced, long-faced”
dichotomy in our phylogram. The “long-faced” lineage is polyphyletic in our
reconstruction although the three members of this group ( Vampyrops, Uroderma,
and Chiroderma ) are derived from a single genus, Artibeus. Furthermore, our
phylogenetic hypothesis suggests that short-faced is the primitive condition for
stenodermines. We have examined too few genera to be certain of this point,
but the data at hand indicate that long faces represent parallel derived characters.

Our sample of stenodermine taxa, however, is sufficient to suggest a polarity
for Baker’s (1973) phylogeny of the subfamily based on gross karyotypic
characters. His fig. 5 is quite similar to our Fig. 3 if the root of his phylogram is
displaced to the right and if one ignores the absence of Sturnira. Karyotypically,
Chiroderma and Uroderma are not related as closely to each other as elec¬
trophoretic data indicate; further study could identify additional areas of dis¬
agreement. It is, however, reassuring to find the same basic phylogenetic frame¬
work emerging from two different and independent data sources.

Phyllostomus and Carollia

There is little that can be said of the association of Carollia and Phyllostomus
presented in Fig. 3. These two genera are not closely related but probably do
represent a distinct lineage within the family. Walton and Walton (1968) suggested
a similar relationship based on their study of postcranial osteology. There is no
indication in our data of close phylogenetic ties between Carollia and Glossophaga
(sensu Smith, 1976).

The divergence of the two species of Phyllostomus appears to have occurred 8
million years ago during the mid-Pliocene (D=0.29). The morphological and
ecological differences between P. hastatus and P. discolor are much greater than
those between the two species of Macrotus studied by Greenbaum and Baker
(1976), even though the latter are separated by a greater genetic distance (D =
0.41-0.50). This represents another of the growing number of cases where genetic
and morphological measures of divergence are found to be discordant (King
and Wilson, 1975; Avise, 1976).

Glossophagines, Desmodus, and Sturnira

This group forms the most heterogeneous branch of our phylogenetic tree, and
the relationships within it are difficult to reconcile with morphological evidence

172

SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

and current concepts of phyllostomatid systematics. Anoura and Glossophaga are
somewhat closely related, based on electrophoretic data, although this association
is overshadowed by the greater amount of protein evolution along the Anoura
branch. Our placement of Sturnira is at variance with current taxonomic opinion.
Walton and Walton (1968) postulated a relationship between Sturnira and the
glossophagines, following a comparison of postcranial morphology. Addition
of more genera to this data set would not result in a closer association of Sturnira
and the stenodermines because additional data would not decrease the large
genetic distances between these groups (Table 5). Based on our electrophoretic
sample, we are left with the conclusion that Sturnira is not genetically a stenoder-
mine bat and is not closely related to any one of the lineages represented in this
study.

A close relationship between Desmodus and the glossophagines, based on
chromosomal, immunological, and sperm morphology data, was proposed by
Forman et al. (1968). Our data also suggest such a relationship between Glos¬
sophaga and Desmodus (Fig. 3; Table 5). Because of the difference in evolution¬
ary rates along the two branches, it is difficult to estimate the age of this diver¬
gence, but we suggest that it is 10 million years. This is consistent with the fossil
record to the extent that fossil desmodontines are not known prior to about 1.5
million years bp (Hutchison, 1967).

An overview of the genic and morphological data from this family suggests
that there are several examples where there is discordance in the rates of evolu¬
tion of genic and classical morphological characters. One hypothesis that at¬
tempts to reconcile genetic and morphological data assumes that the morphologi¬
cal modifications leading to a specialized taxon have been due to changes in
regulatory genes affecting developmental pathways. Such changes, which one
would not expect to be reflected in the structural genes assayed in electrophoresis,
could result in major and rapid morphological evolution. This form of quantum
evolution ( sensu Simpson, 1953) has recently been invoked by King and Wilson
(1975) to explain the small genetic distance between Homo and Pan. If this hypo¬
thesis reflects the true path of evolution followed in these discordant examples,
we would predict, following King and Wilson (1975), that DNA hybridization
between such taxa would show similarity in the unique DNA fraction consistent
with that found electrophoretically and a larger difference in the presumably
regulatory medium repeated DNA fraction.

Phylogenetic reconstruction is as much a science as it has been portrayed an
art. One proceeds by constructing hypotheses of relationships from different
data sources and searching for one that subsumes the others and provides an
explanation of their differences. This consistent hypothesis is accepted as “true”
either until a more general one is produced or conflicting data are found. The
phylogenetic hypotheses of Smith (1976) and those reflected by the checklist of
Jones and Carter (1976) are not in accordance with the genetic relationships
indicated by our electrophoretic data. We do not view these electrophoretic
results as a procrustean bed of truth into which the morphological evidence
must be forced in agreement. Rather, they generate a phylogenetic hypothesis

Table 5. — Nei’s genetic distance (D, upper half matrix) and Rogers’ genetic similarity (S, lower half matrix) for hat populations from Trinidad.
Where more than one population of a species is listed, numerical designations are as in Table 3. I indicates an infinite value for D ( Nei's genetic

identity 1=0.00).

BIOLOGY OF THE PHYLLOSTOMATIDAE

173

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sufficiently different from others that have been proposed to indicate that “the
great deal of uncertainty and contradictory evidence” (Smith, 1976) surrounding
phyllostomatid phylogency will continue in the future. We still lack an hypothesis
of the phylogeny of the Phyllostomatidae that is consistent with available data
and that also identifies the evolutionary processes producing the differences
between morphological and genetic findings.

[Note added in galley. — Additional work by us suggests that the distance we
report between Glossophaga and Desmodus is too low. Examination of new ma¬
terial, both at Lubbock and Berkeley, shows that Desmodus and Glossophaga
share very few alleles. ]

Conclusions

Biochemical genetics has proven valuable in evolutionary biology through the
characterization of population structure in space and time and through generation
of phylogenetic hypotheses. By examining the genetic structure of populations,
important evolutionary parameters can be identified and quantified to provide a
bridge between genetic phylogenies and more traditional evolutionary recon¬
structions. The study of chiropteran genetics is only 10 years old; yet, in that time
it has provided information that both challenges and supports the traditional view
of chiropteran evolution. The dynamics of population structure of vespertilionid
and phyllostomatid bats does not appear to be the same, although studies of
temporal structure will be necessary to confirm this conclusion. The mode of
evolution, as reflected by electrophoretic parameters, appears to be different
between some lineages of phyllostomatids, particularly the desmodontines. When
more genetic data are available on phyllostomatid bats, an integration of genetic,
karyotypic, and morphological data should produce a consistent model of
evolution in this group, which might be surprising in its complexity.

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of neotropical bat hemoglobins. Comp. Biochem. Physiol., 30:117-122.

Walton, D. W., and G. W. Walton. 1968. Comparative osteology of the pelvic and
pectoral girdles of the Phyllostomatidae (Chiroptera: Mammalia). J. Grad.
Res. Center, Southern Methodist Univ., 37:1-35.

SPERM MORPHOLOGY

G. Lawrence Forman and Hugh H. Genoways

Bishop and Austin (1957) in their study of variation in mammalian spermatozoa
suggested that the sperm of each mammalian species was probably unique.
Although complete volumes have been written on the ultrastructure of spermatozoa
(for example Baccetti, 1970), particularly of humans and domestic animals,
there is still relatively little information available on the comparative gross
morphology of spermatozoa. McFarlane (1963), Forman (1968), and Forman
et al. (1968) made significant contributions to our understanding of the use of
sperm morphology in establishing systematic and phylogenetic relationships of
birds and mammals. However, there have been very few similar studies published
to this date.

The use of sperm morphology as a systematic character among mammals, is
relatively new, beginning with the study of British murid rodents by Friend (1936)
Other studies dealing with rodent sperm include those of Braden (1959), Hirth
(1960), Wooley and Beaty (1967), Genoways (1973), Helm and Bowers (1973),
and Linzey and Layne (1974). Hughes (1964, 1965) compared the morphology
of sperm of 18 species of marsupials representing five families, and Biggers and
Delamater (1965) and Biggers (1966) reported on the spermatozoa of several
genera of American marsupials. Griffiths (1968) presented data on the sperm
of the echidna and Bedford (1967) reported observations on the fine structure
of the spermatozoa of two primates in addition to man. An especially important
contribution is that of Martin et al. (1975). They used scanning electron
microscopy to compare spermatozoa of 16 species of primates representing
four families and concluded that sperm morphology might be valuable in gaining
better understanding of intrageneric relationships among primates.

Six studies have described the sperm of Rhinolophus ferrumequinum, and
Hirth (1960), Fawcett and Ito (1965), Wimsatt et al. (1966), and Forman
(1968) reported on various aspects of the spermatozoa of species of vespertilionid
bats. Forman (1968) was the first to present information on the sperm of members
of the family Phyllostomatidae. In his study, he presented information on eight
species representing four of the six subfamilies. In the same year, Forman et al.
(1968) reported on two additional phyllostomatid species, Desmodus rotundus
and Diphylla ecaudata, of a fifth subfamily, the Desmodontinae.

Over the past seven years, we have accumulated data on the sperm of
phyllostomatid bats in the course of several other studies of this family. This has
resulted in material for 35 species, 28 of which have not been studied previously.
Through new staining techniques, we also have been able to acquire new
information on the seven species for which some data were presented previously.
The results of our studies and their systematic implications are discussed below.

177

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Methods and Materials

The spermatozoa of 35 species belonging to six subfamilies of phyllostomatids
were examined. To obtain spermatozoa, the epididymides of freshly-killed bats
were removed. A small amount of fluid containing mature sperm was taken and
suspended in an isotonic solution of sodium citrate. A few drops of the sodium
citrate and spermatozoon solution were placed on a microscope slide and allowed
to air-dry. Dilution of the spermatozoa with sodium citrate was necessary so that
individual spermatozoa would be dispersed for study and photographing.
Spermatozoa on slides were fixed with a solution of one part acetic acid and
four parts absolute methyl alcohol. Slides were allowed to fix for 10 to 15 seconds
and then shaken dry. Fixing for a longer period resulted in destruction of the
acrosome.

Slides were stained with Toluidine Blue O and counterstained with PAS.
Counterstaining resulted in delineation of the acrosomal material so that the
outline of the headcap could be observed. The procedure outlined below was
followed in staining slides:

1. fix in solution of acetic acid and methyl alcohol;

2. rinse three times in distilled water;

3. place in 15% Periodic Acid for 10 minutes;

4. rinse in tap water for 10 minutes;

5. rinse briefly in distilled water;

6. place in Schiffs' Reagent for 10 minutes;

7. rinse in metabisulfite with three changes at three minutes each;

8. rinse in tap water for 5 minutes;

9. rinse briefly in distilled water;

10. place in .02% Toluidine Blue O for 30 minutes;

1 1. place in acetone for 2 minutes;

12. place in solution of acetone plus xylene (1:1) for 2 minutes;

13. place in xylene for two changes at 3 minutes each;

14. mount using cover slip and Permount.

The following characters were measured: total length of head, length of
acrosome, nuclear length, head width, midpiece length. The mean, range (in
parentheses), and one standard deviation for the aforementioned characters are
given beyond in the species descriptions whenever possible. Measurements were
taken by means of a Unitron Filar widefield dial micrometer attached to an AO
microstar Series 10 research microscope. Measurements are given in microns.

The terms dorsal and ventral refer to the flattened surfaces of the head and
midpiece, whereas lateral refers to the narrow sides of the sperm. Length of
head included both the acrosome and nuclear area. Width of the head was measured
as the distance between extremities when observed in dorsal or lateral view. The
tails of sperm were not considered in this study.

Characters considered in this study included: shape of head; shape of apices of
acrosome and nucleus; shape of base of head; symmetry of acrosome and head;
length of acrosome as compared with nucleus; location of posterior edge of
acrosome; placement of the attachment of the neck and midpiece to head; relative
amount of acrosome anterior to nucleus; thickness, relative length, and degree

BIOLOGY OF THE PHYLLOSTOMATIDAE

179

Table 1. — Calculated ratios comparing the dimensions of the spermatozoa of 35 species of

phyllostomatid hats.

Midpiece

length/

head

Species length

Head

length/

head

width

Head

length/

acrosome

length

Midpiece

length/

acrosome

length

Nuclear

length/

head

width

Midpiece

length/

nuclear

length

Head

length/

nuclear

length

Nuclear

length/

acrosome

length

Micronycteris megalotis

1.91

1.53

1.78

3.41

1.19

2.45

1.28

1.40

Micronycteris nicefori

2.01

1.18

1.71

3.44

1.11

2.13

1.06

1.62

Macrotus waterhousii

2.00

1.29

1.50

3.00

0.82

3.12

1.56

0.96

Tonatia bidens

2.44

1.56

1.25

3.05

1.25

Mimon crenulatum

1.66

1.38

1.62

2.69

1.03

2.23

1.35

1.20

Phyllostomus discolor

1.73

1.46

1.67

2.89

1.12

2.25

1.30

1.28

Glossophaga soricina

2.12

1.19

2.53

0.90

2.83

1.33

0.90

Anoura geoffroyi

1.44

1.28

1.82

2.62

0.98

1.89

1.31

1.39

Choeronycteris mexicana

Carollia brevicauda

1.51

1.46

1.48

2.24

0.97

2.27

1.50

0.99

Carollia perspicillata

1.63

1.56

1.59

2.60

1.07

2.39

1.46

1.09

Sturnira lilium

1.92

1.65

1.71

3.27

1.17

2.71

1.41

1.21

Sturnira tildae

1.81

1.59

1.73

3.13

1.27

2.26

1.25

1.38

Uroderma bilobatum

1.86

1.48

2.30

4.29

1.08

2.56

1.37

1.68

Vampyrops helleri

1.72

1.62

1.60

2.76

1.24

2.26

1.31

1.22

Vampyrodes caraccioli

1.69

1.64

1.76

2.98

1.25

2.21

1.31

1.35

Chiroderma improvisum

Chiroderma trinitatum

1.82

1.39

1.62

2.95

1.18

2.23

1.23

1.32

Mesophylla macconnelli

1.65

1.36

1.63

2.69

1.03

2.19

1.32

1.23

Artibeus cinereus

1.94

1.29

1.42

2.75

1.07

2.35

1.21

1.17

Artibeus toltecus

1.75

1.57

1.66

2.91

1.23

2.23

1.28

1.30

Artibues jamaicensis'

1.94

1.36

1.64

3.17

1.09

2.42

1.25

1.31

Artibeus lituratus

1.73

1.48

1.45

2.51

1.11

2.30

1.33

1.09

Ardops nichollsi

2.09

1.35

1.76

3.67

1.03

2.76

1.32

1.33

Phillops haitiensis

1.79

1.45

1.73

3.11

1.11

2.34

1.30

1.33

Ariteus flavescens

1.97

1.41

1.63

3.21

1.04

2.73

1.36

1.04

Stenoderma rufum

1.86

1.43

1.57

2.91

1.08

2.46

1.32

1.08

Centurio senex

1.72

1.22

1.66

2.85

1.01

2.07

1.21

1.37

Brachyphylla cavernarum

Erophylla bombifrons

1.44

1.42

1.49

2.15

1.09

1.88

1.30

1.14

Erophylla sezekorni

1.59

1.70

2.80

1.15

2.19

1.38

1.23

Phyllonycteris poeyi

1.34

1.40

1.55

2.09

1.03

1.82

1.35

1.15

Desmodus rotundus

2.47

1.74

1.58

3.91

1.42

3.03

1.23

1.29

Diaemus youngii

2.23

1.80

1.75

3.91

1.45

2.78

1.23

1.41

Diphylla ecaudata

2.10

1.32

1.58

3.32

2.09

1.00

1.58

of tapering of midpiece. Table 1 gives statistical ratios based on measurements
taken. Figs. 1-5 compare the total head length, nuclear length, and midpiece
length of the species studied. Voucher specimens are deposited in The Museum
of Texas Tech University (TTU) and Carnegie Museum of Natural History (CM).
Most specimens were collected under a grant from the National Science Foundation
(GB-41 105) to Robert J. Baker and Hugh H. Genoways.

Accounts of Species
Subfamily Phyllostomatinae
Micronycteris megalotis (Gray, 1 842)

Description (Fig. 1A). — Head oval, rear portion tapered slightly but con¬
siderably more than that of Macrotus\ bilaterally symmetrical; apex narrowly
rounded; acrosome no wider than nucleus; base slightly convex; nuclear portion

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PH YLLOSTOMATINAE

Fig. 1. — Sperm of six phyllostomatine bats. A) Micronycteris megalot is", B) Micronycteris
nicefori; C) Macrotus waterhousii', D) Tonatia b ideas', E) Mimon crenulatum', F) Phyllostomus
discolor. Scale equals 5 microns.

has blunt apex, more rounded than that of acrosome; acrosome longer than nucleus
and constituting a substantial portion of the head length; head length 4.46(4.19-
4.65) ±0.138, 4.87(4.56-5.12) ±0.237, acrosome length 3.00(2.79-3.07)

±0.102, 2.73(2.42-2.98) ±0.188, nuclear length 3.65(3.44-3.91) ±0.160,
3.81(3.17-3.19) ±0.072, head width 2.92(2.79-3.07) ±0.088, 3.19(3.07-3.35)
±0.091. Neck short, joins head midway at base of head. Midpiece extremely
thin, relatively long, length 9.45(9. 1 1 -9.95) ± 0.286, 9.32(8.84-9.58) ± 0.25 1 .

Remarks. — Morphology of the sperm head of Micronycteris megalotis is
substantially different from that of Macrotus waterhousii, with the sperm head
of M. megalotis considerably narrower than that of M. waterhousii.

BIOLOGY OF THE PHYLLOSTOMATIDAE

181

Specimens examined. — Trinidad: Blanchisseuse, St. George, 1 (TTU 23754); Maracas,
St. George, 1 (TTU 23759).

Micronycteris nicefori Sanborn, 1949

Description (Fig. IB). — Head wider than that of M. megalotis, more rounded;
bilaterally symmetrical; base flattened, not convex; acrosome substantially
shorter than nucleus, in sharp contrast to condition found in M. megalotis; nuclear
portion extremely rounded; apex of acrosome and nucleus similar in shape; head
length, 4.00(3.72-4.37) ±0.299, acrosome length 2.34(2.23-2.60) ±0.145,
nuclear length 3.78(3.62-3.91) ±0.092, head width 3.40(2.98-3.72) ±0.177.
Neck short, not joining head midway along base. Midpiece extremely narrow,
difficult to distinguish from tail; length 8.04(7.91-8.18) ±0.1 15.

Remarks. — Morphology of the spermatozoa of M. nicefori is similar to that
of M. megalotis but does differ in several ways. Most noticeably, the acrosome
is shorter than the nucleus in M. nicefori but longer than the nucleus in M.
megalotis. M. nicefori also has a wider sperm head than megalotis and a flattened
rather than convex base of head.

Specimen examined. — Trinidad: 2 mi. N, 2 mi. E Valencia, St. Andrew, 1 (TTU 23768).

Macrotus waterhousii Gray, 1843

Description (Fig. 1C). — Head not rounded, triangular; bilaterally symmetrical;
base strongly convex; apex of acrosome broadly rounded, bullet-shaped; posterior
border of acrosome sharply defined; acrosome no wider than nucleus and similar
in length; nuclear portion small, with extremely blunt apex, and more rounded than
apex of acrosome; head length 3.73(3.53-4.00) ±0.150, 3.67(3.44-3.81) ±0.100,
nuclear length 2.39(2.32-2.70) ±0.132, 2.49(2.32-2.79) ±0.156, acrosome
length 2.49(2.32-2.70) ±0.178, 2.23(2. 14-2.32) ±0.068, head width 2.90(2.70-
3.16) ±0.156, 2.95(2.79-3.07) ±0.1 12. Neck short, joining head midway at
base of head. Midpiece extremely short; demarcation with tail distinctive; length
7.46(7.34-7.63) ±0.1 12, 7.66(7.16-7.91) ±0.183.

Remarks. — The form of the sperm head in this species is unique with no
comparable conformation found in any other genus. Also of interest is the ex¬
tremely short midpiece.

Specimens examined. — Jamaica: Green Grotto, 2 mi. E Discovery Bay, St. Ann Parish,
3 (TTU 21501-02, 21504).

Tonatia bidens (Spix, 1 823)

Description (Fig. ID). — Head rounded to broadly oval; acrosome can con¬
tribute markedly to total length of head; acrosome bilaterally symmetrical, rear
terminus only slightly beyond apex of nucleus; apex of acrosome broadly rounded
but less so than nucleus; acrosome considerably shorter than nucleus and never
wider than nucleus; nucleus rounded, with extremely blunt apex; base of head
concave; head length approximately 4.64(4.46-4.84), nuclear length 3.72(3.58-
4.00), head width 2.98(2.88-3.07). Neck relatively long and slightly off center of

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point of attachment to head. Midpiece relatively long, anterior portion broad,
tapering sharply posteriorly; length 1 1.36(9.49-1 1.25).

Remarks. — Among the phyllostomatines, the head of the spermatozoon
of T. bidens is most similar in general shape (acrosome and nucleus) to
Micronycteris nicefori and Phyllostomus discolor.

Specimen examined. — Trinidad: 2 mi. N, 2 mi. E Valencia, St. Andrew, 1 (TTU 23794).

Minion crenulatum (E. Geoffroy St.-Hilaire, 1810)

Description (Fig. IE). — Head bluntly rounded; acrosome keel-shaped, ex¬
tremely asymmetrical; acrosome slightly broader, at widest point, than nucleus;
acrosome terminates posteriorly about midway along length of nucleus, adding
about 25 per cent to length of head; nucleus slightly longer than acrosome; nuclear
portion extremely rounded, apex narrowly rounded terminating in broad point;
base of nucleus rounded but slightly concave; head length 5.42(5. 1 2-5.86) ± 0. 1 94,
acrosome length 3.34(3. 16-3.53) ±0.1 34; nuclear length 4.02(3.91-4.09) ±0.068,
head width 3.92(3.72-4.09) ±0.1 19. Neck short with attachment to head slightly
off center. Midpiece of moderate breadth anteriorly; moderate length; length
8.98(8.56-9.39) ±0.213.

Remarks. — The sperm head of Mimon differs in general morphology from both
Macrotus and Micronycteris and is exceptionally large. The asymmetry of the
acrosome is in striking contrast to the generally symmetrical acrosome of other
phyllostomatines.

Specimen examined. — Trinidad: 2 mi. E San Rafael, St. George, 1 (TTU 23770).
Phyllostomus discolor (Wagner, 1 843)

Description (Fig. IF). — Head narrowly rounded; acrosome only slightly
asymmetrical, shorter than nucleus, and terminating posteriorly about half-way
along length of nucleus; acrosome slightly wider, at widest point, than is nucleus;
nucleus triangular in shape with broad base, apex narrowly rounded, pointed; base
of nucleus slightly concave; head length 5.19(4.93-5.58) ±0.239, acrosome length
3.1 1(2.79-3.44) ±0.240, nuclear length 3.99(3.53-4.37) ±0.230, head width
3.55(3.26-3.72) ±0.159. Neck extremely short, junction with head considerably
off center; joins head on same side as most distinct portions of the apex of the
acrosome. Midpiece of moderate length, thin, tapering gradually to distinctive
junction with tail; length 8.98(8.56-9.58) ±0.316.

Remarks. — The head of the spermatozoon of Phyllostomus discolor has
morphological similarities with both Mimon and Micronycteris but is identical
to neither; the head is most similar to that of M. nicefori except that the acrosome
is slightly asymmetrical. The nucleus is narrower than in Mimon with broad,
triangular base as in M. nicefori.

Previous study. — Two specimens from Nicaragua (Forman, 1968:905).

Specimen examined. — Trinidad: Las Cuevas, St. George, 1 (TTU 23777).

BIOLOGY OF THE PHYLLOSTOMATIDAE

183

Subfamily Glossophaginae
Glossophaga soricina (Pallas, 1766)

Description (Fig. 2A). — Head extremely small, short, and quite rounded;
base of head broad giving a shovellike shape; base has well-developed concavity;
apex of acrosome nearly symmetrical, being somewhat more narrowly rounded
than the broadly rounded apex of nucleus; acrosome nearly as long as nucleus;
posterior limit of acrosome considerably behind midpoint of nucleus; only a small
portion of acrosome occurs anterior to nucleus; acrosome never wider than nucleus;
head length 3.80(3.53-4.00) ±0.162, acrosome length 3. 19(3.09-3.26) ±0.202,
nuclear length 2.86(2.70-3.26) ±0.268, head width 3.19(3.07-3.26) ±0.091.
Neck moderate in length, junction with head only slightly off center. Midpiece
extremely broad, tapering gradually posteriorly; junction with tail quite distinctive;
length 8.08(7.63-8.46) ±0.316.

Remarks. — Sperm morphology in this species is notably similar to that of
Anoura; heads are extremely small compared to those of most other species.

Previous study. — Four specimens from Chiapas (Forman, 1968).

Specimens examined. — Veracruz: 4 km. W, 5 km. S Sontecomapa, 1 (TTU 28900);
Yucatan: Merida, 1.

Anoura cultrata Handley, 1960

Description (after Forman, 1968). — Head rounded, its breadth approximately
seven-eighths of length, broadest in basal region, bluntly rounded at apex; base
slightly concave (the acrosome was not examined in the previous study). Neck not
observed. Midpiece short when compared to length of tail; width uniform through¬
out.

Remarks. — The spermatozoa of Anoura cultrata are distinct from those of
Glossophaga soricina. The head is broader in A. cultrata than in G. soricina, the
ratio of length to breadth being 1.15 as opposed to 1.28 in G. soricina (Forman,
1968).

Previous study. — Two specimens from Panama (Forman, 1968).

Anoura geoffroyi Gray, 1838

Description (Fig. 2B, 2C). — Head quite rounded; base slightly convex;
acrosome slightly asymmetrical, with apex occasionally somewhat pointed;
acrosome shorter than nucleus and contributing markedly to total head length;
acrosome only slightly broader than nucleus at widest point; apices of acrosome
and nucleus usually broadly rounded, that of the nucleus particularly so; head
length 3.92(3.53-4.09) ±0.184, 4.05(3.91-4.37) ±0.151, acrosome length 2.23
(2.05-2.32) ±0.09, 2.23(2.05-2.42) ±0.1 16, nuclear length 3.08(2.79-3.44) ±
0.216, 3.09(2.88-3.35) ±0.165, head width 3.14(2.88-3.26) ±0.128, 3.16(2.98-
3.35) ±0.104. Neck of moderate length, junction with head slightly off center;
attachment to head on same side as longest portion of acrosome. Midpiece ex-

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SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

GLOSSOPHAGINAE

CAROLLI IN AE

Fig. 2. — Sperm of some glossophagine
B-C) Anoura geoffroyr, D) Choeronvcteris
perspicilhita. Scale equals 5 microns.

and carol liine bats. A) Glossophaga soricina, ;
mexicana; E) Carollia brevicauda ; F) Carollici

BIOLOGY OF THE PHYLLOSTOMATIDAE

185

tremely wide at anterior end, tapering abruptly towards posterior end; junction
with tail distinctive; length 5.57(4.93-6.14) ±0.358, 5.84(5.58-6.05) ±0.149.

Remarks. — The spermatazoon of this species is quite similar to that of
Glossophaga soricina , the only species of the genus examined.

Specimens examined. — Hidalgo: 13 km. WSW Tehuetlan, 2 (TTU 15477-78). Trinidad:
2 mi. N, 2 mi. E Valencia, St. Andrew, 1 (TTU 23802); Las Cuevas, St. George, 1 (TTU
23798).

Choeronycteris mexicana Tschudi, 1844

Description (Fig. 2D). — Head oval, somewhat triangular or shovel shaped;
extremely large (in length and breadth); acrosome symmetrical, relatively long,
posterior terminus well posterior to midpoint of head, and apex broadly rounded;
acrosome difficult to distinguish from nucleus, blending in at the sides of the head;
acrosome adds only slightly to total length of head; nucleus extremely rounded,
apex rounded; base concave, corners rounded; head length 5.09(4.74-5.58) ±
0.259, acrosome length 3.37(3.26-3.44) ±0.089, nuclear length 4.26(4.00-4.46)
±0.158, head width 3.99(3.62-4. 19) ±0. 145. Neck short, attached to base of
head nearly at its midpoint. Midpiece narrow, moderate length, tapering only
slightly posteriorly; length 8.59(8.37-9.02) ±0.182.

Remarks. — Spermatozoa from Choeronycteris mexicana are easily dis¬
tinguishable by their larger size from those of other glossophagines. Glossoph-
agines examined to date appear relatively consistent and uniform in sperm mor¬
phology.

Specimen examined. — Tlaxcala: 5 km. E, 3 km. N Tlaxcala, 1 (TTU 25347).

Subfamily Carolliinae
Carollia castanea H. Allen, 1890

Description (after Forman, 1968:909). — Head rounded, somewhat heart-
shaped; apex broadly rounded; base concave and symmetrical, narrowing
laterally at point of junction with neck (acrosome not observed in this study).
Neck short but distinct; junction with head near center of base. Midpiece short,
anterior end at distinct angle to base of head, tapering only slightly posteriorly.

Remarks. — A spiraled midpiece was observed in this species, confirming the
existence of such a structure in at least one member of the Phyllostomatidae
(Forman, 1968).

Previous study. — Three specimens from Panama (Forman, 1968).

Carollia brevicauda (Schinz, 1821)

Description (Fig. 2E). — Head rounded; acrosome long, posterior border located
from midway to two-thirds back along the length of the nucleus; acrosome slightly
asymmetrical and terminating in broadly rounded apex; acrosome extremely large
and longer than nucleus, possibly somewhat wider than nucleus at its widest
point; nucleus rounded with broadly rounded apex; base of head slightly concave;

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head length 5.22(4.84-5.49) ±0.180, acrosome length 3.53(3.26-3.81) ±0.167,
nuclear length 3.48(3.26-3.72) ±0. 1 1 8, head width 3.58(3.44-3.81) ±0.135.
Neck of moderate length; attachment to head off center, with attachment on same
side as longest portion of acrosome. Midpiece narrow, moderate length, tapering
gradually to posterior; junction with tail distinctive; length 7.90(7.53-8.28) ±
0.208.

Remarks. — Overall shape of the sperm head in C. brevicauda is more rounded,
wider, and generally greater in size than that of C. perspicillata. C. brevicauda
shares several characteristics with C. perspicillata , including an acrosome that is
often longer than the nucleus and a nucleus that is rounded with a broadly rounded
apex.

Specimen examined. — Veracruz: 4 km. W, 5 km. S Sontecomapa, 1 (TTU 28901).
Carollia perspicillata (Linnaeus, 1758)

Description (Fig. 2F). — Head relatively narrow (because significant amount
of acrosome is anterior to apex of nucleus; portion of acrosome anterior to apex
of nucleus may exceed 30 per cent of total length of head); acrosome slightly
asymmetrical, as long as or slightly longer than the nucleus in many cases;
acrosome terminates posteriorly about 40 to 50 per cent of way back along the
length of the nucleus; acrosome only slightly wider than nucleus at its widest point;
nucleus rounded, base concave, and apex broadly rounded; head length 5.23
(5.02-5.39) ±0.103, acrosome length, 3.29(3.07-3.53) ±0. 148, nuclear length
3.58(3.26-3.81) ±0.201, head width 3.35(3.16-3.53) ±0.131. Neck short,
attached to base of head slightly off center. Midpiece of moderate length, gradually
tapering; junction with tail distinctive; length 8.55(8.1 8-9. 1 1 ) ± 0.28 1 .

Remarks. — Morphology of the spermatozoon of Carollia perspicillata resembles
that of Micronycteris megalotis, but the head differs in several respects from that
of C. brevicauda. Large sperm heads might be characteristic of the genus
Carollia.

Specimens examined. — Quintana Roo: 14 km. NE Playa del Carmen, 1 (TTU 18421);
Trinidad: Blanchisseuse, St. George, 1 (TTU 23859).

Subfamily Stenoderminae
Sturnira lilium (E. Geoffroy St. -Hilaire, 1810)

Description (Fig. 3A). — Head large, relatively narrow oval; acrosome
symmetrical, shorter than nucleus; acrosome large, terminating anteriorly in
moderately rounded apex and posteriorly about halfway along length of nucleus;
distinctive portion of acrosome lies anterior to nucleus; acrosome may be narrower
at base than nucleus at its widest point or they may be of equal breadth; nucleus
oval, apex more broadly rounded than that of the acrosome; base extremely narrow
(relative to greatest breadth of nucleus) and concave; head length 5.15(4.93-
5.49) ±0.179, acrosome length 3.02(2.70-3.16) ±0.150, nuclear length 3.64
(3.44-4.00) ±0.158; head width 3. 12(2.98-3.26) ±0.085. Neck moderate in

BIOLOGY OF THE PHYLLOSTOMATIDAE

187

length, attached to head slightly off center. Midpiece long, stains dark; broad at
anterior end, sharply tapering posteriorly; junction with tail distinctive; length
9.87(9.39-10.14) ±0.224.

Remarks. — The overall similarity of sperm from Sturnira lilium to that found
in other stenodermines supports the inclusion of this genus within the subfamily.

Previous study. — Two specimens from Chiapas (Forman, 1968).

Specimens examined. — Trinidad: 2 mi. N, 2 mi. E Valencia, St. Andrew, 1 (TTU 23901);
Blanchisseuse, St. George, 1 (TTU 23899).

Sturnira tildae de la Torre, 1959

Description (Fig. 3B). — Head similar in structure to S. lilium but differs from
it in several ways; base of head less concave than that of S. lilium and sometimes
lacking concavity; apex of acrosome symmetrical, as much as half of the acrosome
occurring anterior to nucleus; acrosome covers only a very small portion of the
nucleus; nucleus ovoid; head length 4.81(4.56-5.02) ±0.121, 4.82(4.65-4.93)
±0.151, acrosome length 2.78(2.51-3.07) ±0.149, 2.43(2.23-2.70) ±0.177,
nuclear length 3.85(3.62-4.37) ±0. 1 86, 3.78(3.44-4.09) ±0.237, head width
3.02(2.88-3.26) ±0.136, 3.00(2.79-3.35) ±0.162. Neck relatively long, attached
to middle of base. Midpiece slightly shorter than that of S. lilium, extremely
narrow, and tapering slightly posteriorly; length 8.71(8.28-9.1 1) ±0.250,
8.81(8.37-9.02) ±0.293.

Remarks. — Spermatazoa of Sturnira tildae differ from those of species in this
genus mainly in that base of head is less concave and midpiece shorter. The small
acrosome may be unique to S. tildae, but that possibility awaits examination of
the acrosome of Sturnira ludovici. The nucleus is similar in configuration to that
of Artibeus cine reus.

Specimens examined. — Trinidad: 2 mi. N, 2 mi. E Valencia, St. Andrew, 1 (TTU 23907);
Blanchisseuse, St. George, 1 (TTU 23904).

Sturnira ludovici Anthony, 1 924

Description (after Forman, 1968). — Head much as in S. lilium, differing only
in proportions; apex blunt; no concavity in base (acrosome not examined). Neck
not discernible. Midpiece broad, nonhelical, and long.

Remarks. — The gross morphology of spermatozoa of Sturnira ludovici is
similar to that of S. lilium. However, according to measurements given by Forman
(1968), length of sperm head and length of midpiece are greater in S. ludovici.

Previous study. — Eight specimens from Panama (Forman, 1968).

Uroderma bilobatum Peters, 1866

Description (Fig. 3C). — Head similar in overall morphology to that of
Artibeus jamaicensis; relatively narrow; acrosome symmetrical or slightly
asymmetrical, narrowly rounded at apex; acrosome notable in being extremely
short terminating posteriorly one-third or less the way along the length of the

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STENODERMIN AE

Fig. 3. — Sperm of some stenodermine bats. A) Sturnira l ilium; B) Sturnira tildae ; C)
Vroderma bilobatum; D) Vcimpyrops helleri; E) Vampyrodes caraccioli ; F) Chiroderma
improvisum ; G) Chiroderma trinitatum ; H) Mesophylla macconnelli. Scale equals 5 microns.

BIOLOGY OF THE PHYLLOSTOMATIDAE

189

nucleus; approximately half of acrosome visible anterior to nuclear apex; acrosome
also appearing to be narrower in width than the nucleus; nucleus ovoid; base of
nucleus flattened or slightly concave with pointed corners; head length 4.56
(4.09-4.84) ±0.23, acrosome length 1.98(1.67-2.32) ±0.27, nuclear length
3.32(3.16-3.53) ±0. 12, head length 3.08(2.98-3.16) ±0.08. Neck extremely
short, junction with head well off center. Midpiece of moderate length; thin but
tapering slightly posteriorly; length 8.49(7.91-8.84) ±0.29.

Remarks. — Morphology of the sperm head of Uroderma resembles most
closely that of Artibeus, particularly A. jamaicensis. The acrosome of this species
is unusually short and covers an extremely small portion of the nucleus. The
flattened base of the head is an unusual feature.

Specimen examined. — Trinidad: Guayaguayare, Mayaro, 1 (TTU 24046).

Vampyrops helleri Peters, 1867

Description (Fig. 3D). — Head long and narrow, nucleus relatively long com¬
pared with other species; acrosome narrow and asymmetrical (terminus of apex
on same side of head as the attachment of the midpiece to the head), appears ter be
slightly narrower than nucleus; apex of acrosome narrowly rounded and may be
somewhat pointed; posterior limit of acrosome terminates midway along the length
of the nucleus; a substantial portion of acrosome occurs anterior to the apex of
the nucleus; nucleus strongly ovoid with rounded base that is strongly concave;
apex of nucleus rounded; head length 5.54(5.39-5.77) ±0.14, acrosome length
3.46(3.16-3.72) ±0.26, nuclear length 4.22(4.09-4.37) ±0.1 1, head width
3.41(3.26-3.53) ±0.14. Neck short, junction with head only slightly off center.
Midpiece long, extremely thin; junction with tail distinctive; length 9.54(8.74-
10.14) ±0.41.

Remarks. — Structure and size of the sperm head within this species is unique
among those studied because it is unusually long; it closely resembles that of
Artibeus jamaicensis.

Specimen examined. — Trinidad: Guayaguayare, Mayaro, 1 (TTU 24063).

Vampyrodes caraccioli (Thomas, 1 889)

Description (Fig. 3E). — Head most complete oval of any phyllostomatid
studied with base of head extremely narrow; head egg-shaped, long, relatively
narrow, similar in size but slightly smaller than that of Vampyrops-, nucleus and
acrosome usually with a symmetrical apex at anterior end, apices narrowly
rounded or pointed, acrosomal apex especially pointed; acrosome usually
symmetrical and equal in width to nucleus, in some cases nucleus appears to be
only slightly longer than accompanying acrosome; posterior limit of acrosome
sometimes behind midpoint of nucleus; substantial portion of acrosome occurs
anterior to apex of nucleus; base of head extremely narrow and flattened to
concave, with pointed corners; head length 5.25(4.84-5.49) ±0.202, acrosome
length 2.98(2.79-3.16) ±0.13, nuclear length 4.02(3.72-4.28) ±0.16, head width

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3.21(3.07-3.44) ±0. 1 3. Neck extremely short, attachment to base of head only
slightly off center or is centered. Midpiece of moderate length and breadth;
length 8.89(8.28-9.21) ±0.39.

Remarks. — Head morphology is unique in being long and having an unusually
narrow apex and base. Sperm resembles somewhat that of Vampyrops, but unlike
Vampyrops, Vampyrodes has a symmetrical acrosome and an extremely narrow
and flattened head base.

Specimen examined. — Trinidad: Blanchisseuse, St. George, 1 (TTU 24060).

Chiroderma improvisum Baker and Genoways, 1976

Description (Fig. 3F). — Head similar to that of C. trinitatum, but slightly less
rounded; acrosome sometimes appears to be asymmetrical, short, and with a small
portion extending anterior to nucleus; posterior limit of acrosome lies in front of
midpoint of nucleus and appears less arched than in C. trinitatum ; nucleus ovoid,
apex considerably more rounded than the more pointed apex of the acrosome;
base of head asymmetrical, but less so than in C. trinitatum ; base slightly concave;
head length 4.74(4.37-5.30) ±0.28, acrosome length 2.65(2.60-2.79) ±0.08,
nuclear length 3.96(3.81-4.19) ±0.14, head width 3.17(2.88-3.26) ±0.18.
Neck relatively long, junction with head well off center as in C. trinitatum. Mid¬
piece of moderate breadth, tapering posteriorly; length 8.64(7.53-9.95) ±0.71.

Remarks. — Although similar to that of Chiroderma trinitatum, the sperm head
in C. improvisum is slightly less rounded, its base less asymmetrical, and it pos¬
sesses a shorter acrosome. The spermatozoa of species of Chiroderma can be
distinguised easily from other stenodermines.

Specimen examined. — Guadeloupe: 2 km. S, 2 km. E Baie-Mahault, Basse-Terre,
1 (TTU 19900).

Chiroderma trinitatum Goodwin, 1958

Description (Fig. 3G) — Head morphology generally variable; shape ovoid
to rounded; nucleus ovoid with pointed apex; acrosome nearly symmetrical,
short, with apex only slightly more rounded than that of nucleus; terminal border
of acrosome appears to be slightly arched with apex directed anteriorly; acrosome
terminates posteriorly at midpoint of nucleus and extends anteriorly only very
slightly beyond apex of nucleus; base of head flattened or very slightly concave
and is unusual in being asymmetrical with the greatest posterior extension
occurring on the side of the head that is in contact with the neck; base of head
narrower than girth of head, with corners pointed; head length 4.87(4.56-5.39)
±0.26, acrosome length 3.00(2.70-3.35) ±0.23, nuclear length 3.97(3.62-
4.28) ±0.25, head width 3.37(3.07-3.62) ±0.175. Neck relatively long, junction
with head well off center and nearly to the edge of base of head. Midpiece thin,
tapering gradually posteriorly and short relative to length of head; length 8.84
(8.56-9.1 1) ±0.21.

Remarks. — The morphology of the spermatozoa head in this species, although
variable, is distinctly different from that of other stenodermines. Only a very small

.BIOLOGY OF THE PHYLLOSTOMATIDAE

191

portion of the acrosome extends anterior to nucleus, the base of the head is
asymmetrical, and the point of midpiece attachment is substantially off center.

Specimen examined. — Trinidad: 2 mi. N, 2 mi. E Valencia, St. Andrew, 1 (TTU 24026).

Mesophylla macconnelli Thomas, 1901

Description (Fig. 3H). — Head relatively long and narrow, not large; acrosome
with pointed asymmetrical apex, tip of apex on same side of head as attachment
of midpiece; acrosome short and an extremely small portion of it occurs anterior
to the apex of the nucleus; posterior limit of acrosome slightly anterior to mid¬
point of nucleus; acrosome considerably shorter than the nucleus (often only
slightly more than half its length) and the same breadth as the nucleus at its
posterior limit; nucleus ovoid, apex symmetrical; base of head flattened with
slight concavity; base of head narrower than its girth, asymmetrical with corner
nearest the midpiece being more pointed than the other; head length 4.71(4.56-
5.02) ±0.14, 4.68(4.28-4.93) ±0.19, acrosome length 2.73(2.51-2.88) ±0.12,
2.64(2.51-2.88) ±0.13, nuclear length 4.01(3.62-4.19) ±0.15, 3.99(3.81-

4.37) ±0.22, head width 3.13(2.98-3.34) ±0.12, 3.25(3.07-3.44) ±0.10. Neck
relatively long, junction with head well off center and near the pointed corner of
the head base. Midpiece short, broad anteriorly, tapering abruptly posteriorly;
junction with tail indistinct; length 7.61(7.25-7.92) ±0.23, 7.66(7.25-8.18)
±0.27.

Remarks. — Most notable among the characteristics of sperm from Mesophylla
is the minute amount of acrosome anterior to the nuclear apex and the unusual
asymmetry of the base of the head. The head is somewhat similar to that of
Phyllostomus discolor, but the base and apex of the nucleus are dissimilar.
An extremely short midpiece distinguishes M. macconnelli from other stenoder-
mines, with the exception of Centurio.

Specimen examined. — Trinidad: Guayaguayare, Mayaro, 2 (TTU 24039, 24044).

Artibeus cinereus (Gervais, 1855)

Description (Fig. 4A). — Head broad in midsection, tapering distinctively both
anteriorly and posteriorly; acrosome extremely pointed, nearly cone-shaped,
slightly shorter than nucleus, and terminating posteriorly about midway along
nucleus; nucleus rounded; base of head slightly convex or often lacking concavity,
base of head notably rounded at the corners; head length 4.59(4.28-4.84) ±0.495,
acrosome length 2.93(2.51-3.26) ±0.339, nuclear length 3.62(3.35-3.91) ±0.104,
head width 3.15(2.98-3.26) ±0.084. Neck short, junction with head very slightly
off center. Midpiece broad anteriorly, tapering gradually posteriorly; length 8.74
(8.37-9.02) ±0.342.

Remarks. — Sperm morphology in this species is very similar to that of Artibeus
jamaicensis, Ardops nichollsi, and Ariteus flavescens. The most unusual feature
is the extremely pointed, exceptionally tapered apex to the symmetrical acrosome.

Specimens examined. — Trinidad: Guayaguayare, Mayaro, 1 (TTU 23924); 2 mi. E San
Rafael, St. George, 1 (TTU 23936).

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STENODER M I N AE

E F G H

Fig. 4. — Sperm of some stenodermine bats. A) Artibeus cinereus ; B) Artibeus toltecus,
C) Artibeus jamaicensir, D) Artibeus lituratus ; E) Ardops nichollsi ; F) Phyllops haitiensis ,
G) Ariteus flavescens , H) Stenoderma rufum; I) Centurio senex. Scale equals 5 microns.

BIOLOGY OF THE PHYLLOSTOMATIDAE

193

Artibeus toltecus (Saussure, 1 860)

Description (Fig. 4B). — Head quite long, appearing relatively narrow, similar
to other species of Artibeus ; nucleus ovoid with relatively narrow apex and base;
acrosome symmetrical and pointed at apex; posterior limit of acrosome extending
to midway along length of nucleus; head length 4.96(4.84-5.12) ±0.10, acrosome
length 2.99(2.79-3.07) ±0.09, nuclear length 3.88(3.62-4.19) ±0.20, head
width 3.16(2.98-3.35) ±0.12. Neck short, junction with head well off center.
Midpiece short, compared to length of head, and narrow; tapering posteriorly;
length 8.69(8.37-9.02) ±0.21.

Remarks. — General shape of head similar to Ardops, Ariteus, and other
species of Artibeus, particularly A. lituratus\ however, the head in general is less
rounded than in other species. Heads of spermatozoa from A. toltecus are longer
than in other stenodermines.

Specimen examined. — Veracruz: 4 km. W, 5 km. S Sontecomapa, 1 (TTU 28902).

Artibeus jamaicensis Leach, 1821

Description (Fig. 4C). — Head similar in morphology to that of Ardops and
Ariteus-, acrosome usually symmetrical, but, if asymmetrical, only slightly so;
apex of acrosome narrowly rounded to nearly pointed; portion of acrosome
anterior to nucleus always less than in Ariteus and Ardops-, nucleus narrowly
rounded at apex; base of nucleus broad and slightly concave; head length 4.48
(4.28-4.65) ±0.1 19, acrosome length 2.74(2.51-2.98) ±0.148, nuclear length
3.59(3.35-4.00) ±0.159, head width 3.30(3. 16-3.44) ±0.089. Neck short,
junction with head off center. Midpiece nearly twice head length, thick anteriorly,
and tapering posteriorly; length 8.69(8.09-9.21) ±0.316.

Remarks. — Morphology of the heads of spermatozoa from A. jamaicensis is
quite similar to that of both Ariteus and Ardops, but the portion of the acrosome
anterior to the nucleus was always less in A. jamaicensis. The acrosome has less
symmetry than other species of Artibeus that have been examined.

Previous study. — One specimen from Dominica and one specimen from
Nayarit (Forman, 1968).

Specimen examined. — Haiti: 1 km. E Lebrun, Dept, du Sud, 1 (TTU 22649).

Artibeus lituratus (Olfers, 1818)

Description (Fig. 4D). — Head similar to other Artibeus-, acrosome relatively
larger (as compared with nucleus) than that of other species within the genus;
acrosome only slightly shorter than the nucleus, with somewhat narrowly rounded,
symmetrical apex; acrosome distinctly triangular, its posterior limit consistently
well behind the midpoint of the nucleus; acrosome sometimes slightly narrower
than nucleus, otherwise equivalent in width at its posterior limit; distinctive por¬
tion of acrosome found anterior to nuclear apex; apex of nucleus rounded but
rarely as narrowly as acrosome; base of head asymmetrical with corner nearest
neck slightly more posterior than the rounded corner on the other side of the base;

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base slightly concave; head length 4.77(4.46-5.21) ±0.229, acrosome length
3.30(3.16-3.53) ±0.132, nuclear length 3.59(3.35-3.72) ±0.103, head width
3.23(3.07-3.35) ±0.140. Neck relatively long, junction with head off center.
Midpiece length similar to other species of Artibeus ; tapering gradually pos¬
teriorly; junction with tail quite distinctive; length 8.27(7.91-8.46) ±0.158.

Remarks. — Head morphology of sperm of A. lituratus is similar to that of
other species of Artibeus but is most like A. toltecus, A. jamaicensis, and
Vampyrops helleri.

Previous study. — Two specimens from Chiapas (Forman, 1968).

Specimen examined. — Trinidad: Guayaguayare, Mayaro, 1 (TTU 24010).

Ardops nichollsi (Thomas, 1891)

Description (Fig. 4E). — Head bullet shaped with pointed apex; acrosome
asymmetrical (but sometimes nearly symmetrical); apex pointed or very narrowly
rounded; a moderate portion of acrosome extends forward beyond nucleus; some
acrosomes narrower than nucleus; acrosome shorter than nucleus, terminating
posteriorly at a point slightly anterior to midpoint of nucleus; nucleus extremely
rounded at apex; base broad and deeply concave; head length 4.25(4.00-4.65)
±0.150, 4.31(3.81-4.56) ±0.262, acrosome length 2.42(2.32-2.60) ±0.132,
2.58(2.42-2.70) ±0.1 17, nuclear length 3.22(3.07-3.44) ±0.125, 3.37(3.16-
3.53) ±0.1 15, head width 3.14(2.88-3.44) ±0.260, 3.03(2.88-3.16) ±0.096.
Neck short, junction with head off center. Midpiece of moderate length, thin,
gradually tapering posteriorly; junction with tail not distinctive; length 8.88(8.74-
9.30) ±0.192, 8.54(8.09-9.02) ±0.277.

Remarks. — The symmetry of the acrosome appears to be variable in this
species. In some spermatozoa, acrosomes are asymmetrical, but in others,
nearly symmetrical. Spermatozoa are similar to those of Ariteus and Artibeus.

Specimens examined. — Guadeloupe: 1 km. S Basse-Terre, Basse-Terre, 1 (TTU 20816);
1 km. N, 1 km. W St. Franfois, Grande-Terre, 1 (TTU 20847).

Phyllops haitiensis (J. A. Allen, 1908)

Description (Fig. 4F). — Head usually somewhat triangular in shape; acrosome
only slightly asymmetrical; posterior terminus of acrosome at midpoint of
nucleus; substantial portion of acrosome occurring anterior to the apex of the
nucleus; acrosome shorter than nucleus with similar morphology and placement
(orientation) on the nucleus as Artibeus, Ardops, and Ariteus-, nucleus rounded
with broadly rounded apex; base of nucleus with rounded corners and a slight
concavity or no concavity in center of basal border; head length 4.90(4.28-
5. 12) ±0.23, 4.82(4.65-5.12) ±0.13, acrosome length 2.80(2.51-3.07) ±0.16,
2.78(2.60-2.98) ±0.14, nuclear length 3.76(3.62-3.91) ±0.20, 3.70(3.53-

3.91) ±0.11, head width 3.57(3.35-3.72) ±0.13, 3.32(3.26-3.44) ±0.06. Neck
extremely short, junction with head only slightly off center. Midpiece of moderate
length and breadth, tapering only slightly posteriorly; junction with tail distinctive;
midpiece length 8.74(8.37-9.30) ±0.31, 8.64(8.37-9.1 1) ±0.23.

BIOLOGY OF THE PHYLLOSTOMATIDAE

195

Remarks. — The morphology of the sperm head of Phyllops is similar to that
of Artibeus, Ariteus, and Ardops. Nuclear morphology is most like that of
Artibeus cinereus, but the base of the nucleus is less concave than in most species
of Artibeus.

Specimens examined. — Haiti: 2 km. N, 2 km. E Lebrun, Dept, du Sud, 1 (TTU 22672);
1 km. S, 1 km. E Legrun, Dept, du Sud, 1 (TTU 22697); 4 km. S Lebrun, Dept, du Sud,
1 (TTU 22733).

Ariteus flavescens (Gray, 1831)

Description (Fig. 4G). — Head nearly identical in morphology to that of
Ardops nichollsi-, triangular; acrosome extremely pointed at apex and acrosome
can be asymmetrical or symmetrical; acrosome shorter than nucleus; base of head
broad and concave; head length 4.60(4.37-4.84) ±0.156, acrosome length 2.83
(2.60-3.16) ±0.233, nuclear length 3.27(2.88-3.53) ±0.208, head width 3.49
(3.26-3.62) ±0.136. Neck short, junction with head off center. Midpiece of
moderate breadth anteriorly, tapering posteriorly; junction with tail distinctive;
length 9.08(8.56-9.30) ±0.304.

Remarks. — The head of spermatozoa from this species bears a striking re¬
semblance to that of Ardops nichollsi. The two also are extremely similar in
dimensions of the nucleus, acrosome, and length of midpiece.

Specimens examined. — Jamaica: Queenhythe, St. Ann Parish, 1 (TTU 21774); Duanvale,
Trelawny Parish, 1 (TTU 21781).

Stenoderma rufum Desmarest, 1 820

Description (Fig. 4H). — Head most similar in shape to those of Ariteus,
Ardops, and Artibeus-, more or less triangular, both nucleus and acrosome
generally symmetrical; acrosome short and usually quite pointed at apex; acrosome
usually narrower at base than is nucleus at its widest point; acrosome can be
slightly asymmetrical at apex in that sometimes it is offset to side of head with
attachment to neck; one-third to half of acrosome occurring anterior to the apex
of nucleus; posterior border of acrosome lies anterior to midpoint of nucleus;
nucleus nearly triangular with broadly rounded apex and quite rounded corners
at the base; base slightly concave to nearly flattened; head length 4.58(4.19-
4.84) ±0.18, 4.48(4.37-4.65) ±0.13, acrosome length 2.92(2.60-3.26) ±0.16,
2.81(2.42-2.98) ±0.17, nuclear length 3.46(3.26-3.81) ±0.13, 3.56(3.44-

3.81) ±0.11, head width 3.20(2.88-3.35) ±0.15, 3.21(3.07-3.35) ±0.10. Neck
relatively long, junction with base of head moderately off center. Midpiece
relatively broad, tapering gradually posteriorly; junction of midpiece and tail
distinctive; length 8.50(8.18-8.84) ±0.20, 8.33(8.09-8.65) ±0.16.

Remarks. — Head of sperm in this species is most similar to that of Ariteus
flavescenes, Ardops nichollsi, and members of the genus Artibeus but is dis¬
tinguishable from all of them. The most unusual feature of the spermatozoa of this
species is the narrowness of the acrosome relative to the breadth of the nucleus.
Also, the nucleus and acrosome are extremely similar in outline, a situation rarely
observed.

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SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

Specimens examined. — Puerto Rico: El Verde, 2 (TTU 22361, 22362).

Centurio senex Gray, 1 842

Description (Fig. 41). — Head short, nuclear portion extremely rounded;
acrosome symmetrical with extremely pointed apex, forming an isosceles triangle,
as wide as nucleus; posterior limit of acrosome lies in front of center of nucleus;
acrosome shorter than nucleus; moderate portion of acrosome occurs anterior
to the nuclear apex, which is narrowly rounded; nucleus usually as wide as it is
long with its anterior border often appearing flattened on either or both sides; base
of head flattened or even slightly convex, giving base a rounded appearance; head
length 4.44(4. 19-4.74) ±0.20, acrosome length 2.68(2.42-3.07) ±0.28, nuclear
length 3.68(3.44-4.00) ±0. 1 8, head width 3.65(3.35-3.91) ±0.17. Neck long,
junction with head well off center. Midpiece extremely thin, short; length 7.36
(7.34-7.91) ±0.20.

Remarks. — The morphology of the sperm head in Centurio senex is distinctive
and unique. The acrosome is extremely pointed, the nucleus nearly circular.
Perhaps the greatest contrast in degree of pointedness of nuclear and acrosomal
apices is observed in this species.

Specimen examined. — Trinidad: Blanchisseuse, St. George, 1 (TTU 24019).

Subfamily Phyllonycterinae
Brachyphylla cavernarum Gray, 1 834

Description (Fig. 5A). — Head of moderate length, narrow; acrosome
symmetrical, considerably shorter than nucleus, and with its posterior limit well
anterior to midpoint of nucleus; nucleus more ovoid than that of Ardops, Ariteus,
and Artibeus\ base slightly concave; head length 4.60, 5.12, acrosome length
2.79, 2.79, nuclear length 3.26, 3.53, head width 2.79, 1.98. Neck short, junction
with head near center. Midpiece of moderate width, long, tapering posteriorly;
junction with tail distinctive.

Remarks. — The sperm of Brachyphylla is different from other phyllonycterines
and does not possess features generally found among other members of the sub¬
family (for example, Brachyphylla differs in shape and size of the acrosome,
relative length of the midpiece, symmetry of the head).

Specimens examined. — Guadeloupe: 1 km. S Basse-Terre, Basse-Terre, 1 (TTU 20966);
1 km. N, 1 km. W St. Francois, Grande-Terre, 1 (TTU 20976).

Erophylla bombifrons (Miller, 1899)

Description (Fig. 5B). — Head extremely long, ovoid and generally robust;
acrosome large and encompassing a distinctive portion of the head; acrosome
with slight asymmetry, anteriormost limit of apex on the same side of head as
attachment of tail, and with an apex quite similar in shape to that of the nucleus;
acrosome only slightly wider than the nucleus, terminating posteriorly just beyond
midpoint of nucleus; acrosome only slightly shorter than nucleus; nucleus broad

BIOLOGY OF THE PHYLLOSTOMATIDAE

197

PHYLLONYCTERINAE

DESMODONTINAE

E F G

Fig. 5. — Sperm of some phyllonycterine and desmodontine bats. A) Brachyphylla
cavernarum; B) Erophylla bombifrons\ C) Erophylla sezekornr, D) Pbyllonycteris poeyr,
E) Desmodus rotundas , F) Diaemus youngii; G) Diphylla ecaudata. Scale equals 5 microns.

and usually rounded, apex symmetrical; base of nucleus strongly asymmetrical
and concave, with corner nearest attachment of midpiece often less rounded than
other corner; head length 5.14(4.84-5.30) ±0.148, acrosome length 3.45(3.26-
3.62) ±0.142, nuclear length 3.95(3.62-4.09) ±0.146, head width 3.62(3.53-

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3.81) ±0.09. Neck appears extremely long; junction with base of head off center.
Midpiece broad anteriorly, tapering abruptly; length 7.42(7.07-8.37) ±0.379.

Remarks. — The head of the sperm of Erophylla bombifrons is similar to that
of Phyllonycteris poeyi', however, the acrosome of E. bombifrons is smaller and
not so asymmetrical. The midpiece of this species is exceptionally thick at its
anterior end.

Specimens examined. — Puerto Rico: 1 mi. W Corozal, 2 (TTU 22426, 22429).

Erophylla sezekorni (Gundlach, 1861)

Description (Fig. 5C). — Head narrow and long, oval in general shape; acrosome
exceedingly asymmetrical with apex on same side of head as midpiece attachment;
acrosome slightly wider than nucleus at its widest point; posterior terminus of
acrosome at or slightly posterior to midpoint of head; acrosome shorter than
nucleus by small amount and with apex more narrowly rounded than that of
nucleus; nucleus an egg-shaped, rounded oval with broadly rounded apex; base
of head rounded or slightly concave; amount of acrosome anterior to nucleus
variable but generally amount is moderate; head length 4.74, 4.84, acrosome
length 2.79, 2.98, nuclear length 3.44, 3.53, head width 2.98, 3.07. Neck moderate
in length, junction with head slightly off center. Midpiece short, broad anteriorly
(but considerably less so than in E. bombifrons), and tapering gradually pos¬
teriorly; junction with tail indistinct; length, 7.53.

Remarks. — The head of the sperm of Erophylla sezekorni is like that of E.
bombifrons but is more similar to that of Phyllonycteris poeyi in general char¬
acteristics. The acrosome in Erophylla is much smaller than in Phyllonycteris
and with considerably less exposed acrosome than in sperm of Phyllonycteris. The
thickened area of the tail just distal to the midpiece in P. poeyi was not observed
in either species of Erophylla.

Specimen examined. — Jamaica: Orange Valley, St. Ann Parish, 1 (TTU 21894).

Phyllonycteris poeyi

Description (Fig. 5D). — Head extremely long and broad because of enormous
asymmetrical acrosome; acrosome slightly wider than long with apex extremely
broad and on same side of head as midpiece attachment; apex of acrosome even
more removed from the midline of nucleus than midpiece, with result that the
apex is often so far off center as to be outside the axis of the nucleus; acrosome
broadest of any phyllostomatid studied and broader than nucleus; acrosome
terminates posteriorly slightly beyond the midpoint of nucleus; nucleus bilaterally
symmetrical except for base; nucleus a broad oval, being slightly longer than
acrosome; base of nucleus concave, and of moderate breadth, apex rounded; head
length 6.42(6. 14-6.98) ±0.214, 6.67(6.32-6.88) ±0.204, acrosome length

4. 13(3.8 1-4.50) ±0.2 15, 4.56(4.28-5.02) ±0.234, nuclear length 4.74(4.56-
5.02) ±0.156, 4.73(4.46-5.02) ±0.201, head width 4.60(4.19-4.74) ±0.169,
4.57(4.19-4.74) ±0.157. Neck short, junction with head off center. Midpiece of
moderate length; broad anteriorly and tapering posteriorly; unusual tapered

BIOLOGY OF THE PHYLLOSTOMATIDAE

199

thickening of tail just distal to junction of tail and midpiece; length 8.63
(8.18-8.84) ±0.204, 8.63(8.28-8.93) ±0.237.

Remarks. — The sperm of P. poeyi exhibits several unique characteristics. The
acrosome has an unusual morphology including having the apex far offset and
being the broadest of any species studied. This is the only species examined in
which over half of the area of the acrosome occurs anterior to the apex of the
nucleus. There is an unusual thickening in the tail of all specimens that occurs just
distal to the junction of the tail and midpiece; the thickened area tapers posteriorly
into a narrow tail.

Specimens examined. — Haiti: 1 km. E Lebrun, Dept, du Sud, 1 (TTU 22773); 1 km.
S Lebrun, Dept, du Sud, 1 (TTU 22782); 4 km. S Lebrun, Dept, du Sud, 1 (TTU 22798).

Subfamily Desmodontinae
Desmodus rotundus (E. Geoffroy St. -Hilaire, 1810)

Description (Fig. 5E). — Head long, narrow, and extremely ovoid with narrowly
rounded apex and narrow base; acrosome long, terminating posteriorly well
behind midpoint of nucleus, apex symmetrical; most of acrosome in contact
with nucleus, only an extremely minute portion anterior to nuclear apex; viewed
dorsally, nucleus comprises most of head; acrosome no wider than nucleus, apex
of acrosome slightly more rounded than that of nucleus; base of head quite narrow,
with distinctive concavity at junction with neck; head length 4.71(4.46-4.93) ±
0.183, acrosome length 2.98(2.88-3.07) ±0.067, nuclear length 3.84(3.62-4.09)
±0.162, head width 2.71(2.51-2.88) ±0.103. Neck extremely short; attaches at
center of head. Midpiece extremely long, thickened or even flared at neck; tapers
gradually posteriorly; junction with tail moderately distinctive; length 11.64
(1 1.16-12.18) ±0.277.

Remarks. — The heads of the spermatozoa of Desmodus rotundus show much
greater symmetry than other phyllostomatid subfamilies. The other unique
features of the sperm of this species include the relatively long and narrow head,
long midpiece that is flared at the anterior end, and an acrosome closely attached
to the nucleus.

Previous study. — Two specimens from Nicaragua (Forman et al., 1968).

Specimens examined. — Trinidad: 2 mi. N, 2 mi. E Valencia, St. Andrew, 1 (TTU 24086);
Blanchisseuse, St. George, 1 (TTU 24080).

Diaemus youngii (Jentink, 1 893)

Description (Fig. 5F). — Head very similar in structure to that of Desmodus
rotundus, however, acrosome protrudes well anterior of apex of nucleus; acrosome
symmetrical, relatively narrow compared to D. rotundus , and with posterior
limit often well in front of the midpoint of the nucleus; apex of acrosome somewhat
more rounded than that of the nucleus; nucleus longer than acrosome; nucleus
nearly identical to that of Desmodus except base is concave or flattened; head
length 5.61(5.21-5.95) ±0.249, acrosome length 3.20(2.98-3.53) ±0.170,
nuclear length 4.50(4.28-4.74) ±0.135, head width 3.1 1(2.98-3.35) ±0.104.

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Neck extremely short, junction with head at center or very slightly off center.
Midpiece extremely long and extremely broad anteriorly; tapering abruptly then
gradually posteriorly; length 12.51(1 1.81-12.83) ±0.255.

Remarks. — Sperm of Diaemus youngii is very similar to that of Desmodus
rotundus but quite different from the sperm of the third member in the subfamily,
Diphylla ecaudata. The difference in head length between Desmodus and Diaemus
is due, in part, to the position of the acrosome on the nucleus. The midpiece of
Diaemus is longer than any other species of phyllostomatid studied and appears
to lack the flared anterior end found in the sperm of Desmodus.

Specimen examined. — Trinidad: La Brea, St. Patrick, 1 (CM 45371).

Diphylla ecaudata Spix, 1823

Description (Fig. 5G). — Head clearly a shovel-shaped, extremely broad,
rounded triangle; acrosome closely applied to front of nucleus as in Desmodus ;
acrosome barely anterior to the nuclear apex (in some cases it cannot be seen);
acrosome large, generally assumes shape of the nucleus at its apex but can be
more pointed; acrosome terminates posteriorly well beyond the midpoint of the
nucleus as in Desmodus ; acrosome the same width as the nucleus throughout most
of its length; nucleus considerably longer than acrosome, its base asymmetrical,
broad, with corners somewhat pointed; a distinctive depression in base of head at
junction with neck; head length 4.57(4.37-4.84) ±0.160, acrosome length 2.89
(2.70-3.16) ±0.154, nuclear length 4.22(4.02-4.63) ±0.154, head width 3.46
(3.26-3.62) ±0.126. Neck slightly longer and somewhat broader than other
vampires; attachment to base of head at one comer of base. Midpiece long, broad
anteriorly and tapering gradually posteriorly; junction with tail not distinctive;
length 9.60(9.21-10.14) ±0.294.

Remarks. — Morphology of the sperm head of Diphylla ecaudata is quite
different from the other two species of vampires — most distinctive is the great
breadth of the nucleus and the attachment of the head farther off center than noted
for any other species examined.

Previous study. — Two specimens from Nicaragua (Forman et al., 1968).

Specimen examined. — Yucatan: 3 km. S, 1 km. W Calcehtoc, 1 (TTU 18447).

Discussion

The spermatozoa of 35 species representing all six of the subfamilies of the
Phyllostomatidae were examined in this study. Descriptions of three additional
phyllostomatid species are available in the literature (Forman, 1968). The
morphology of all species studied is basically similar, and this serves to distinguish
members of the Phyllostomatidae from those of other families of bats. The
acrosome proved to be the most variable structure, more variable than even the
nuclear region.

Below we will discuss the relationships by subfamily that were observed in this
work.

BIOLOGY OF THE PHYLLOSTOMATIDAE

201

Phyllostomatinae. — Acrosomes within this subfamily were almost universally
asymmetrical and always extended well anterior to the nuclear apex. Sperm
from Mimon crenulatum and Macrotus waterhousii were most dissimilar from
other members of the subfamily and from each other. Mimon possesses a
strikingly enlarged and asymmetrical acrosome, whereas Macrotus is characterized
by the unusual configuration of the nucleus, particularly by its unique broad base.
Sperm of Phyllostomus, Micronycteris, and Tonatia were quite similar, and
Phyllostomus and Micronycteris were characterized further among the phyllosto-
matines by a relatively long midpiece.

Glossophaginae. — Heads of the spermatozoa from this subfamily were rather
rounded. Sperm from Choeronycteris showed a larger head and a substantially
longer midpiece than either Anoura or Glossophaga. Anoura was distinguished
from other glossophagines by a more strongly concave base to the head and from
other phyllostomatids by an unusually short midpiece.

Spermatozoa were found to be no more variable within this subfamily than
they were among the phyllostomatines or desmodontines. Therefore, sperm
morphology does not support the contentions based on karyology (Baker, 1967),
dental anatomy (Phillips, 1971), and immunologic comparisons (Gerber and
Leone, 1971) that the glossophagines are a polyphyletic grouping.

Carolliinae. — The sperm of three species of the genus Carollia that have been
studied were similar, with the nuclei being quite rounded. However, the species
can be distinguished from each other based on overall head morphology.

Stenoderminae. — Morphology of the sperm heads of stenodermines was highly
variable. Acrosomes varied from pointed and nearly symmetrical ( Centurio ) to
broadly rounded at the apex and strongly asymmetrical ( Chiroderma ). There
was considerable variability in the point of attachment of the neck and midpiece
to the base of the head and ranged from nearly central attachment to attachment
near the edge of the base of the head. However, the length and breadth of the mid¬
piece of stenodermines was similar, except for Mesophylla, in which the mid¬
piece was shorter than in other species.

Sperm from Ardops, Ariteus, Stenoderma, Phyllops, and Artibeus were
alike in size and morphology of the nucleus and acrosome. Members of the first
four genera are Antillean endemics characterized by shortened rostra and white
spots on their shoulders. These genera are believed to have resulted from a single
invasion of the Antilles (Baker and Genoways, 1978) with subsequent radiation.
Morphology of the sperm supports this hypothesis and also suggests that members
of this group may share a close ancestor with members of the genus Artibeus.
Uroderma bilobatum is similar in morphology to members of this group, except
that in Uroderma the base of the head is flattened and has pointed corners.

Sperm heads of Centurio senex were unusually triangular in form with the base
of the head unusually broad. In members of the genus Vampyrops, the nucleus
was extremely long, but in Vampyrodes, the distinguishing feature was the narrow
base of the head. In addition to the shortened midpiece, Mesophylla is char¬
acterized by the strongly asymmetrical base of the head.

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The sperm of Chiroderma improvisum and C. trinitatum were the most
unique in head morphology among the stenodermines examined. In both species,
only a very small portion of the acrosome extends beyond the apex of the nucleus.
Furthermore, the base of the nucleus is asymmetrical with the greatest posterior
extension occurring on the side of the head that is in contact with the neck. The
sperm of these two species are similar but C. improvisum can be distinguished
from C. trinitatum by the head of the former being slightly less rounded, acro¬
some shorter, and base of head less asymmetrical.

Until recently, members of the genus Sturnira were placed in a separate sub¬
family, Sturnirinae. However, recent authors (Baker, 1967; Slaughter, 1970;
Jones and Carter, 1976) have placed them in the subfamily Stenoderminae. The
morphology of the sperm of the three species described herein were similar, all
being characterized by nearly symmetrical acrosomes. Sperm head morphology
of species of Sturnira was most similar to that of stenodermines, among the sub¬
families we examined, and we believe our data support placement of members
of the genus Sturnira in the subfamily Stenoderminae. Although the sperm of the
three species of Sturnira were similar, they could be distinguished on
the basis of size and details of morphology.

Phyllonycterinae. — The sperm of Brachyphylla cavernarum was completely
unlike that of any other phyllonycterines examined. Similarity in sperm mor¬
phology does not support placement of Brachyphylla in the Phyllonycterinae,
as suggested by Silva Taboada and Pine (1969) from morphological and be¬
havioral investigations and Baker and Lopez (1970) based on karyology. Our
data indicate that it would be best to follow Miller (1907) and place Brachyphylla
in the subfamily Stenoderminae. Among the stenodermines, the sperm of
Brachyphylla could be distinguished by its long midpiece.

The sperm head of other phyllonycterine species studied was more uniform
than that of species within other subfamilies; heads were all relatively narrow
and acrosomes were large and asymmetrical. Spermatazoa from Erophylla
hombifrons, E. sezekorni, and Phyllonycteris poeyi were especially similar to
those of Anoura and Caro Ilia.

The sperm of Phyllonycteris poeyi possesses a unique enlargement in the tail
just distal to its junction with the midpiece. This structure was not seen in any
other phyllostomatids examined.

Desmodontinae. — Sperm from the three species of vampire bats were markedly
different; the only common feature among the three was a midpiece that proved
to be the longest among the Phyllostomatidae. Diphylla possessed sperm heads
that were substantially broader and more rounded than those of Desmodus and
Diaemus. The nuclear portion of the head was similar in Desmodus and Diaemus\
however, in Diphylla the nucleus was broader. Sperm from Diphylla was also
characterized by the neck and midpiece juncture with the head being placed
farther off center than any other phyllostomatid studied.

Spermatazoa of Desmodus and Diphylla show great similarity in the close
application of the acrosome to the nucleus, with little space between the apices
of the acrosome and the nucleus. The acrosome also extends posteriorly beyond

BIOLOGY OF THE PHYLLOSTOMATIDAE

203

the midpoint of the nucleus. Neither of these two characteristics appear in
Diaemus.

Literature Cited

Baccetti, B. 1970. Comparative spermatology. Academic Press, New York, 573 pp.

Baker, R. J. 1967. Karyotypes of bats of the family Phyllostomidae and their taxonomic
implications. Southwestern Nat., 12:407-428.

Baker, R. J., and H. H. Genoways. 1978. Zoogeography of Antillean bats. Pp. 53-97, in
Zoogeography in the Caribbean (F. B. Gill, ed.). Spec. Publ. Acad. Nat. Sci.
Philadelphia, 1 3 : iii + 1-128.

Baker, R. J., and G. Lopez. 1970. Karyotypic studies of the insular populations of
bats on Puerto Rico. Caryologia, 23:465-472.

Bedford, J. M. 1967. Observations on the fine structure of spermatozoa of the bush baby
( Galago senegalensis), the African green monkey ( Cercopithecus aethiops) and
man. Amer. J. Anat., 121:443-459.

Biggers, J. D. 1966. Reproduction in male marsupials. Pp. 251-280, in Comparative
biology of reproduction in mammals (I. W. Rowlands, ed.), Academic Press, Inc.,
New York, xxi + 559 pp.

Biggers, J. D., and E. D. Delamater. 1965. Marsupial spermatozoa: pairing in the
epididymis of American forms. Nature, 208:402-404.

Bishop, M. W. H., and C. R. Austin. 1957. Mammalian spermatozoa. Endeavour,
16:137-150.

Braden, A. W. H. 1959. Strain differences in the morphology of the gametes of the
mouse. Australian J. Biol. Sci., ser. B, 12:65-71.

Fawcett, D. W., and S. Ito. 1965. The fine structure of bat spermatozoa. Amer. J.
Anat., 1 16:567-610.

Forman, G. L. 1968. Comparative gross morphology of spermatozoa of two families of
North American bats. Univ. Kansas Sci. Bull., 47:901-928.

Forman, G. L., R. J. Baker, and J. D. Gerber. 1968. Comments on the systematic
status of vampire bats (family Desmodontidae). Syst. Zool., 17:417-425.

Friend, G. F. 1936. The sperms of British Muridae. Quart. J. Micros. Sci., 78:419-443.

Genoways, H. H. 1973. Systematics and evolutionary relationships of spiny pocket
mice, genus Liomys. Spec. Publ. Mus., Texas Tech Univ., 5:1-368.

Gerber, J. D., and C. A. Leone. 1971. Immunologic comparisons of the sera of certain
phyllostomatid bats. Syst. Zool., 20:160-166.

Griffiths, M. 1968. Echidnas. Internat. Ser. Monogr. Pure and Applied Biol., Zool.
(G. A. Kerkut, gen. ed.), 38:ix + 1-282.

Helm, J. D., and J. R. Bowers. 1973. Spermatozoa of Tylomys and Ototylomys. J.
Mamm. 54:769-772.

Hirth, H. F. 1960. The spermatozoa of some North American bats and rodents. J.
Morph., 106:77-83.

Hughes, R. L. 1964. Sexual development and spermatozoan morphology in the male
macropod marsupial Potorotts tridactylns ( Ken). Australian J. Zool., 12:42-51.

- . 1965. Comparative morphology of spermatozoa from five marsupial families.

Australian J. Zool., 14:533-543.

Jones, J. K., Jr., and D. C. Carter. 1976. Annotated checklist, with keys to subfamilies
and genera. Pp. 7-38, in Biology of bats of the New World family Phyllostomatidae,
Part I (R. J. Baker, J. K. Jones, Jr., D. C. Carter, eds.), Spec. Publ. Mus., Texas
Tech Univ., 10:1-218.

Linzey, A. V., and J. N. Layne. 1974. Comparative morphology of spermatozoa of the
rodent genus Peromyscus (Muridae). Amer. Mus. Novitates, 2532:1-20.

Martin, D. E„ K. G. Gould, and H. Warner. 1975. Comparative morphology of
primate spermatozoa using scanning electron microscopy. I. Families Hominidae,
Pongidae, Cercopithecidae and Cebidae. J. Human Evol., 4:287-292.

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McFarlane, R. W. 1963. The taxonomic significance of avian sperm. Proc. Internat.
Ornith. Congr., 8:91-102.

Miller, G. S., Jr. 1907. The families and genera of bats. Bull. U.S. Nat. Mus.,

57:xvii + 1-282.

Phillips, C. J. 1971. The dentition of glossophagine bats: development, morphological
characteristics, variation, pathology, and evolution. Misc. Publ. Mus. Nat.
Hist., Univ. Kansas, 54:1-138.

Silva Taboada, G., and R. H. Pine. 1969. Morphological and behavioral evidence for
the relationship between the bat genus Brachyphylla and the Phyllonycterinae.
Biotropica, 1:10-19.

Slaughter, B. H. 1970. Evolutionary trends of chiropteran dentitions. Pp. 51-83,

in About bats (B. H. Slaughter and D. W. Walton, eds.), Southern Methodist Univ.
Press, Dallas, Texas, vii + 339 pp.

Wimsatt, W. A., P. H. Krutzsch, and L. Napolitano. 1966. Studies on sperm survival
mechanics in the female reproductive tract of hibernating bats. I. Cytology and
ultrastructure of intrauterine spermatozoa in Myotis lucifugus. Amer. J. Anat.,
191:25-59.

Wooley, D. M., and R. A. Beaty. 1967. Inheritance of midpiece length in mouse
spermatozoa. Nature, 215:94-95.

ALIMENTARY TRACT

G. Lawrence Forman, Carleton J. Phillips, and C. Stanley Rouk

Bats of the family Phyllostomatidae have extremely diversified dietary habits.
Although accurate and detailed dietary data often are unavailable, there never¬
theless are generalizations that can be made and certain trends seem obvious
(Gardner, 1977; Phillips et ai, 1977). In addition to differences in diet, there also
are differences in feeding behavior and in feeding strategies. Nonalimentary
structural specializations such as reduced dentitions, elongate tongues (Phillips
et al., 1977), elaborate lip ridges, and complex palatal topography also are
common in leaf-nosed bats.

In view of the great variability in alimentary function, it is reasonable to
hypothesize that the gut tube itself might be unusually variable within the
Phyllostomatidae. This is especially true in comparison to other families of bats,
in which the dietary habits are not nearly so diversified. Current data suggest that
at least certain portions of the alimentary tract are in fact highly variable.

This account reviews what already is known about gastrointestinal structure
in phyllostomatids and reports new information, particularly with regard to
histology and histochemistry of the stomach. However, certain alimentary regions,
such as the intestine and esophagus, still require investigation for almost nothing is
known about them. A survey of esophageal structure could prove particularly
interesting because of the wide array of food items ingested by leaf-nosed bats. In
all likelihood, the esophagus will reflect diet-specific morphological adaptations.
Continuing comparative analysis of digestive tract morphology undoubtedly will
prove important to our understanding of systematic relationships as well as to our
understanding of the evolutionary process.

Materials and Methods

Some information presented in this chapter was extracted from a Ph.D.
dissertation by Rouk (1973). In that study, the following histological and histo-
chemical procedures were employed: fixation — 10 per cent neutral, buffered
formalin; straining of sections — a, Harris hematoxylin and eosin; b, aldehyde-
fuchsin for elastin and acid mucopolysaccharides; c, Hale’s colloidal iron followed
by acid fuchsin, Ponceau 2R, and phosphotungstic acid sequence for acid muco¬
polysaccharides and chief cells; and d, Masson’s triple connective tissue stain.

Esophagus

The histological organization of the esophagus in phyllostomatids is similar
to that of other bats and other kinds of mammals as well. As is typical for the
Chiroptera, the phyllostomatid esophagus in preserved specimens appears
to be unusually narrow. The luminal surface is characterized by protruding

205

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longitudinal folds of stratified squamous epithelium. The esophagus of large¬
sized phyllostomatids can be relatively narrower than that of smaller species;
for example, Robin (1881) found that the esophagus of one species of Artibeus
was only slightly broader than that of a species of Glossophaga, even though the
body of the former was three times that of the latter.

Kolb (1954), who reviewed esophageal structure in bats, found some specific
variation in the amount of cornification (keratinization) of the esophageal
epithelium. He (Kolb, 1954) thought that such variation could reflect adaptations
for the ingestion of particular foods. A similar finding was reported for the oral
cavity (Phillips et al., 1977), and it also was suggested that the degree of
cornification could be a local response to a given amount of surface stress rather
than a specific, inherited feature. The most complete histological study of the
esophagus of a phyllostomatid is that by Moller (1932), who investigated
Glossophaga soricina. As might be predicted, he found that the esophagus of
G. soricina lacked significant corneum, particularly in the lower abdominal
portion. Cells lining the esophageal lumen had ovoid nuclei, unlike those
characteristic of dead, cornified cells. This feature probably is reflective of the
general absence of abrasive food in the diet of Glossophaga and certainly is in
contrast to the histology of insectivorus species in which the esophogeal surface
is cornified.

Stomach

Comparative gastrointestinal structure and function is of particular interest
because of the variability in diet among phyllostomatid species. It is because of
this diversity in diet that the phyllostomatids have been subjects of more detailed
studies of alimentary structure (especially the stomach) than have other
families of bats. The following account, therefore, deals predominantly with
morphology of the stomach because knowledge of variability in this structure in
leaf-nosed bats even exceeds that for most other groups of small mammals.
Comments on the small intestine, insofar as data are available, also are included.

In most cases, stomachs of phyllostomatids can be described in terminology
that has been applied to other mammals. In those instances in this account
where unusual or less familiar terms apply, a brief explanation parenthetically
follows the term.

In all species thus far studied, the stomach has the form of a local dilation
of the enteron. Torsion produces a saclike structure with a lesser curvature
(anterior) and a greater curvature (posterior). Specific variability in topography,
therefore, has been accomplished by evolutionary modification of this general
plan. Gastric glands occur throughout the mucosa of all species studied. Squamous
epithelium, on the other hand, has been lacking. The summary given in the
following paragraphs is based predominantly on the works of Forman (1971a,
1971b, 1972, 1973), Rouk and Glass (1970), and Rouk(1968, 1973).

BIOLOGY OF THE PHYLLOSTOM ATIDAE

207

Fig. 1. — Semidiagramatic representations of the stomachs of selected phyllostomatines.
The hatched area indicates the region of pylofundic transition glands: a, Micronycteris
megalot is; b, Macrophylum macrophylum; c, Tonatia b ideas, d, Phyllostomus discolor,
e, Phylloderma septentrionalis, f, Vampyrum spectrum. Scale is 10 mm.; upper scale is
for figs, a to e; lower scale, f.

Gross Morphology
Phyllostomatinae

The phyllostomatines have the simplest and least specialized stomachs.
This probably relates to their somewhat unspecialized or primitive feeding
habits that include insectivorous, carnivorous, and omnivorous diets. The
stomach in Micronycteris is extremely simple in configuration; a cardiac
vestibule usually is lacking. The pyloric tube (portion between the esophagus
and duodenum) usually is short, with that of M. nicefori being relatively longer
than that of M. hirsuta or M. megalotis (Fig. la). The fundic caecum ( = cardiac
caecum) is modestly developed in all three of these species. The stomachs of
Macrotus waterhousii and Macrophyllum macrophyllum (Fig. lb) also are
simple and generally resemble those of Micronycteris.

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The stomach in Tonatia varies somewhat from those previously mentioned,
and that of Tonatia minuta varies intraspecifically. For example, the stomach of
T. minuta may have a poorly developed fundic caecum. Additionally, the pyloric
tube is bent at a right angle to the general orientation of the stomach, as observed
in T. bidens (Fig. lc), or it may more closely approximate the simple, symmetrical
configuration found in species of Micronycteris. The esophageal entrance is located
about midway along the lesser curvature. The pyloric tube in Chrotopterus auritus
differs from that in species of Micronycteris only in being relatively longer.

The stomach of Phylloderma stenops (Fig. le) is more globular than those
of other phyllostomatines, but otherwise it does not differ substantially from those
found in species of Micronycteris. The stomach of Trachops is Micronycteris- like
but still is more tubular, and the lesser and greater curvatures are nearly parallel.

The stomachs of several other phyllostomatines differ more distinctively
from the Micronycteris- like configuration. For example, in Phyllostomus
discolor (Fig. Id) and P. hastatus the fundic caecum is well developed and
often is dilated at its terminus. The pyloric portion is distinctively elongated
and sometimes there is a prominent constriction in front of the gastroduodenal
junction. A small, but perceptable cardiac vestibule occurs between the lesser
curvature and the gastroesophageal junction. Although this vestibule is not
nearly so expansive as that in some frugivores, it nevertheless is more
distinctive than that of phyllostomatines described above. The stomach of P.
hastatus generally resembles that of P. discolor , except for its considerably larger
size. The greater and lesser curvatures are nearly parallel in both species.

The stomach of Vampyrum spectrum (Fig. If), a carnivore that often feeds
on other bats (see Rouk, 1973), is noticeably pearshaped with a moderately
developed fundic caecum and a long, well differentiated pyloric tube. A cardiac
vestibule is lacking and the lesser curvature is longer than in other phyllosto¬
matines. This is because the pyloric tube exits to the side (right side of the body)
with only very slight anterior recurvature of the terminal portion of the stomach.
The stomach of this species, with its straight pyloric tube, has a strong resemblance
to those of many species of the Insectivora (see Allison, 1948; Myrcha, 1967).

Simplicity of stomach form is evident in the Phyllostomatinae. Some elongation
of the pyloric portion, along with some dilation of the caecum also, is evident
in comparison with stomachs of insectivorous bats of other families. These slight
modifications likely are associated with increased volume of food ingested.

Glossophaginae

The stomach of Glossophaga soricina (Fig. 2a) is large and saccular.
Although its diet includes insects along with nectar, pollen, and fruit, the
stomach is decidedly more specialized than that of any of the Phyllostomatinae,
including the omnivorous Phyllostomus discolor.

The fundic caecum in G. soricina is dilated and bulbar. The caecum can be
distinguished from the remainder of the stomach by a distinctive furrow or
sulcus on the dorsal surface. The stomach is curved in both frontal and transverse

BIOLOGY OF THE PHYLLOSTOMATIDAE

209

d e

Fig. 2. — Semidiagramatic representations of the stomachs of selected glossophagines
and a carolliine. The hatched area indicates the region of pylofundic transition glands:
a, Glossophaga soricina; b, Hylonycteris underwoodr, c, Lonchophylla robusta; d,
Lichonycteris obscurer, e, Carol! ia perspicillata. Scale is 10 mm. for e; for all others,
8 mm. Symbols are FC, fundic caecum; CV, cardiac vestibule.

planes. A small cardiac vestibule has been observed in some specimens, but
seems to be absent in others. This variable feature possibly is an individual
response to opportunistic feeding by this species. Glossophaga commissarisi
has a stomach that is similar to that of G. soricina except for its even more
distinctive cardiac vestibule. The fundic caecum is relatively longer and narrower
than that of G. soricina. The pyloric tube is enlongated and more distinctive
than in G. soricina.

Even though stomachs of Hylonycteris underwoodi (Fig. 2b), Lonchophylla
robusta, Anoura geoffroyi, Choeronycteris mexicana, and Leptonycteris all
bear a general resemblance to those in Glossophaga , distinguishing characteristics
can be observed in most. For example, Hylonycteris has a relatively long, narrow
fundic caecum (Fig. 2b) that is nearly tubular and is marked by numerous deep
sulci. The extemely broad pyloric tube is short, but decidedly arched from left
to right. The stomachs of Anoura geoffroyi and Choeronycteris mexicana bear
striking resemblance to those of Glossophaga. In comparison to the other
glossophagines, Lonchophylla robusta has an unusual stomach (Fig. 2c) in
that both the cardiac vestibule and fundic caecum are developed distinctively.
The gross morphology of this stomach approaches that of some fruit-eating
stenodermines.

The stomachs of Leptonycteris nivalis and L. sanborni are nearly identical.
They also are somewhat distinctive because of an unusually elongated, extremely
pointed fundic caecum. Also, the terminal portion of the stomach (pylorus) is
tubular and elongated to the point of being recurved to lie juxtaposed to the
cardiac vestibule. Therefore, the stomach assumes a C-shaped configuration

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when viewed from the front. This striking elongation and recurvature of the
pyloric stomach in Leptonycteris and in Lichonycteris as well (and to a lesser
extent in Choeroniscus and Glossophaga) might represent an adaptation to
permit intake of an increased percentage of plant material in the diet. Increased
length of the pyloric tube is one way to increase gastric volume.

The stomachs of Choeroniscus godmani and Lichonycteris obscura (Fig. 2d)
possess well-developed cardiac vestibules and broad terminal portions that
can be recurved sharply toward the gastroesophageal junction. The fundic
caeca of these two species are shallow; unlike the other species of glossophagines,
the caeca are not delineated by a sulcus ( = incisura cardiaca) from the cardiac
vestibule. Therefore, the vestibule merges gradually into the caecum on the
greater curvature of these two.

Carolliinae

The stomachs of two species from this subfamily have been examined.
Carol lia perspicillata (Fig. 2e) and C. castanea generally are quite similar
but apparently are individually variable in gross morphology. The terminal
( = pyloric) portion is elongate and strongly recurved anteriorly. This recurvature
possibly functions to retard gastric emptying. A cardiac vestibule is present;
in some specimens it is moderately developed, whereas in others it is quite small.
The caecum is baglike and dilated and is more prominent in C. castanea than
it is in C. perspicillata. Overall, the stomachs of these two species are in many
ways intermediate between those of glossophagines and those of stenodermines.
The Carolliinae exhibit the overall simplicity of most glossophagine stomachs
in combination with some specialization of the caecum (especially the pyloric
tube), which is characteristic of fruit-eating stenodermines.

Stenoderminae

An extensive array of stenodermine species, most of which are considered
to be frugivores, have been studied. The stomachs of stenodermines are
substantially more complex and more specialized than those of the previously
described species. Virtually all gross features of the stomach are enlarged or
lengthened, especially in comparison with the simpler stomachs of the
phyllostomatines and glossophagines.

The stomachs of Sturnira lilium and 5. ludovici (Fig. 3a) are similar to one
another. In S. lilium, which is typical, the cardiac vestibule is elongate and tapers
so that the gastroesophageal junction lies well superior to the gastroduodenal
junction. The fundic caecum is saccular and thinwalled, forming a spacious
chamber with an apex that varies from being rounded to being tapered. A fold
of the stomach wall distinguishes the cardiac vestibule from the fundic caecum.
The tubular ( = pyloric) portion of the stomach is long and narrow (S. ludovici
has a shorter pylorus and a somewhat larger cardiac vestibule giving the stomach
a more robust appearance than that of S. lilium). The stomach from a single

BIOLOGY OF THE PHYLLOSTOMATIDAE

211

Fig. 3. — Semidiagramatic representations of the stomachs of selected stenodermines.
The hatched area indicates the region of pylofundic transition glands: a, Sturnira ludovicr,
b, Uroderma magnirostrunr, c, Artibeus lituratus ; d, Centurio senex ; e, Vampyrodes
caraccioli ; f, Chiroderma villosum. Scale is 10 mm. Symbols are identified in Fig. 2.

specimen of S. mordax was examined by Rouk (1973) who found it to have a
a considerably simpler gross morphology than those of other species of
Sturnira. Rouk (1973) reported that the terminal portion was relatively
unspecialized and that the caecum was poorly developed. However, the
stomach in S. mordax does possess a moderately large cardiac vestibule.

The remaining stenodermines for which stomachs have been examined show
increased specialization by way of elongation or enlargement of one or more
portions of the stomach. The stomachs of seven species of Artibeus ( aztecus ,
inopinatus, jamaicensis, lituratus, phaeotis, toltecus, and watsoni) have been
studied (see Fig. 3c). These seven, along with that of Centurio senex (Fig. 3d),

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have tremendously enlarged cardiac vestibules that permit temporary storage
of large amounts of plant material. In Vampyressa, Vampyrops, Uroderma,
Vampyrodes, and Chiroderma, the cardiac vestibule varies from small to
moderately large, with the fundic caecum being variously drawn out into a baglike
or nearly tubelike structure.

Stomachs of Uroderma bilobatum and U. magnirostrum (Fig. 3b) share
gross characteristics with Sturnira, as well as with Artibeus, and could be
said to be intermediate between the two. The caecum is elongate and narrowed
nearly to a point at its apex. The stomachs of Vampyrops helleri and V. vittatus
differ from that of Uroderma only slightly in that the caecum of V. helleri
and V. vittatus is somewhat broader.

The stomachs of Vampyressa pusilla and V. nymphaea are nearly identical
to one another. The cardiac vestibule is small in comparison with most of the
other stenodermines. The elongate fundic caecum is recurved anteriorly, as
it is in Uroderma, Vampyrops, Vampyrodes, Chiroderma , and some Artibeus,
and it is dilated at its terminus.

The remaining two species to be discussed in this account, Vampyrodes
caraccioli (Fig. 3e) and Chiroderma villosum (Fig. 3f), possess greatly
enlarged fundic caeca. The stomach of Vampyrodes somewhat resembles that
of Uroderma except that the cardiac vestibule is much reduced. A distinctive
narrowing occurs between the cardiac vestibule and fundic caecum of both
species so that there is only a small region where the two are contiguous. The
fundic caecum of Vampyrodes is about 1.5 times the length of the remainder
of the stomach, and that of Chiroderma is in excess of twice the length.

The stomach of C. villosum, which has a tubular caecum, represents perhaps the
most extreme specialization for plant feeding in the Phyllostomatidae. This con¬
dition closely parallels that observed in some Old World megachiropterans. The
caecum is marked externally by a series of parallel constrictions that surround it
for nearly its entire length. The duodenum at the gastrointestinal junction is unusual
in being grossly dilated on the lesser curvature to produce what amounts to a small
ampulla or caecum. The function of this dilation is unknown.

It would appear that there are two adaptive trends within the Stenoderminae.
Each apparently represents a different response to increased need for stomach
volume in these frugivores. One trend, which is best illustrated in Artibeus and in
Centurio, was to increase size of the cardiac vestibule while minimizing the impor¬
tance of the fundic caecum. The other approach, seen so vividly in such genera as
Vampyressa, Vampyrodes, and Chiroderma, was to minimize, or even to nearly
eliminate, the cardiac vestibule while correspondingly enlarging the caecum into
an obviously useful storage chamber. Both trends would permit increased con¬
sumption or storage, or both, of plant materials that presumably are difficult to
digest.

Phyllonycterinae

Rouk (1973) examined the stomach of only one member of this subfamily,
Brachyphylla cavernarum (Fig. 4a). The esophagus enters the stomach quite near

BIOLOGY OF THE PHYLLOSTOMATIDAE

213

the gastroduodenal junction. Therefore, the lesser curvature between esophagus
and duodenum is extremely short. The fundic caecum is extremely well developed
into a “bag” that appears to be nearly compartmentalized into a two-chambered
structure. The caecum bends abruptly anteriorly about midway along its length.
At this location, there is a suggestion of a sphincter, although this constriction in
the muscularis externa has not been demonstrated to have a sphincteric function.
The duodenum is quite enlarged at its junction with the stomach, which is separated
from the intestine by a distinctive constriction. The stomach of Brachyphylla
clearly is distinctive among phyllostomatids. Other phyllonycterines should be
examined to determine if this distinctive form is consistent within the group.

Desmodontinae

The gastric morphology of Desmodus rotundus (Fig. 4b) has been variously
described and illustrated by a number of workers (Huxley, 1865; Rouk and
Glass, 1970; Hart, 1971; Forman, 1972). Its simple, tubular form is
predominately an elongate caecum of generally uniform breadth that lacks
a cardiac vestibule or demonstrable pyloric portion (although pyloric glands
are present). The terminal-most part of the caecum frequently is dilated into
a thin-walled sac; the distal one-half is folded back upon the proximal one-half.
There is no conclusive evidence of any sphincters within the stomach, except
for that adjacent to the duodenum.

In Diaemus youngii (Fig. 4c), the stomach bears strong resemblance to that of
Desmodus except that the caecum may be less tubular and more conical in this
species. The terminal part of the caecum is slightly dilated. In the stomach of
Diphylla ecaudata (Fig. 4d), numerous semilunar folds within the distal one-half
of the caecum divide it into smaller compartments. The caecum, with its haustra
coli, therefore, bears strong resemblance to the colon of man. The “pouches” thus
formed in the caecum of Diphylla would tend to retard gastric emptying, important
in vampires because the stomach is specialized for absorption. Additionally, the
folds in the caecum would tend to increase the surface area to volume ratio, thereby
increasing the efficiency of absorption from the stomach.

Gastric Mucosa

The stomachs of all species of phyllostomatids are completely lined with a
glandular mucosa. There is no uncornified or cornified squamous epithelium
in the stomach. A zone, usually narrow, of mucuous-producing cardiac glands
is found at the gastroesophageal junction. A broader zone of pyloric glands,
which also are mucuous producing and which are similar in structure to cardiac
glands, are located at the gastroduodenal junction in all species. The remainder
of the mucosa is occupied by a broad region of fundic glands composed of
mucous cells, parietal cells, and chief ( = zymogenic) cells. A zone of
transitional glands that is extremely variable in length occurs between fundic
and pyloric mucosa. This transitional area is rather broad in species of the
Glossophaginae but is relatively narrow in the Stenoderminae. Species of

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Fig. 4. — Semidiagrammatic representations of the stomachs of one phyllonycterine
and three desmodontines. The hatched area indicates the region of pylofundic transition
glands: a, Brachyphylla cavernarunr, b, Desmodus rotundas, ; c, Diaemus youngii, d,
Diphylla ecaudata. Scale is 10 mm.; upper scale is for a; lower scale, all others.

Artibeus, along with Centurio and Vampyrodes, consistently have extremely
narrow “transition” zones. This narrowness of the transition zone seems to be
due to a relatively extensive proximal advancement of pyloric glands within
the pyloric tube.

Depth of the gastric mucosa varies slightly within stomachs and among
species. The mucosa is shallowest in the vampires, with fundic glands being

BIOLOGY OF THE PHYLLOSTOMATIDAE

215

only 50 to 75 micrometers in Desmodus. The gastric glands of vampires are
reduced to shallow acini in comparison to the tubular form of other species.
This is accompanied by a general reduction in all cellular constituents,
although zymogenic, parietal, argentiffin, and mucous neck cells all are
present. Mucous neck cells comprise the most abundant cellular component
of the mucosa, whereas parietal ( = HCl-producing) cells are extremely sparse.

The gastric mucosa of other species varies from 100 to 600 micrometers,
in depth, although 200 to 250 micrometers is most commonplace. Pyloric
glands frequently are longer than are the fundic glands within a species; for
example, in Artibeus they are 50 to 80 per cent longer. In many species, the
fundic glands are somewhat longer at the apices of rugae than on the stomach
wall proper. In striking contrast is the fundic portion of the mucosa of
stenodermines, such as Artibeus and Centurio, in which the glands are of
extremely uniform depth. Relative constancy of cell frequency accounts for the
uniformity of mucosal depth. In some phyllostomatines, especially Micronycteris
and Chrotopterus, the fundic mucosa is quite shallow at the apex of the caecum.

The stomach wall of all species is thrown into rugae, which occur everywhere
within the stomach. These folds generally are oriented along the longitudinal
axis and are arranged in parallel rows in the terminal, tubular stomach. They
occur in wavy, parallel rows throughout the remainder of the stomach in many
other species. In stenodermines, all species that have been examined with respect
to rugal organization reveal some degree of “complication” or interdigitation
of folds. In Vampyressa, V ampyrops, Chiroderma, and Sturnira, they are
distributed diagonally (toward the pyloric sphincter), but only within the caecum.
Rugae are slanted only within the midregion of Uroderma. In most stenodermines
that have been studied, folds interdigitate only to a moderate degree, but in
Artibeus and Centurio an extremely complex interlocking of folds produces
an elaborate maze because folds are highly branched. This arrangement likely
would be effective in retarding gastric emptying, a particularly important
digestive adaptation in obligate plant feeders.

Histochemistry of the Gastric Mucosa

Few systematic groupings of mammals have been examined comparatively
with respect to the histochemistry or cytochemistry of the stomach lining.
Phyllostomatids are an exception to this in that the mucous cells and their
secretory products have been studied with a variety of techniques. Procedures
have been employed that elucidate acid as well as neutral mucopolysaccharides.

A positive periodic acid-Schiff (PAS) reaction is thought to indicate an
abundance of mucosubstance and, thus, it provides an overall estimate of
the quantity of mucus within or on the surface of cells in the stomach or intestine
(see Lillie, 1965). In all examined species phyllostomatids, there is a moderate
to intense coloration of mucous material in the apical portion of the cytoplasm
of surface columnar cells. In Desmodus rotundus (the only desmodontine exam¬
ined to date), the intensity of this reaction in surface mucus is somewhat reduced

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in comparison with that of other phyllostomatids. In many species having well-
developed fundic caeca, the staining is stronger in the foveolae of the fundic
glands of the caecum than elsewhere in the fundus. Mucus possibly accumulates
to a greater extent in the caecum than elsewhere in phyllostomatids.

Mucous cells beneath the surface (the so-called mucous neck cells), which are
scattered among the parietal cells, react much more variably to the PAS reaction
than do the surface columnar cells. Mucous neck cells of frugivorous species
generally are less reactive than are those of carnivorous and omnivorous kinds.
Those of Desmodus (and perhaps the other desmodontines) react only weakly.

The upper portions of the tubules of cardiac and pyloric glands stain intensely
with PAS. There is only slight variability among species. As in the case of fundic
glands, reactivity in these upper portions is somewhat reduced in frugivorous
species. Among studied species, the most intense reaction has been found in an
omnivore, Phyllostomus discolor. The quantity of gastric mucus in this species
exceeds that of frugivorous phyllostomatids. On the other hand, in Desmodus
the reactivity is weak in comparison with nondesmodontine phyllostomatids.

Two procedures, or their variants, have been employed in an effort to elaborate
the relatively acidic components of gastric mucus in phyllostomatids. Forman
(1972) employed Alcian blue 8GX, and Rouk (1973) and Forman (19716)
used Hale’s colloidal iron procedure in efforts to categorize acid
mucopolysaccharides in stomachs of selected species of phyllostomatids.
A summary of their results is presented here.

Acid mucopolysaccharides are found most consistently in the cardiac glands
(those at the gastroesophageal junction) and within the few transitional and
fundic glands adjacent to the cardiac glands. Nearly all species of phyllostomatids
studied to date showed some positive staining of cardiac glands. The only
exceptions are species of Sturnira (including S. lilium, S. ludovici, and S.
mordax ). In these species, the cardiac glands are either weakly reactive or
non reactive to procedures intended to demonstrate the presence of acid
mucopolysaccharides. Present evidence also suggests that Centurio and Desmodus
have reduced amounts of acid mucopolysaccharides in their cardiac glands.
The reaction of the pyloric glands to Hale’s colloidal iron and Alcian blue is
similar to that of the cardiac glands. There is, however, less consistency among
species, less uniformity within the zone of pyloric glands, and often less intensity
in comparison to the histologically similar cardiac glands.

In most species of phyllostomatines, the pyloric glands are nonreactive;
the exception is Vampyrum spectrum, in which these glands are weakly
reactive with Hale’s colloidal iron.

In the glossophagines, there are two general conditions of stainability of the
pyloric glands with Alcian blue and Hale’s colloidal iron. With Hale’s iron
(as employed by Rouk, 1973) pyloric glands stain intensely within the basal
one-third of the tubules in Glossophaga soricina and Lonchophylla robusta.
Forman (19716) studied glossophagine cardiac glands with Alcian blue. In his
study of five species of glossophagines, the lower portion of each pyloric gland
tubule was Alcian blue positive in three ( Glossophaga soricina, G. commissarisi,

BIOLOGY OF THE PHYLLOSTOMATIDAE

217

and Anoura geoffroyi) but negative in two others ( Choeroniscus godmani and
Lichonycteris obscura).

Among the phyllostomatids, the most widespread and distinctive reactivity
to procedures for acid mucopolysacchardies in the stomach are found in certain
of the carolliines and stenodermines. For example, pyloric glands in Vampyrodes,
Vampyressa, Chiroderma, Centurio, and in seven species of Artibeus that have
been studied, react intensively with Hale’s colloidal iron either throughout or
nearly throughout the length of the tubule. Rouk (1973) determined that nearly
all glands in the stomach of Vampyressa pusilla contain noteworthy amounts
of Hale positive mucin. In these same stenodermines, as well as in Uroderma,
V ampyrops, and Sturnira mordax, the mucous neck cells within the upper
portions of fundic gland tubules also react moderately or strongly with Hale’s
iron. Reactivity in these cells rarely has been observed in nonstenodermines.

These results suggest that a relationship might exist between gastric acid
mucopolysaccharides and plant feeding in phyllostomatids. Whether their
function is protective, digestive, or both remains to be determined.

Pyloric Sphincter

The muscular portion of the sphincter at the gastroduodenal junction is
unusually variable in form in phyllosotmatids. Numerous variations in
the form of this circular muscle mass have been observed in leaf-nosed bats,
and at least part of this variability appears to be related to diet. The sphincter
is in some way asymmetrical in the majority of species that have been examined.
In kinds where asymmetry is present, the valve on the greater curvature is larger
than that portion on the lesser curvature. This condition always prevails in
insectivorous and carnivorous species. The valves of Macrotus, Micronycteris,
Tonatia minuta , and Glossophaga are short to moderate in length and generally
are robust with broadly rounded apices. In Centurio, the valve of the greater
curvature is fully three times the mass of the “lesser” valve. This form of valvular
asymmetry is maximized in Tonatia minuta in which the greater valve is long and
extremely thick, whereas the lesser valve is absent, or nearly so. Two noteworthy
instances in which the valve is greatest in mass on the lesser curvature are found in
Uroderma bilobatum and in Chiroderma villosum. This asymmetry might result
in some sort of “milking” action that permits slow release of stomach contents into
the duodenum.

Two trends in pyloric sphincter morphology are evident in frugivorous
species as well as in some pollenivorous and nectarivorous kinds. One trend
involves increased symmetry, whereas the other involves the amount of muscular
contribution to the valve.

First, the pyloric valve of some fruit-eating stenodermines and carolliines,
including Artibeus, Sturnira, Vampyressa, Carollia perspicillata and perhaps
others, is of nearly uniform length throughout its circumference. It would
appear that increased symmetry of the valve in these species is related to
consumption of plant material. None of the insectivorous or carnivorous kinds
has a symmetrical valve; indeed, the most pronounced asymmetry always is

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observed in these species. The pyloric valve of Desmodus rotundus is reduced
in bulk, as compared with other phyllostomatids, but it also is nearly symmetrical.
It is possible that symmetry may be related to passage of liquid food into the
duodenum, both in vampires and in plant feeders.

Second, bats that consume plant material including fruit, nectar, and
pollen have a valve flap that nearly always is longer and thinner than valves
of bats that eat animal material. This feature is particularly well developed in
stenodermines and in Brachyphylla cavernarum. In species of Artibeus,
Centurio, Chiroderma, Uroderma, Vampyressa, Vampyrops, and Vampyrodes,
the flap achieves such length that its apex is directed up into the duodenum.
This results in valve flaps that are parallel with the intestinal wall. In addition,
the apex of the muscular flap is quite pointed in species of Artibeus. Most
glossophagines that have been examined, including species of Lonchophylla,
Lichonycteris, Choeronycteris, and Hylonycteris, but excluding Glossophaga,
have thin valves that are similar to those of stenodermines. Anoura and
Leptonycteris are intermediate between the Glossophaga- type and stenodermine-
type valve, but most similar to the latter. It is reasonable to hypothesize that
these longer, thinner, often symmetrical valve flaps might improve the
efficiency of gastric closure, thus delaying gastric emptying and improving
digestion (by increasing time) in these plant feeders.

The pyloric sphincters of Sturnira lilium and S. ludovici, although symmetrical,
are unique in that identifiable muscular flaps either are absent or nearly so as
barely to be perceptable. The functional significance of this apparent degeneracy
is unknown.

Tunica Muscularis

All stomachs of phyllostomatids possess two layers in the tunica muscularis,
an outer longitudinal and an inner circular one. An extremely thin muscularis
mucosae occurs just inside the external tunic. It is separated from the outer
musculature by an extremely sparse complement of loose submucosa. Both
external muscle layers often are variably thicker on the greater curvature than
on the lesser curvature. The musculature generally is thicker in phyllostomatines
and phyllonycterines than in the other subfamilies.

Considerable variability in the relative thickness of the two outer layers has
been observed within the stomachs of phyllostomatids. In most species, the
layers are subequal, with the circular layer being the more robust of the two.
The circular layer is not infrequently organized into bundles, cross-sections of
which are easily viewed in longitudinal stomach sections. This “bundling” is
most pronounced in the caecum (when present) where it is prominent in the
greater curvature in the majority of stenodermines that have been examined.
In a variety of leaf-nosed bats, particularly glossophagines and stenodermines,
these bundles are particularly thick just beneath the folds ( = rugae) in the stomach
lining. In Chiroderma villosum, circumferential, parallel, external constrictions
occur in the elongate caecum as a result of the distinctively thickened circular
bands beneath the rugae.

BIOLOGY OF THE PHYLLOSTOMATIDAE

219

The circular layer clearly is the dominant portion within the aboral pyloric
tube of nearly all species. Macrophyllum macrophyllum is a noteworthy
exception because in this species the aboral circular layer is thinner than
elsewhere in the stomach. In stenodermines, the pyloric circular layer thickens
progressively from cardiac vestibule to pyloric sphincter.

Species that feed predominantly or exclusively on plant material have
enlarged cardiac vestibules and fundic caeca. This development of “sub¬
compartments” is accompanied by a progressive reduction in the thickness
of the muscularis extemis in the enlarged areas. In species that apparently
are omnivorous (for example, Glossophaga soricina, Phyllostomus discolor ,
and species of Micronycteris ), the muscularis externis is reduced in thickness
in the apex of the caecum. Such a reduction could be regarded as an intermediate
condition or as reflective of a trend toward a frugivorous diet.

Intestine

Bats most often have short, small intestines in comparison to other kinds of
small mammals. Most comparative measurements of intestinal lengths in
bats (see Eisentraut, 1950; Robin, 1881) have revealed that frugivores usually
have relatively long intestines (in relation to body length) when compared to
insectivorous, carnivorous, or nectarivorous species. This finding applies to
Phyllostomatidae as well as to the Microchiroptera in general.

Eisentraut (1950) noted that of numerous species of bats with a variety of
feeding habits, those with an intestinal length greater than four times the body
length always were fruit-eating phyllostomatids, and that others had intestines
of relatively lesser length. Among species with the longest intestines (relative
to body length) are Chiroderma villosum, Vampyrops vittatus, and several
species of Artibeus and Brachyphylla (Forman, unpublished data). Vampires
have intestines of moderate length. Based on only scattered and incomplete
data, those few glossophagines for which measurements are available generally
have relatively short intestinal tubes.

In general morphological features, the intestine differs little from that of
most other groups of small mammals. Both “small” and “large” intestinal
segments are present and a short duodenum is distinguished by noteworthy
breadth. One noteable feature, shared with other groups of bats, is the lack of
an ascending or transverse colon so that the large intestine is restricted to a
relatively short descending colon.

A caecum always is lacking. However, at the junction of small and large
intestines there frequently is a small ampulla formed as a result of a hypertrophic
dilation of the muscularis externa. Abundant lymphoid tissue (nodules of
Peyer’s patches) always are present within the ampulla, which is displaced
well away from mesenteric attachment to the gut (Forman, 1974a, 19746).
This ampulla first was observed in Carollia perspicillata (Schultz, 1965).
Schultz likened this “protrusion,” in size and location, to the abbreviated
ileocolonic caecum in species of the Old World microchiropterans Rhinopoma
and Megaderma.

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Fig. 5. — Scanning electron micrograph of intestinal villi in the middle portion of the
small intestine of Phyllonycteris aphylla. The arrow indicates a plane of orientation of
rows of villi that is diagonal with respect to the intestinal wall. Note the generally pyramidal
shape of each villous and that villi in one row lie between villi in an adjacent row when
they are viewed directly from left to right. Scale is 0.25 mm.

Although an ileocaecal valve is lacking in all species that have been examined,
Schultz (1965) noted the presence of a valvelike flap within the middle portion
of the intestine in Diphylla ecaudata. Whether this structure functions as a valve
is not known.

The complete gastrointestinal tracts of six species of phyllostomatids were
figured by Schultz (1965) in his monograph on blood vessel supply to the digestive
tract of bats. Several of his figures of the gut of Glossophaga soricina reveal an
extremely complex “looping” of the intestine in this species. The first loop of
the intestine is joined to the terminal portion of the ileum by a mesenteric
ligament. The intestine then proceeds into considerable looping, the extent of
which is a function of intestinal length. The attachment of the first intestinal
loop to the terminal ileum by a ligament also was illustrated by Schultz in a
figure of the gut of Carollia perspicillata.

Torsion is extensive in the intestine of most phyllostomatids. In Carollia and
Glossophaga, it is as much as 270° (Schultz, 1965). In most phyllostomatids,
the intestine is considerably displaced to the right within the abdominal cavity.
One exception is Macrotus californicus, in which the intestine is not displaced.

BIOLOGY OF THE PHYLLOSTOMATIDAE

221

Fig. 6. — Surface view of intestinal folds ( = villi) in one specimen of Artibeus
jamaicensis. Note the complex interdigitation and maze-like organization of ridges. Also,
note the narrow channels (C) within the intestinal epithelium. These are most distinctive,
in breadth and depth, at the angles or bends of the intestinal folds. Scale is 0.5 mm.

The small intestines of vampires ( Desmodus and Diphylla ) are not grossly
different from those of other phyllostomatids. However, twisting is only slight,
and most of the intestine is folded back upon itself in a series of numerous compact
winding folds.

The topography of the mucous membrane of the large intestine generally
is uniform among the few species that have been examined. Folds are longitudinal
and usually have smooth surfaces with an abundance of goblet cells.

Considerable variation in the topography of the mucous membrane of the
small intestine occurs within and among species of phyllostomatids. Projections
of the membrane into the lumen can be in the form of fingerlike villi, nearly
continuous transverse folds, or projections of a form to some degree intermediate
between the other two extremes. Although variation is extensive, a review of
the literature, along with some observations of gut morphology in phyllostomatids
by the senior author, reveals one apparent pattern of villous distribution within
the family. This pattern occurs most consistently within fruit-eating species.
Fingerlike villi, if present, usually are located within the distal-most portion
or ileum. As one progresses upward toward the gastroduodenal junction,
“pyramid-shaped” projections, which are oriented in transverse rows, become

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Fig. 7. — Surface view of intestinal folds ( = villi) in one specimen of Carollia
perspicillata. Scale is 0.5 mm.

Fig. 8. — Surface view of intestinal folds ( = villi) in one specimen of Chrotopterus
auritus. Note the simplicity of folds as compared with those of Artibeus jamaicensis and
Carollia perspicillata. Scale is 0.5 mm.

BIOLOGY OF THE PHYLLOSTOMATIDAE

223

Fig. 9. — Several forms of villi observed within the small intestine (middle portion)
of Mcicrotus waterhousii. Arrows indicate the presence of a groove on the surface of
some villi. All villi are drawn to scale.

abundant and increase in lateral dimension. These projections are distributed
in rows that assume a zig-zag configuration when viewed from the top. The
zig-zags in most kinds become progressively more flattened from the middle
portion of the intestine through the duodenum. Also, the transverse folds or
“pennant-shaped villi” (after Schultz, 1965), which interdigitate with and are
interrupted by one another within the lower portions of their distribution,
often loose much of this complexity in the upper portions of the small
intestine.

The most detailed descriptions of intestinal mucous membrane topography
of phyllostomatids are those of Mathis (1928) and Schultz (1965). Mathis

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described the villous pattern in Phyllostomus hastatus and Glossophaga
soricina and reported that in his view villi, as such, were lacking in portions
of the intestine in Phyllostomus. Also, the broad villi in the uppermost
intestine were set in oblique rows. This latter pattern also occurs in other species
within the family (see Fig. 5). Mathis reported that the villi of G. soricina
in some areas can be tightly compact without any arrangement into rows.
Digitate or club-shaped villi may be interspersed among “transverse folds”
and be of somewhat greater height than the folds. Schultz’s (1965) description
of villous morphology in G. soricina generally agrees with that of Mathis
(1928); Schultz further stated that the configuration in Anoura geoffroyi is
“just as with G. soricina .” The extent to which the pattern as observed in these
two species can be applied to other glossophagines is unknown.

Intestinal villi of the fruit-eating stenodermines frequently are arranged in
extremely elaborate interdigitating networks (see Fig. 6). This complicated
arrangement likely helps to impede transport of food. Other fruit-eating
species have less elaborate villous arrangements (Fig. 7). One carnivorous
kind (Fig. 8) has villi uncomplicated in cross-sectional configuration and
nearly fingerlike in their appearance.

Villi often are arched from side to side. This feature in combination with
staggered arrangement of villi in oblique rows produces a mechanism for
entrapment of food material between villi at their bases. This likely results
in improved food assimilation or absorption inasmuch as food would tend
to be retained in the small intestine for longer periods of time.

Some variability in the structure of villi occurs within localized portions
of the small intestine in phyllostomatids. Villi within the middle portion of the
intestine of Mac rot us waterhousii can have narrowly rounded (Fig. 9 a) or
relatively broadly rounded apices (Fig. 9c). The “arching” of villi, with
subsequent entrapment of food material, might be augmented by an apparent
groove on the superior surface of some villi (see Figs. 9c, h). Food could become
trapped at the base of these folds.

The intestinal topography of Desmodus is not known to be particularly
specialized. Villi are known to be present in the intestines of both Desmodus
and Diphylla (Schultz, 1965) but generally are not fingerlike, and they are
arranged in a pattern of interdigitation. Rouk and Lane (1970) reported that
the crypts of Lieberkuhn appear to be reduced in comparison to other species.

The types of cells present within the small intestine of phyllostomatids
essentially are the same as those of other groups of bats and other eutherians.
The Paneth cells of bats have been examined by Schaaf (1970) in relation to
food habits. Schaaf s study group included three insectivorous species as
well as Artibeus jamaicensis, Bachyphylla nana, Phyllonycteris poeyi, and
Monophyllus redmani. The results of selected histochemical tests were
uniform for prosecretion granules and mucopolysaccharides in all species.
Strong acidophilia was present in the cells indicating the probable presence
of lysosomes. Secretion granules contained a mixture of protein and carbohydrates.
The results agree well with those for other species of mammals. Therefore,

BIOLOGY OF THE PHYLLOSTOMATIDAE

225

Paneth cells presently are not known to be specialized to permit the assimilation
of large quantities of any particular food material by phyllostomatids, for
which food habits are highly varied but generally obligate.

The glands of Brunner are mucus producing and generally restricted in
distribution to an extremely narrow submucosal ring at the gastroduodenal
junction. Several unusual conditions with respect to Brunner’s glands occur
within the Phyllostomatidae. These conditions might relate to the varied
food habits that occur within the family.

The stomachs of Sturnira lilium and S. ludovici have cells within the bases
of the pyloric glands that are histologically identical to the submucosal glands
of Brunner within the uppermost duodenum. Several species of Artibeus
(Forman, 1972; Rouk, 1973) have similar cells within their pyloric stomachs.
Cells of Brunner’s glands in the duodenum and those cells at the base of
pyloric glands stain identically with the periodic acid Schiff reaction for neutral
mucopolysaccharides. This staining is considerably different from that
within remaining cells of the pyloric glands. Cells such as those of Brunner’s
glands may provide for better protection of the pyloric mucosa from large
amounts of hydochloric acid that likely are produced by the considerable
number of parietal cells in some fruit-eating phyllostomatids.

Of those studied, the Brunner’s glands of Phyllostomus hastatus and P.
discolor are best developed. Other species of phyllostomatines (those of
Tonatia, Micronycteris , and Chrotopterus ) have relatively numerous Brunner’s
glands but they nevertheless are less distinctive than are those of Phyllostomus.

The numerous species of stenodermines, carolliines, and some species of
glossophagines are in marked contrast to the phyllostomatines. Although
only a few species of Artibeus have been examined, it is known that the
Brunner’s glands of A. lituratus and A. jamaicensis are extremely sparse in the
most proximal portion of the duodenum and that they are absent in at least
some specimens of Artibeus phaeotis and in A. inopinatus. It is reasonable to
hypothesize that other species of Artibeus harbor few of these glands. In addition
to species of Artibeus, the following bats have been reported to lack Brunner’s
glands at the gastroduodenal junction: Centurio senex, Chiroderma villosum,
Uroderma bilobatum, Vampyrodes caraccioli, Vampyressa pusilla, V. nymphaea,
and V ampyrops helleri. Artibeus toltecus and Vampyrops vittatus are reported to
have numerous Brunner’s glands at the gastrointestinal junction. The basal cells
of the pyloric glands in Centurio senex are histologically similar to the Brunner’s
glands of Artibeus lituratus. Also, it is noteworthy that all species of stenodermines
that lack Brunner’s glands in the upper duodenum, except for Chiroderma, have
relatively extensive zones of pyloric mucosa in the stomach. It is reasonable at this
point to suggest that the pyloric mucosa in these animals may be performing the
“neutralization” action on the food bolus that ordinarily is believed to be performed
by the glands of Brunner in other species of mammals.

Additionally, several species of nectar-feeding glossophagines ( Lichonycteris
obscura and Choeroniscus godmani ) have been observed to have few Brunner’s
glands (Forman, 1971a). The only phyllonycterine that has been examined,

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SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

Brachyphylla cavernarum, has no glands of Brunner. These observations,
along with those on stenodermines, clearly indicate that the conditions in
certain phyllostomatids do not support the widely held view that mammals
consuming plant material have more abundant glands of Brunner than do
animals eating animal material.

The connective tissue of the intestine of bats generally is extremely
sparse. The intestine of Desmodus rotundus (and perhaps the other two
sanguivorous species) is a noteworthy exception. Both the submucosa and the
lamina propria of the villi are unusually thick and dense. They are highly
vascularized and harbor a considerable lymphatic network.

Studies of organized gut-associated lymphoid tissue (Peyer’s patches) in
New World bats (Forman, 1974a, 19746) have revealed differences in
abundance, distribution, and morphology of this tissue within the
Phyllostomatidae. These differences possibly relate to diet. For example, fruit¬
eating species usually have the most patches when compared with nectarivorous
or with carnivorous and insectivorous kinds. Also, the patches can occur
almost anywhere along the length of the small intestine in fruit eaters, frequently
including the duodenum. These patches have relatively large nodules with
extremely large geminal centers. The patches and nodules of insect eaters
and carnivores, in contrast, are relatively small with small germinal centers
typically indicating a low state of activity. Patches in these species usually are
restricted to the submucosa of the ileum.

These observations suggest that at least within the family Phyllostomatidae
organized lymphoid tissue within the gut might be differentially responsive to
intestinal contents including food material and associated microbial populations
as well.

Literature Cited

Allison, A. C. 1948. The stomach in South African Insectivora, with notes on the
organization of mammalian gastric glands. J. Anat., 82:249-261.

Eisentraut, M. 1950. Die Ernahrung de Fledermausen (Microchiroptera). Zool.
Jahrb., Jena., 79: 1 14-177.

Forman, G. L. 1971a. Gastric morphology in selected mormoopid and glossophagine
bats as related to systematic problems. Trans. Illinois Acad. Sci., 64:273-282.

- . 19716. Histochemical differences in gastric mucus of bats. J. Mamm.,

52:191-193.

- . 1972. Comparative morphological and histochemical studies of stomachs

of selected American bats. Univ. Kansas Sci. Bull., 49:591-729.

- . 1973. Studies of gastric morphology in North American Chiroptera

(Emballonuridae, Noctilionidae, and Phyllostomatidae). J. Mamm., 54:909-923.

- . 1974a. Comparative studies of organized gut-associated lymphoid tissue in

mammals with diverse food habits. Distribution, size, and organization of
Peyer’s patches in New World bats. Trans. Illinois Acad. Sci., 67:152-156.

- . 19746. The structure of Peyer's patches and their associated nodules in New

World bats in relation to food habits. J. Mamm., 55:738-746.

Hart, L. A. 1971. Structure of the gastric mucosa as related to feeding habits in
selected species of New World bats. Unpublished Ph.D. dissertation,
Virginia Polytechnic Inst., 65 pp.

BIOLOGY OF THE PHYLLOSTOMATIDAE

227

Gardner, A. L. 1977. Feeding habits. Pp. 293-350, in Biology of bats of the New
World family Phyllostomatidae. Part II (R. J. Baker, J. K. Jones, Jr., and D. C.
Carter, eds.). Spec. Publ. Mus., Texas Tech Univ., 13:1-364.

Huxley, T. H. 1865. On the structure of the stomach in Desmodus rufus. Proc.

Zool. Soc., London, pp. 386-390.

Kolb, A. 1954. Biological Beobachtungen an Fledermausen. Saugetiekundl. Mitt.,
2:15-26.

Lillie, R. D. 1965. Histopathologic technic and practical histochemistry. McGraw-
Hill, New York, xii + 775 pp.

Mathis, J. 1928. Beitrag zur Kenntnis des Fledermausdarmes. Z. Mik. Anat.

Forsch., 12:594-647.

Moller, W. 1932. Das Epithel der Speiserohrenschleimhaut der blutenbesuchenden

Fledermaus Glossophaga soricina im Vergleich zu insektenfressenden
Chiropteren. Zeit. Mikr. Anat., 29:637-653.

Myrcha, A. 1967. Comparative studies on the morphology of the stomach in the

Insectivora. Acta Theriol. 12:223-244.

Phillips, C. J., G. W. Grimes, and G. L. Forman. 1977. Oral biology. Pp. 121-246,
in Biology of bats of the New World family Phyllostomatidae. Part II (R. J.
Baker, J. K. Jones, Jr., and D. C. Carter, eds.). Spec. Publ. Mus., Texas Tech
Univ., 13:1-364.

Robin, H. A. 1881. Recherches anatomique sur les mammiferes de l’ordre des Chi-
ropteres. Ann. Sci. Nat. Zool., 12:1-180.

Rouk, C. S. 1968. Comparative gastric histology of selected American bats. Unpublished
M.S. thesis, Oklahoma State Univ., 51 pp.

- . 1973. Gastric morphology and adaptive radiation in the Phyllostomatidae.

Unpublished Ph.D. dissertation, Texas Tech University, 85 pp.

Rouk, C. S., and B. P. Glass. 1970. Comparative gastric histology of American bats.
J. Mamm., 51:455-472.

Rouk, C. S„ and W. L. Lane. 1970. Comparative histology of intestines of selected
American bats. Abstracts of Papers Presented at the 50th Annual Meeting,
American Society of Mammalogists, College Station, Texas, June.

Schaaf, V. P. 1970. Untersuchungen uber das histochemische Verhalten der Panethschen
Kornerzellen bei mittelamerikanischan Fledermausarten mit unterschiedlichen
Ernahrungsweisen. Anat. Anz. Bd., 126:275-277.

Schultz, W. 1965. Studien uber den Magen-Darm-Kanal der Chiropteren. Ein
Beitrag zum Problem der Homolgisierung von abschnitten des Saugetierdams. Ziet.
Wissenschaft. Zool., 171:241-391.

MORPHOMETRIC ANALYSIS OF
CHIROPTERAN WINGS

James Dale Smith and Andrew Starrett

Bats are unique among mammals in their possession of wings. The evolution
and adaptation of these anatomically complex structures along with the develop¬
ment of an acute ability to orient acoustically has contributed markedly to one
of the most interesting examples of adaptive radiation in vertebrate history. Yet
the morphometric properties of bat wings have remained poorly understood.
Biologists have described chiropteran diversity and faunal complexity throughout
the world, but the flight behavior of only a few species has been reported (see
Eisentraut, 1936; Dwyer, 1965; Kulzer, 1968; Norberg, 1970, 1976a, 1976b;
Pennycuick, 1 97 1 ; Schnitzler, 1971).

Revilliod (1916) was the first to attempt to describe the morphometries of
chiropteran wings. In this much overlooked paper, he utilized several indices
to demonstrate the degree of adaptation to flight by several families of bats. Poole
(1936) was among the earliest investigators to report wing loading values for bats,
and Struhsaker (1961) was the first to calculate aspect ratios of bat wings. Bader
and Hall (1960) were the first investigators to use computer techniques to analyze
the osteometric variation of bat wings. In this study, they employed correlation
coefficients to assess the interrelationships among the skeletal elements of the wing
and foot of Myotis lucifugus and M. sodalis.

Other studies, although important contributions, have been limited in their scope
and coverage. Among these are Vaughan’s (1959) detailed anatomical analysis
of three bat species from North America; a more recent survey of the skeletal
and muscular system and aerodynamics appears in Vaughan (1970a, 1970b,
1970c). Hartman (1963), Gaisler (1964), Farney and Fleharty (1969), and
Jones and Suttkus (1971) have reported wing loading and aspect ratios for num¬
erous species of bats. Pearson et al. (1952), Orr (1954), Short (1961), and Jones
(1967) have contributed important information relative to the growth and de¬
velopment of chiropteran wings. Seasonal changes in wing loading of several
North American species were examined by Davis (1969) and O’Farrell and
Studier (1976), and Norbert (1969, 1972) reported on functional osteology and
myology of the wings of several bats.

By far, the most extensive analysis of the morphometric properties of bat wings
is that by Findley et al. (1972). In this study, they relied on regression and
correlation procedures as well as factor analysis to examine the wings of
approximately 1 35 species. Our initial goal was to expand on this study with our
primary focus on the bats of the family Phyllostomatidae. However, it soon became
apparent to us that a meaningful interpretation of the morphometries of phyllosto-
matid wings required a broader understanding of the overall variation in size
and shape of wings in the Chiroptera.

229

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SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

Methods

Methods of deriving the form and extent of chiropteran wings for the purpose
of studying size and shape have been variable. For example, some workers have
traced the outline of the extended wing of freshly killed bats or individuals pre¬
served in alcohol. From such tracings, they have derived the area of the wing and
other aerodynamic parameters by using a planimeter or by some other time-
consuming procedure. While these efforts are to be commended, such tech¬
niques do not readily permit an overall consideration of the diversity of the
chiropteran fauna of the world.

In addition, most past studies of wing morphology have neglected to consider
the influence of the fourth digit in determining the size and shape of the wing.
Typically, the lengths of the forearm and digit III are taken to describe the span
of the wing, and the length of digit V, its width. These measurements have been
used to derive the aspect ratio and wing loading of chiropteran wings, which,
characterized in this manner, are assumed to be rectangular in shape. For deter¬
mination of wing loading, such calculations tend to result in over-estimates of
area due to the inclusion of an intrinsic portion of the rectangular shape that,
in fact, does not exist in the real wing (Fig. 1). These calculations also may lead
to mistaken estimates of similarity between markedly different wings and may
mask subtle differences between similarly shaped wings. Furthermore, most
past studies have considered only the total lengths of digits rather than examining
the variability of digital composition and its influence on wing size and shape.

In this study, 1 1 wing measurements, length of the head and body, and weight,
were obtained from 1456 museum specimens, which comprised 433 species and
147 genera from 17 families of bats. Most of these specimens were conventional
study skins, although in some cases only specimens preserved in alcohol were
available. The wing measurements included the length of the forearm (as described
by Smith, 1972) and the individual lengths of the metacarpal and phalangeal
elements of digits III, IV, and V. The length of the often curved and cartilaginous
portion of the terminal phalange of the third digit was recorded as the greatest
radius of the arc. When available, the length of the head and body and the weight
of the specimens were recorded from the specimen label. Head and body length
was measured directly on specimens preserved in alcohol. The weights of many
specimens, especially those in alcohol, were not recorded at the time of capture.
In these cases, weights were estimated (see below). All measurements were re¬
corded in millimeters (by means of dial calipers, calibrated in twentieths of a
millimeter) or grams.

Derived Variables

At the outset of our analysis, we, like many others before us, converted our raw
variables, a priori , into a number of derived variables such as aspect ratio,
wing loading, tip index, and so forth. The subsequent analysis of these derived
variables was beset with a number of problems. Foremost among these were
inflated correlations, which resulted from linear dependence of the derived
variables. This resulted in obscuring the sources of dependency. Atchley (1978),

BIOLOGY OF THE PHYLLOSTOMATIDAE

231

Fig. 1. — Diagrammatic comparison of an actual wing and the construct of the wing
(stippled area) used in this study. The dotted line indicates the assumed shape of the wing
if only the length (forearm plus digit III) and width (digit V) are considered.

Atchley and Anderson (1978), Atchley et al. (1976), and Pimentel (1978) re¬
cently presented discussions regarding the statistical properties of derived vari¬
ables such as ratios and indices. Although derived variables can be useful in some
cases, they should be scrutinized closely and avoided when possible. Because the
goal of our investigation was to examine, insofar as possible, the interactions
among wing components and because these interactions were largely masked by
the difficulties noted above, we chose to analyze only our original raw variables.
However, after these analyses were completed ( a posteriori), we found that some
of our derived variables could be used in a generalized descriptive sense. Those
which were found to be most useful are presented in the Appendix (Tables Al-
A21) and are described below.

Weight. — This variable was essential to the computation of wing loading. To
circumvent the problem of missing data, Findley et u/.(1972, table 3) utilized
the predicting qualities of a simple linear regression to derive estimated weight
from head and body length. We also examined this relationship for 1082 specimens
using a similar regression model on known head and body length (20 and weight
( Y) and found that the residuals ( Y- Y') were lowest at the small-sized end of the
variation. However, the residuals increased markedly at the large-sized end of
the spectrum. In an attempt to reduce these overestimates, we computed a second
degree polynomial regression. This reduced the magnitude of the residuals in
the upper range of variation, but the analysis did not provide, in our opinion,
totally satisfactory results. As did Findley et al. (1972), we partitioned our data
into recognized taxonomic groups corresponding to familial and subfamilial
categories and obtained different functions for nearly every grouping (Table 1).

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SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

Table 1 . — Results of the second degree polynomial regression analysis of head and body length
(X-axis) and weight (Y-axis). Symbols are: correlation coefficient, r; Y-intercept, A; linear
regression coefficient, Bl; and quadratic regression coefficient, B2. Significant correlation
coefficients and F -values are indicated with an asterisk.

Taxon

Pteropodidae

119

0.967*

26.53

-0.892 + 0.256

0.011+0.001

928.24*

Emballonuridae

0.877*

26.89

-1.104 + 0.720

0.014 + 0.005

60.35*

Rhinolophidae

153

0.978*

2.37

-0.375 + 0.162

0.010 + 0.001

1680.94*

Nycteridae

0.651*

-45.59

1.789+1.508

-0.013 + 0.014

12.66

Megadermatidae

0.573*

-239.29

6.235 + 2.784

-0.035 + 0.017

7.46

Noctilionidae

0.925*

-218.56

5.229+1.340

-0.023 + 0.008

121.54*

Phyllostomatinae

168

0.943*

-12.33

0.024 + 0.184

0.007 + 0.001

659.23*

Glossophaginae

0.865*

24.61

-0.813 + 0.297

0.010 + 0.002

136.63*

Carolliinae

0.967*

20.76

-0.830 + 0.658

0.012 + 0.006

76.95*

Stenoderminae

128

0.955*

26.59

-1.196 + 0.476

0.017 + 0.003

678.55*

Desmodontinae

0.833*

-10.85

-0.123 + 1.898

0.008 + 0.012

44.91*

Phyllostomatidae'

391

0.912*

-12.78

0.017 + 0.130

0.007 + 0.008

1212.65*

Vespertilionidae

157

0.933*

0.45

-0.116 + 0.120

0.005 + 0.001

524.80*

Molossidae

120

0.979*

1.54

-0.227+0.092

0.006 + 0.001

1442.02*

All bats

1108

0.961*

6.52

-0.422 + 0.045

0.009 + 0.001

1990.05*

'Combined sample of the family Phyllostomatidae.

The results of our linear regression model (not shown) agreed, for the most
part, with those presented by Findley et al. (1972). We found in our regression
analyses that the regression coefficients (Bl or B2) had relatively little effect
on the slope of the line. More importantly, the F-intercept values ( A ) varied
greatly, in both our analysis and theirs, and in the majority of cases these intercept
values departed, negatively, from zero (the theoretical intercept in these analyses).
Therefore, these models predicted extremely low or even negative weights for
bats of extremely small body size. In those cases where the departure of the F-
intercept was positive, weight would be given to a bat that had zero head and
body length. An a priori manipulation of the regression model certainly might
improve the “fit” of the line, but we suspect biologic reality is quickly obscured
by such practice; biological meaning is not automatically ascribed by statistical
significance. Furthermore, we suspect that the complexity of the relationships
of weight to head and body length and other meristic parameters is more com¬
plicated than can be measured precisely with regression/correlation statistics,
and we strongly caution other investigators against placing much faith in such
predictions. With an awareness of these difficulties in mind, we utilized the
predictions of weights generated by our polynomial regression model. However,
the weight values obtained in this manner were used only to compute wing loading
for comparative purposes and these were not used in any further rigorous
analyses. In those groups where there were insufficient numbers to compute a
regression function, we utilized the function of the most closely related group
for which there was a function. All weights (actual or estimated) were converted
to Newtons (Nt).

Wing areas. — The computation of the area of the wings was necessary for the
calculations of both aspect ratio and wing loading. The area of the plagiopatagium
was calculated as the area of a rectangle (length of forearm X length of digit V).

BIOLOGY OF THE PHYLLOSTOMATIDAE

233

In deriving the area of the wing tip, we attempted to consider an attenuated
(polygonal) tip rather than a simplistic, rectangular tip as has been the practice.
To accomplish this, using measurements from museum material, we considered
a construct of the wing (Fig. 1 ) in which the fourth digit was an integral component.
We noted from empirical observations that the posture of this digit varied among
species and that estimates of the tip area varied with this posture. In addition,
we found that in most instances, when the wing was fully extended, the fifth digit
projected at approximately a right angle from the leading edge (forearm and
digit III). Although our testing of empirical data was limited, we found that we
could geometrically estimate the angle of projection of digit IV (alpha angle), with
90 per cent confidence, when the panel areas A1 and A2 (Fig. 2) were considered
to be equal or nearly equal. More precisely, the alpha angle equals the arc
tangent of (length of digit V/length of digit III). Alpha angles are given in
degrees of rotation from digit III. Bats with relatively long fifth digits tended to
possess large alpha angles, whereas those with relatively long third digits had
lower alpha angles (Table Al).

Once the alpha angle was determined, calculating the area of the two triangles
Ai and A2 (Fig. 2) was simply: area of the wing panel between digits III and IV
equals (cosine alpha angle X (length of digit III X digit IV) and area of the
wing panel between digit IV and V equals sine alpha angle X (length of digit
IV X digit V). The total area of the wing, or any portion thereof, was derived by
summing the respective areas and multiplying by 2. All areas were converted into
square meters (m2).

Wing loading. — This variable was obtained by weight (Nt)/total area of the
wing (m2). Wing loads are reported as Newtons per square meter (Nt/m2) (Table
A6).

Aspect ratio. — We followed Hartman (1963) in computing this variable: over¬
all aspect ratio — 2 (length of forearm plus length of digit III)2/total area of the
wing. We partitioned the aspect ratio into two additional ratios as follows: 1)
aspect ratio of the plagiopatagium — (length of the forearm X 2)2/area of
the plagiopatagium, and 2) aspect ratio of the wing tip — (length of digit III X
2)2/area of the wing tip. These ratios are presented in Tables A3-A5.

Tip index. — The tip of the chiropatagium is the principal propulsive portion
of the chiropteran wing (Vaughan, 1970c). The tip index (Findley et al., 1972)
is the ratio of length of digit Ill/length of forearm. A high tip index (2.00) indicates
a proportionately long third digit, whereas a low index (1.00) reflects a relatively
short wing tip (Table A2).

Relative lengths of the wing elements. — We followed Findley et al. (1972) in
computing the relative length of the wing, which is (length of forearm plus length
of digit III)Aength of the head and body. In similar fashion, we computed the
relative lengths of the forearm and digits III-V (Tables A7-A1 1).

Percentage of digital composition. — In an a priori effort to characterize the
varying composition of digits III-V, we computed the percentage that each digital
element contributed to the total length of its respective digit. These values proved
a posteriori to be useful guidelines in the interpretation of the discriminant
analysis (Tables A 1 2-A2 1 ).

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SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

Fig. 2. — Diagrammatic representation of the derivation of the alpha angle. See text
(methods) for discussion.

Statistical Procedure

Past studies of the morphometries of chiropteran wings have been rather
limited in the sophistication of their statistical analysis. Most report only simple
descriptive statistics such as mean, range, standard deviation, and in some cases,
coefficient of variation. As noted earlier, Bader and Hall (1960) and Findley
et al. (1972) have applied more detailed statistical procedures; the latter employed
both correlation and regression statistics as well as factor analysis.

In our initial statistical analysis of the morphometric properties of bat wings,
we computed such simple statistics as mean, range, one standard error of the mean,
and coefficient of variation for all variables. As noted above, these descriptive
statistics for selected derived variables are presented in the Appendix (Tables
A1-A21). In these tables, taxonomic groups are ranked by the magnitude of
their variable means (largest to smallest) rather than in phylogenetic order.
Within the family Phyllostomatidae, subfamilies were allowed to rank in this
fashion as were genera within subfamilies. The mean for “all bats” also was al¬
lowed to take its appropriate position within the familial ranking.

BIOLOGY OF THE PHYLLOSTOMATIDAE

235

We used regression and correlation analyses from BIOMED (Dixon, 1973)
and SPSS (Nie et al., 1975) in our examination of the relationships between head
and body length and weight. However, in the main portion of our study, we
employed the multivariate procedures of principal components (PCA) and
discriminant analyses to assess the morphometric interactions among the twelve
original variables and their effects on size and shape of chiropteran wings.
Descriptions of these multivariate procedures may be found in Koons (1962),
Cooley and Lohnes (1971), and Pimentel (1978). The computations of these
procedures were accomplished in the Computer Center, California Polytechnic
University, San Luis Obispo, using an unpublished program (DISANAL)
written by Richard A. Pimentel.

Interpretation of the component graphs and variable vectors. — We suspect
that many readers might not be completely familiar with the graphical repre¬
sentations that we have employed in this study. It is difficult to portray visually
the multidimensional patterns of variation computed by the multivariate statistical
procedures used in this study, which assess variation among all ^-variables in
p-dimensional space. We have used component graphs that are two dimensional
views of portions of these multidimensional spaces. In the figures beyond, we
have plotted the first and second (1X2) axes to show the length /width character
of the dispersion. Height of the dispersion is shown in the graphs in which axes
one and three (1x3) are plotted. Viewed together, each set of component graphs
depicts the dispersion of centroids in three dimensions. The coordinates used
to plot these graphs (Figs. 3, 5, 6) are given in Tables 3 and 5, respectively.

In Fig. 4, we have plotted the direction cosines (PCA) and canonical vectors
for the twelve original variables in much the same manner as described for the
component graphs. The coordinates used to plot these vectors are given in Tables
2 and 4, respectively. To avoid confusion, only the positive end of each vector
is shown. The tail or negative end of a vector passes through the ordinate of each
graph for an equal length in the opposite direction. The influence that any one
vector has on the location of the group centroids is determined by the magnitude
or length of that vector and the proximity of its point (positive end) or tail (negative
end) to the various centroids. Long vectors exert a strong influence on the location,
whereas shorter vectors exhibit weaker effects. In these analyses, an association
with the positive end of a vector implies large size (longness) and proximity to
the tail of a vector indicates small size (shortness).

It is important to bear in mind continually the fact that the overall ordination
of groups (Figs. 3, 5, 6) is the result of synergistic interplay among variables (Fig. 4)
and not the result of any one or two of these. We have attempted to illustrate and
set these figures in such a way as to facilitate the reader’s perception of the
dimensionality of the variation on the dispersion of groups. To facilitate further
an interpretation of the component graphs, the reader may wish to make a xerox
transparency of Fig. 4 and overlay this on the corresponding component graphs.
In addition, this overlay may be used to interpret Figs. 11 to 16.

236

SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

For those readers who wish to see the finer aspects of the ordination, we strongly
encourage the construction of three-dimensional models. This may be ac¬
complished easily by xeroxing the 1X2 component graphs and attaching these
to a styrofoam base. Sticks may be cut to an appropriate length by using the 1 X 3
component graphs to determine the height of particular centroids. Leave a
sufficient excess on these sticks to allow placement in the styrofoam base at their
respective (1X2) centroid positions. A three-dimensional model of the variable
vectors may be constructed by pushing wires through a styrofoam ball and
using Fig. 4 for proper orientation.

Specimens Examined

In the following list of specimens examined, a total of 1456, bold-faced letters
preceding the familial or subfamilial name will be used to identify the respective
group centroids in the component graphs (Figs. 5, 6, 10). Within the Phyllosto-
matidae, bold-faced numbers indicate the species identity in the component
graphs (Figs. 11-16). Any variations from this scheme will be noted in the
respective legends. Numbers following scientific names indicate sample size.
The mnemonic acronyms (for example, PTEROP for Pteropodidae and
PHYNYC for Phyllonycterinae) used in the figures are sufficiently phonetic to
provide easy interpretation.

A. Pteropodidae (172); Aethalops alecto, 5; Chironax melanocephalus, 4; Cynopterus
archipelagus, 4; C. brachyotis , 5; C. sphinx , 4; Dobsonia inermis, 2; D. minor, 2; D.
moluccensis, 1; D. praedatrix, 1; Dyacopterus spadiceus, 3; Eidolon helvum, 2; Epomophorus
labiatus, 1; E. minor, 4; E. wahlbergi, 1; Balionycteris maculata, 5; Epomops dobsoni,
1; Haplonycterisfisheri, 3; Hypsignathus monstrosus, 1; Megaerops ecaudatus, 3; M. wetmorei,
3; Micropteropus pusillus, 3; Myonycteris torquatu, 2; Nanonycteris veldkampi, 1;
Penthetor lucasi, 7; Ptenochirus jagori, 6; Pteropus alecto, 2; P. anetianus, 1; P. giganteus,
3; P. hypomelanus, 4; P. lylei, 1; P. melanotus, 1; P. rufus, 1; P. tonganus, 5; P. woodford i,
7; Rousettus amplexicaudatus, 1; R. angolensis, 1; R. arabicus, 2; R. leschenaulti, 3; R.
obliviosus, 1; Scotonycteris zenkeri, 1; Eonycteris spelaea, 3; Macroglossus lagochilus, 4;
M. minimus, 3; Megaloglossus woermanni, 8; Melonycteris melanops, 2; M. woodfordi,
4; Notonycteris macdonaldi, 11; Syconycteris crassa, 4; Nyctimene albiventer, 6; N.
cephalotes, 3; N. major, 6; N. robinsoni, 6; Paranyctimene raptor, 3; Harpionycteris
white head i, 1.

B. Rhinopomatidae (9): Rhinopoma hardwickei, 5; R. microphyllum, 1; R. muscatellum,
3.

C. Craseonycteridae (5): Craseonycteris thonglongyai, 5.

D. Emballonuridae (90): Centronycteris maximilliani, 1; Coleura afra, 5; Cormura
brevirostris, 2; Emballonura atrata, 1; E. beccarii, 6; E. monticola, 3; E. nigrescens, 3;
E. raffrayana, 2; E. semicaudata, 5; Peropteryx kappleri, 1; P. macrotis, 2; P. leucopterus, 2;
Rhynchonycteris naso, 2; Saccopteryx bilineata, 3; Taphozous australis, 2; T. flaviventris,
3; T. hamiltoni, 2; T. hildegardeae, 2; T. longimanus, 2; T. mauritianus, 3; 7. melanopogon,
8; T. nudiventris, 4; T. peli, 3; T. perforatus, 9; T. pluto, 5; T. saccolaimus, 5; Cyttarops alecto,
1; Depanycteris isabella, 1; Did id urns scutatus, 1; D. albus, 1.

E. Rhinolophidae (140): Rhinolophus acuminatus, 2; R. affinis, 2; R. alcyone, 6; R.
arcuatus, 2; R. blassi, 2; R. borneensis, 1; R. creaghi, 2; R. capensis, 2; R. clivosus, 4; R.
cornutus, 2; R. deckeni, 2; R. denti, 2; R. euryale, 3; R. euryotis, 2; R. ferrumequinum, 4;
R. fumigatus, 2; R. hildebrandti, 2; R. hipposideros, 5; R. keyensis, 3; R. landeri, 3; R. lepidus,
2; R. I act us, 2; R. macrotis, 1; R. madurensis, 1; R. malayanus, 1; R. megaphyllus, 2; R.

BIOLOGY OF THE PHYLLOSTOMATIDAE

237

mehelyi, 1; R. pearsoni, 2; R. philippinensis, 2; R. pusillus, 2; refulgens, 4; R. rouxi, 2;
R. shameli, 1; R. simulator , 1; R. stheno, 2; R. subbad ins, 2; R. swinnyi , 2; Hipposideros
armiger , 3; H. bicolor, 4; H. caffer, 8; H. camerunensis, 2; H. cineraceous, 2; H. commersoni,
4; H. cyclops, 4; H. diadema, 2; H. galeritus, 1; H. lankadiva, 3; H. larvatus, 3; H. lylei, 2;
H. pratti, 1; H. speoris, 2; Aselliscus tricuspidatus, 2; Asellia tridens, 3; Cloeotis percivali,
5; Coelops frithii, 2; Triaenops persicus, 2.

F. Nycteridae (26): Nycteris urge, 2; N. grandis, 2; N. hispida, 4; N. javanica, 3; N.
macrotis, 6; N. thebaica, 5; N. tragata, 1; N. woodi, 3.

G. Megadermatidae (14): Cardioderma cor, 4; Lavia frons, 2; Macroderma gigas, 1;
Megaderma lyra, 5; M. spasma, 2.

PL Noctilionidae (6): Noctilio albiventris, 4; N. leporinus, 2.

I. Mormoopidae (8): Pteronotus parnellii, 2; P. davyi, 2; P. gymnonotus, 2; Mormoops
blainvillii, 1; M. megalophylla, 1.

J. Phyllostomatinae (183): 1-2 Micronycteris megalot is, 14; 3 M. schmidtorum, 8; 4
M. minuta, 8; 5 M. hirsuta, 4; 6-7 M. brachyotis, 6; 8 M. pusilla, 1; 9 M. nicefori, 8; 10 M.
sylvestris, 4; 11 M. behni, 2; 12 M. daviesi, 5; 13 Macrotus waterhousii, 4; 14 M. cali-
fornicus, 10; 15-16 Lonchorhina aurita, 13; 17 L. orinocensis, 1; 18-19 Macrophyllum
macrophyllum, 8; 20-21 Tonatia bidens, 7; 22 T. brasiliensis, 3; 23 T. carrikeri, 3; 24 T.
nicaraguae, 5; 25 T. silvicola, 8; 26 T. venezuelae, 3; 27 Mimon bennetti, 1; 28 M. cozumelae,
4; 29-30 M. crenulatum, 12; 31 M. koepckeae, 1; 32-33 Phyllostomus discolor, 10; 34 P.
hastatus, 6; 35 P. elongatus, 4; 36 P. latifolius, 2; 37 Phylloderma stenops, 2; 38 Trachops
cirrhosus, 10; 39 Chrotopterus auritus, 3; 40 Vampyrum spectrum, 5.

K. Glossophaginae (156): 1 Glossophaga soricina, 6; 2 G. alticola, 5; 3 G. commissarisi,
10; 4 G. longirostris, 5; 5 Monophyllus redmani, 6; 6 M. plethodon, 5; 7 Leptonycteris nivalis,
10; 8 L. sanborni, 3; 9 L. curasoae, 5; 10 Lonchophylla Hesperia, 8; 11 L. mordax, 10; 12 L.
concava, 2; 13 L. robusta, 7; 14 L. thomasi, 10; 15 Lionycteris spurrelli, 4; 16
Anoura geoffroyi, 5; 17 A. caudifera, 3; 18 A. cultrata, 5; 19 A. werckleae, 2; 20 A.
brevirostrum, 2; 21 Sderonycteris ega, 1; 22 Lichonycteris degener, 1; 23-24 L. obscura,
7; 25 Hylonycteris underwoodi, 5; 26 Platalina genovensium, 6; 27 Choeroniscus godmani, 3;
28 C. minor, 3; 29 C. intermedins, 6; 30 C. inca, 3; 31 C. periosus, 1; 32 Choeronycteris
mexicana, 10; 33 Musonycteris harrisoni, 3.

L. Carolliinae (23): 41 Carollia castanea, 6; 42 C. subrufa, 2; 43 c. brevicauda, 4; 44 C.
perspicillata, 4; 45 RHinophylla pumilio, 2; 46 R. alethina, 2; 47 R. fischerae, 3.

M. Stenoderminae (276): 1 Sturnira lilium, 5; 2 S. thomasi, 3; 3 S. tildae, 5; 4 S. magna,
6; 5 S. mordax, 1; 6 S. bidens, 6; 7 S. nana, 5; 8 S. aratathomasi, 3; 9 S. ludovici, 10; 10 5.
erythromos, 6; 11 Uroderma bilobatum, 10; 12 U. magnirostrum, 2; 13 Vampyrops infuscus,
5; 14 V. vittatus, 4; 15 V. dorsalis, 6; 16 V. aurarius, 6; 17 V. nigellus, 2; 18 V. brachycephalus,
1; 19 V. helleri, 6; 20 V. lineatus, 5; 21 V. recifinus, 2; 22 Vampyrops sp. (new species, fide
Gardner and Handley), 5; 23 Vampyrodes caraccioli, 5; 24 Vampyressa pusilla, 4; 25 V.
melissa, 6; 26 V. nymphaea, 3; 27 V. brocki, 1; 28 V. bidens, 3; 29 Chiroderma doriae, 2;
30 C. villosum, 6; 31 C. salvini, 4; 32 C. trinatatum, 6; 33 C. improvisum, 1; 34 Ectophylla
macconnelli, 6; 35 Artibeus cinereus, 6; 36 A. glaucus, 2; 37 A. watsoni, 4; 38 A. pliaeotis, 6;
39 A. toltecus, 5; 40 A. aztecus, 5; 41 A. hirsutus, 6; 42 A. inopinatus, 5; 43 A. concolor, 5;
44 A. jamaicensis, 8; 45 A. planirostris, 10; 46 A. lituratus, 8; 47 Artibeus sp. (undescribed
species, fide D. R. Patten), 10; 48 Enchisthenes harti, 6; 49 Ardops nichollsi, 6; 50 Phyllops
falcatus, 1; 51 P. haitiensis, 4; 52 Ariteus flavescens, 6; 53 Stenoderma rufum, 6; 54
Pygoderma bilabiatum, 1; 55 Ametrida centurio, 8; 56 Sphaeronycteris toxophyllum, 2;
57 Centurio senex, 5.

N. Phyllonycterinae (27): 58 BrachyphyUa cavernarum, 6; 59 B. nana, 3; 60 Erophylla
bombifrons, 3; 61 E. sezekorni, 5; 62 Phyllonycteris poeyi, 4; 63 P. aphylla, 6.

O. Desmodontinae (13): 48-49 Desmodus rotundas, 5; 50 Diaemus youngi, 5; 51 Dipliylla
ecaudata, 3.

P. Natalidae (4): Natalus stramineus, 3; N. micropus, 1.

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SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

Q. Thyropteridae (3): Thyroptera discifera, 2; T. tricolor, 1.

Furipteridae (1): Furipterus horrens, 1; sample too small for analysis.

R Myzapodidae (2): Myzapoda aurita, 2.

S. Vespertilionidae (178): Barbastella barbastellus, 5; Chcdinolobus gouldi, 1; C. tuber-
culatus, 2; C. variegatus, 2; Eptesicus bottae, 6; E. hottentotus, 1; E. serotinus, 4; E.
sonialiscus, 1; E. tenuipinnis, 3; Euderma maculatum, 2; Endiscopus denticulus, 1;
Hesperoptenus tickelli, 2; Histiotus montanus, 1; Laephotis botswanae, 2; Lasionycteris
noctivagans, 1; Lasiurus borealis, 3; L. cinereus, 2; L. egregius, 1; L. intermedins, 2; L.
seminolus, 2; Minetillus moloneyi, 3; Myotis adversus, 2; M. austroriparius, 2; M. bech-
steini, 3; M. blythi, 3; M. brandti, 1; M. capaccinii, 3; M. daubentonii, 3; M. evotis, 1;
M. formosus, 2; M. muricola, 3; M. myotis, 3; M. mystacinus, 4; M. nattereri, 2; M. emar-
ginatus, 3; M. ricketti, 1; M. scotti, 4; M. welwitschii, 2; Nycticeius humeralis, 2; N. schlief-
feni, 1; Nycatalus aviator, 3; N. azoreum, 1; N. lasiopterus, 4; N. leisleri, 2; N. noctula, 5;
Otonycteris hemprichi, 5; Scotoecus hirundo, 2; Pliiletor brachypterus, 4; Pipistrellus imbri-
catus, 4; P. kuhlii, 1; P. nanulus, I; P. pipistrellus, 5; P. savii, 1; P. subflavus, 3; Plecotus
auritus, 5; P. phyllotis, 2; P. townsendii, 2; Scotomanes ornatus, 1; Scotophilus gigas, 2; S.
heathi, 3; S. leucogaster, 2; Tylonycteris pachypus, 3; T. robustula, 10; Vespertilio superans,
1; Miniopterus medius, 2; M. schreibersi, 4; Harpiocephalus harpia, 1; Marina aurata, 3; M.
cyclotis, 2; M. huttoni, 1; M. leucogaster, 1; Kerivoula cuprosa, 1; K. hardwickei, 2; K.
picta, 1; Nyctophilus geoffroyi, 1.

T. Mystacinidae (8): Mystacina tuberculata, 8.

U. Molossidae (1 12): Cheiromeles torquatus, 3; Eomops albatus, 1; Eumops auripendulus,
3; E. bonariensis, 1; E. glaucinus, 1; E. hansae, 1; E. trumbulli, 1; E. underwoodi, 1; Molossops
brachymeles, 1; M. temmincki, 1; M. greenhalli, 1; Molossus ater, 4; M. bondae, 1; M.
crassicaudatus, 1; M. molossus, 6; Otomops martiensseni, 4; O. wroughtoni, 2; Sauromys
petrophilus, 3; Promops centralis, 1; P. davisoni, 1; P. nasutus, 6; Tadarida aegyptiaca, 2;
T. africana, 2; T. aloysiisabaudiae, 2; T. ansorgei, 3; T. aurispinosa, 2; T. australis, 2; T.
bivittata, 2; T. condylura, 3; T. congicus, 2; T. demonstrator, 2; T. doriae, 4; T. femorosacca,
1; T. gallagheri, 1; T. jobensis, 4; T. jugularis, 2; T. laticaudata, 3; T. leonis, 1; T. lobata,
2; T. macrotis, 2; T. major, 1; T. midas, 2; T. nanulus, 2; T. nigeriae, 4; T. norfolkensis, 2;
T. plicata, 1; T. pumila, 3; 7. russata, 2; T. sarasinorum, 5; T. spurrelli, 2; T. teniotus, 3;
T. thersites, 1.

Acknowledgments

We are deeply indebted to the following institutions and persons for making
available the material examined by us: American Museum of Natural History,
Karl F. Koopman; British Museum (Natural History), John E. Hill; Florida
State Museum, Stephen Humphrey; Louisiana State University, Museum of
Zoology, George H. Lowery, Jr.; Museum of Vertebrate Zoology, University of
California, Berkeley, James L. Patton; Museum of Southwestern Biology,
University of New Mexico, James S. Findley; Museum of Natural History,
University of Kansas, Robert S. Hoffmann; The Museum, Texas Tech University,
Hugh H. Genoways; Natural History Museum of Los Angeles County, Lan Lester
and Donald Patten; Naturhistorisch Museum, Wien, Kurt Bauer; Natur-Museum
Senckenberg, Frankfurt, Heinz Felten and Dieter Kock; Royal Ontario Museum,
R. L. Peterson and Judith L. Eger; United States National Museum, including the
Biological Surveys Collection, Alfred L. Gardner, Don E. Wilson, and C. O.
Handley, Jr.

We also wish to thank Russell Benson, Department of Mathematics, California
State University, Fullerton (CSUF), for his assistance in developing the calculation

BIOLOGY OF THE PHYLLOSTOMATIDAE

239

of geometric variables. Steven Eich, James Lamprecht, Monte D’Asta, and Mark
Hartman, Computer Center, CSUF, provided valuable aid and advice in Fortran
programming and computer processing. We are especially grateful to Richard A.
Pimentel, California Polytechnic University, San Luis Obispo, who unselfishly
assisted us with the multivariate analyses, which included using an unpublished
program (DISANAL) that he developed. He also reviewed the manuscript and
provided assistance in its preparation.

Finally, we wish to express our gratitude to Susan E. Smith who sat for many
hours recording measurements, keypunched data, helped with the illustrations,
and most of all provided moral support and companionship to the senior author.

Results and Discussion

The mean (range in parentheses), one standard error, and coefficient of
variation for the raw variables and selected derived variables are given in the
Appendix (Tables A1-A21). A pooled correlation matrix for raw variables was
computed, and all coefficients, except those for the third phalanx of digit III,
were strongly and positively correlated (P< 0.001). This was to be expected owing
to the size/growth nature of these variables. The coefficients for the third phalanx
of digit III were low because this phalanx is not present in all groups of bats
(for example, pteropodids, emballonuroids, rhinolophoids, see Miller, 1907).
The largest coefficients of correlation for this phalanx were shown with the
metacarpal and two phalanges of digit V, 0.405 (P<0.05) and 0.325 (P<0.05),
respectively.

Principal components analysis. — The results of the principal components
analysis are given in Figs. 3 and 4 and Tables 2 and 3. Because of the notorious
susceptibility of the first component axis to size factors, this analysis yields only
broad generalizations concerning the shape of bat wings. The first component,
usually designated the “size component,” exhibits 91.8 per cent of the total
variation (Table 2). Also, the component correlations for all variables are high
for this component. The first three components account for 96.7 per cent of the
total variation. Although component loading extends to the twelfth component,
99.1 per cent is accumulated by the sixth. The majority of the loading, past the
first three components, is contributed by the third phalanx of digit III, which
exhibits high loading in the fourth and seventh component (51.36 and 13.75 per
cent, respectively).

As noted above, the first component contains high loading as the result of
general size. This is illustrated by the complete agreement of signs by all
coefficients in this component (Table 2). The direction of the sign (negative, in
this case) is irrelevant and simply indicates that all variables increase ( + ) or
decrease ( — ) in the same direction (for example, length of the head and body
decreases in consort with length of the forearm or any of the other raw variables).
The fact that the component scores for each variable are of different magnitude
indicates general positive allometry among the variables. The effect of size in
the first component also can be seen in Figs. 3 and 4. In figure 4A-B, the agree-

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Table 2. — Eigenvectors (direction cosines) of principal components for lengths of head and
body and selected wing elements. Only the first three components are shown because most
of the variation is exhibited in these components. The numbers in parentheses following each
component score indicates the percentage of variance contributed by each variable to a

particular component.

Variable

Component Axes

Cumulative
per cent

Head and body

-0.626(96.07)

0.701 ( 3.53)

-0.075 ( 0.03)

99.63

Forearm

-0.427 (96.77)

-0.173 ( 0.46)

0.176 ( 0.38)

97.61

Metacarpal III

-0.313 (92.52)

-0.124 ( 0.42)

0.485 ( 5.13)

98.07

Digit III, phalanx 1

-0.186 (89.16)

-0.038 ( 0.11)

— 0.130 ( 1.00)

90.27

Digit III, phalanx 2

-0.244 (84.53)

— 0.165 ( 1.13)

-0.525 ( 9.03)

94.32

Digit III, phalanx 3

-0.022 ( 6.03)

-0.247 (21.63)

0.014 ( 0.05)

27.71

Metacarpal IV

-0.303 (93.16)

-0.261 ( 2.02)

0.362 ( 3.08)

98.26

Digit IV, phalanx 1

-0.150(86.65)

-0.069 ( 0.53)

-0.139 ( 1.70)

88.88

Digit IV, phalanx 2

-0.133 (70.89)

-0.150 ( 2.62)

-0.440 (17.89)

91.40

Metacarpal V

-0.276 (86.51)

-0.505 ( 8.51)

-0.064 ( 0.11)

95.13

Digit V, phalanx 1

-0.116(82.78)

-0.106 ( 2.04)

-0.094 ( 1.26)

86.08

Digit V, phalanx 2

-0.107 (75.09)

-0.1 14 ( 2.50)

-0.280 (11.90)

89.49

Per cent trace

91.8

2.7

2.1

Cumulative per cent

91.8

94.5

96.7

ment among the signs of the first component scores is manifested by all vectors of
variables (direction cosines) orienting toward the left. Likewise, the ordination of
group centroids along the first component axis (Fig. 3) aligns large-sized bats
(Pteropodidae, A) to the left, and small-sized bats (Craseonycteridae, C) to the
right. Also, it should be noted that the nature of the ordination of groups (Fig. 3)
is greatly influenced, especially in the first two component axes, by the magnitude
of the eigenvalues for head and body length ( — 0.626 and 0.701, Table 2 and
Fig. 4A-C). Other vectors of variables that markedly affect the ordination along
the first component are the lengths of the forearm (B) and the metacarpals of
digits III-V (C, G, J) ( — 0.427, —0.313, and —0.276, respectively).

In the second component, all coefficients, except that for the length of the head
and body, agree in sign (Table 2). This strongly suggests that the size and shape
of bat wings are essentially independent of body size and, presumably, weight.
The fact that all of the coefficients for intrinsic wing elements vary in magnitude
continues to indicate a level of positive allometry. Other than head and body length,
the strongest eigenvalue in this component axis is that for the fifth metacarpal
( — 0.505). It is difficult to evaluate the shape tendencies in the second component
because the correlation structure is rather weak in both this and the third com¬
ponent. In addition, a minor portion of the variation is shown in these two com¬
ponents compared to the overwhelming nature of the first. A cautious in¬
terpretation of the shape trends in the second component might be that shape is
modified by a factor of size.

Influence attributable to shape are much more distinct, albeit weak, among
the coefficients of the third component. Body size, as expressed by the length of

BIOLOGY OF THE PHYLLOSTOMATIDAE

241

Table 3. — Mean coordinates of group centroids from the principal components analysis. These

centroids are plotted in Fig. 3.

Taxon

Code

Component axes

Pteropodidae

-68.73

9.16

-21.46

Rhinopomatidae

16.06

1.95

5.81

Craseonycteridae

60.81

-1.68

-9.15

Emballonuridae

2.23

1.05

10.75

Rhinolophidae

9.84

-2.62

-1.56

Nycteridae

13.07

-4.78

-5.53

Megadermatidae

-17.38

-6.65

-11.30

Noctilionidae

-24.10

-17.87

2.90

Mormoopidae

8.07

-4.62

10.09

Phyllostomatinae

-0.64

-7.09

0.27

Glossophaginae

19.63

2.86

-1.17

Carolliinae

24.47

-4.63

-4.36

Stenoderminae

4.14

-6.08

-1.59

Phyllonycterinae

-2.26

-0.75

3.41

Desmodontinae

-15.05

-5.44

7.14

Natal idae

32.07

-9.13

0.37

Thyropteridae

40.68

-4.96

6.70

Myzapodidae

13.70

-9.45

1.06

Vespertilionidae

19.24

-2.43

6.15

Mystacinidae

18.16

-0.76

6.14

Molossidae

2.35

9.66

9.51

head and body, has little influence in this component, having expended most of
its force in the ordination of the first and second component axes. It will be noted
(Table 2) that several of the wing elements, notably the third and fourth meta-
carpals (C,G) and the second phalanges of digits III-V (E,I,L), have their largest
eigenvalues in the third component. The divergence of variable vectors, caused by
differential signs in the third component axis, further substantiates the shape trends
of this component (Fig. 4B-C). Bearing in mind that only a small portion of the
variation is expressed and the weak correlation structure of the third component,
we cautiously direct attention to several interesting associations among the
variables in this component.

In Figure 4B-C, the vectors for variables of all intrinsic wing elements (B-L)
are directed to the left; the vector for head and body length (A) projects to the
right in the 2X3 graph (Fig. 4C) again indicating the independent nature of this
variable. As noted previously, the general similarity in the direction of orientation
of all vectors for wing elements postulates a general allometric relationship among
wing components in terms of size. However, in the two graphs (1X3 and 2 X 3),
the vectors for wing components diverge into different regions of the graphs (that
is, some orient upward and others are directed downward). This signifies dif¬
ferences in relative independence that ultimately are expressed as shape.

The vectors for the third and fourth metacarpals (C, G) project in the same
general direction and are nearly equal in length (Fig. 4B-C), indicating that their

242

SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

Fig. 3. — Component graph from principal component analysis. Group centroids are
plotted on the 1 X2 axes and 1 X3 axes to illustrate their position in Euclidean three-space.
Coordinates for these centroids are given in Table 3. Stars represent phyllostomatid centroids
(see list of specimens examined or Table 3 for key to alphabetic code). This figure may be
xeroxed and folded on the dotted line to help visualize the three-dimensionality of the dis¬
persion of centroids.

variation is associated. Although somewhat removed, the vector for the forearm
(B) tends to share this same general relationship. It is interesting to note that the
vector for the fifth metacarpal (J) is rather far removed from the third and fourth
metacarpals thereby suggesting a marked divergence in its pattern of variation.
This suggests that the forearm and metacarpals of digits III and IV vary as a unit,
whereas the metacarpal of the fifth digit is somewhat independent. Following
these examples, we can point to several additional interesting sets of vectors that

Fig. 4. — Positive eigenvectors (A-C) and variable vectors (D-F) for the raw variables
computed in the principal components analysis and discriminant analysis, respectively.
Coordinates for these vectors are given in Tables 2 and 4, respectively. Corresponding sets
of vectors from these two analyses are shown side-by-side to allow easy comparison. The
negative portions of the vectors were omitted to avoid confusing the diagram. If shown, they
would project an equal distance in the opposite direction past the zero-zero point. Letters at
the ends of vectors refer to the respective lengths of variables: A, head and body; B, forearm;
C, metacarpal III; D, first phalanx III; E, second phalanx III; F, third phalanx III; G,

BIOLOGY OF THE PHYLLOSTOMATIDAE

243

metacarpal IV; H, first phalanx IV; I, second phalanx IV; J, metacarpal V; K, first phalanx
V; L, second phalanx V. See text for discussion.

244

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have generally associated patterns of variation. The first phalanges of digits III
and IV (D, H) appear to have a similarly related effect on wing shape. Likewise,
the vectors for the second phalanges of digits III to V (E, I, L) suggest a similar
relationship among these phalangeal elements. These two sets of variables, to¬
gether, diverge markedly from the metacarpal elements (C, G, J) of their respective
digits. The vector for the first phalanx of digit V (K) tends to associate with the
fifth metacarpal (J). These patterns of positive allometry generally indicate the
complexities of wing shape.

As we have seen, size greatly influences the ordination of groups in the principal
components analysis. This is exerted strongly in the first component and hardly
at all in subsequent components. The overwhelming effect of size has led many
investigators to attempt to eliminate size as an ordinating factor and thereby in¬
crease the component loading by the “inherent” shaping qualities of their raw
variables. The product of these efforts has been the mathematical adulteration
of raw variables into ratios, indices, and proportions, which may appear to
eliminate size, but which actually obscure or otherwise confound the recognition
of independent patterns of variation. Simply ignoring the first component and
considering components 2-4 is not a satisfactory means of eliminating size, be¬
cause the component correlations are usually even weaker in the fourth compo¬
nent. We submit that in a morphometric analysis such as this, and in fact in all
analyses based on absolute measures of continuous variables, size reflects the
essence of variation. By this, we do not mean absolute size in itself, but the allo-
metric and isometric aspects of size that ultimately are expressed as synergistic
relationships among variables. Therefore, any attempt to strip away the effects of
size seriously risks masking or totally eliminating the interactive relationships
between size and shape.

The centroids computed for each group in the principal components analysis
are given in Table 3 and plotted in Fig. 3. The cigar-shaped dispersion, as noted
earlier, is oriented with the longest axis more or less corresponding to the first
component axis. The shape of this cluster is caused mostly by the effects of gross
size. Most taxa, including the six subfamilies of phyllostomatids (J to O), are
packed in the midregion of the dispersion. By examining the vectors of variables
shown in Fig. 4A-C and the group centroids plotted in Fig. 3, the reader can
begin to appreciate the ordinating effects exerted by the various characters. In
the lower diagram of Fig. 3 (axes 1 X 2), the pteropodids (A) are pushed to the far
left and into the upper quadrant, primarily on the basis of large head and body
length. The noctilionids (H), megadermatids (G), and, to a lesser extent, the
desmodontines (O) also are influenced by the positive force of this vector. The
craseonycterids (C), on the other hand, ordinate into the lower right-hand quadrant
by the opposite (negative) effect of the vector for head and body length. The taxa
in the lower left-hand quadrant are ordinated by the positive (large size) effects of
all vectors of variables for wing elements; especially lengths of the forearm, second
phalanx of digit III, third and fourth metacarpals, and second phalanx of digit
V. The taxa in the upper right-hand quadrant ordinate by the negative (small
size) effects of these wing elements. Note that the phyllostomatines (Fig. 3J)

BIOLOGY OF THE PHYLLOSTOMATIDAE

245

are pushed, almost directly, by the vector for the fifth metacarpal (Fig. 4J), where¬
as the molossids (U) and emballonurids (D) lie along the tail end of this vector.
The majority of the taxa are ordinated into the lower right-hand quadrant, which
results from a complex synergistic interaction among the intrinsic elements of
the wing.

The effects of the vector for variables in the third component may be seen in the
upper diagram of Fig. 3 (axes 1 X 3). In this component graph, the pteropodids
(A) and megadermatids (G) are ordinated into the lower left-hand quadrant by
large-sized, distal phalangeal elements (E, I, L). In these two groups, the meta-
carpals constitute a relatively smaller portion of the total length of the various
digits (Fig. 7; Tables A12, A15, A16). On the other hand, noctilionids (H),
desmodontines (O), and, to a lesser extent, carolliines (N) are characterized by a
generally long forearm (B), third and fourth metacarpal (C, G), and third phalanx
of digit III (F). The taxa positioned in the upper right-hand quadrant generally
reflect a complex synergism among variables.

In summary to this point, principal components analysis is an effective screening
procedure that allows some general insights into the interactive relationships of
size and shape exhibited by the wings of bats. However, this procedure, because
of its sensitivity to gross size, is not well suited to the detection of subtle nuances
in the variation of wing shape among chiropterans. It provides a generalized view
of the tip of the iceberg, so to speak, but does not give a clear perspective of the
underlying complexity of shape. With regard to the phyllostomatids as a group,
little can be said other than that they tend to ordinate amongst the medium to
large-sized bats near the grand centroid.

Discriminant analysis. — The transformation from Euclidean space into
discriminant space effectively reduces the overwhelming influence of general
size on the ordination of group centroids without otherwise adulterating the
intrinsic variation of the raw variables. In Table 4, there is a more equitable
dispersal of the variation across the first six canonical axes. There is much more
symmetry shown by the canonical vectors of variables in Fig. 4D-F than by vectors
from the component analysis (Fig. 4A-C). In addition, the correlations of
canonical vectors and variables are more evenly dispersed across the various
canonical axes rather than being heavily focused in the first axis as was the case
in the principal components analysis.

It should be pointed out that, although the variable vector for the third phalanx
of digit III (F) is not particularly strong as compared to other vectors, its influence
on the dispersion in the first canonical axis essentially segregates taxa into two
groups — those that possess this element and those that do not. The correlation
coefficient for this variable with the first canonical axis is comparatively high
(0.540). This is equalled by the correlation coefficients for the fifth metacapal
(J) and second phalanx of digit V (L), which have their greatest affinity with the
third canonical axis (0.489 and 0.582, respectively). A more detailed discussion
of the effects of these various variable vectors on the size and shape of chiropteran
wings will be presented in the following accounts.

246

SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

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BIOLOGY OF THE PHYLLOSTOMATIDAE

247

Table 5. — Group centroids for the first six canonical axes. The first three axes are plotted

in Figs. 5 and 6.

Canonical axes

Taxon

Code

Pteropodidae

-3.738

-2.621

2.456

0.992

-0.101

-0.294

Rhinopomatidae

-2.791

3.379

-2.446

4.291

0.374

-3.076

Craseonycteridae

-1.660

0.525

-0.725

-2.080

-1.265

-0.199

Emballonuridae

-0.733

0.376

-2.779

2.421

-1.728

1.512

Rhinolophidae

-3.741

1.106

-1.076

-1.379

0.010

-0.547

Nycteridae

-3.711

-0.434

-0.739

-3.183

0.863

2.374

Megadermatidae

-4.678

1.284

-0.556

-4.321

-2.089

1.244

Noctilionidae

3.106

1.231

-3.222

-2.933

-3.010

-3.535

Mormoopidae

2.203

1.986

-0.166

1.015

-2.226

-1.714

Phyllostomatinae

1.364

1.412

1.114

-0.510

-0.535

-0.170

Glossophaginae

1.778

0.770

1.622

-0.059

-0.792

0.518

Carolliinae

1.590

0.298

1.608

-0.233

-0.538

1.227

Stenoderminae

2.752

1.360

2.018

-0.124

-0.294

0.345

Phyllonycterinae

1.028

1.885

1.073

0.891

0.758

-0.912

Desmodontinae

2.207

3.805

1.199

-0.187

-0.700

-1.650

Natal idae

-2.076

1.060

-1.075

-0.762

0.158

2.415

Thyropteridae

-0.024

0.323

-0.695

-0.636

2.004

1.694

Myzapodidae

1.146

0.288

0.051

-0.416

1.367

-1.053

Vespertilionidae

0.781

0.480

-0.843

0.423

1.813

0.291

Mystacinidae

2.315

2.132

1.447

0.774

1.658

-1.224

Molossidae

3.194

-3.510

-1.523

-0.585

-0.120

-0.474

Pteropodidae

Fruit bats are generally the largest chiropterans in terms of absolute size of all
raw variables. We have observed that overall large size greatly effects the
ordination in the principal components analysis. However, in discriminant space,
these overwhelming effects of size are much reduced. Because of their large size,
the pteropodids are especially well suited to illustrate the moderation of size in
the discriminant analysis. The strongest vector in the principal component analysis
was that for head and body length (A) — see Fig. 4A-C. This feature in the dis¬
criminant analysis is one of the least powerful (Fig. 4D-F; Tables 4 and 6). Not
only is the length of the vector short as compared to others such as those for the
lengths of the forearm (B), and third and fifth metacarpal (C, J), for example,
but it is directed away (approximately 90 degrees) from the group centroid for
pteropodids. The canonical coefficient (0.225) for head and body length in the
first canonical axis (Table 4) is positive and near zero, suggesting the denial of
large body size by pteropodids relative to this axis. Although comparatively mi¬
nor, the greatest influence by this variable on the ordination of bats in discrimi¬
nant space occurs in the second and third axes, but here too the vector generally
orients away from the pteropodids. Furthermore, the contribution of this variable
to the overall discriminant functions of the various centroids appears to be minor
(Table 6).

248

SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

Fig. 5. — Canonical graph from discriminant analysis. Group centroids are plotted with
their respective confidence circles (95 per cent) on 1X3 canonical axes of discriminant
space and the coordinates for these are given in Table 5. Stars and stippled area indicate
phyllostomatid centroids and confidence circles. The confidence circle for the Myzapodi-
dae (R) is too large to plot. See list of specimens examined or Table 5 for key to alphabetic
code and text for discussion.

The length of the forearm (B) is one of the more powerful forces in the overall
ordination of groups (Tables 4, and 6). In the first two canonical axes, the vector
for this variable lies approximately perpendicular to the pteropodid centroid.
Relative to the molossid centroid (U), this vector may be interpreted as exerting
a positive force on the pteropodid centroid. However, the peripheral position of
this centroid to the vector suggests a weak influence by this variable. In the third

BIOLOGY OF THE PHYLLOSTOMATIDAE

249

Fig. 6. — Canonical graph from discriminant analysis. Group centroids are plotted with
their respective confidence circles (95 per cent) on 1X2 canonical axes of discriminant
space and the coordinates for these are given in Table 5. Stars and stippled area indicate
phyllostomatid centroids and confidence circles. The confidence circle for the Myzapodidae
(R) is too large to plot. See list of specimens examined or Table 5 for key to alphabetic
code and text for discussion.

canonical axis (Figs. 4E, 5), the effect of this variable is somewhat more direct
(negative). A heuristic interpretation of this variable vector would suggest a
medium to short forearm for the pteropodids.

An examination of the relative lengths of the wing and forearm (Tables A7,
A8) clarify and substantiate this interpretation. Although the absolute lengths of
all wing elements are large, the wings of pteropodids average proportionately

250

SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

shorter than those of other chiropterans (1.80). The mean relative length of the
forearm is second smallest for the order (0.65); only that of molossids is smaller
(0.63). In terms of the composition of wing span, the forearm of pteropodids
contributes an average of 35.98 per cent (range, 32.69-38.84) to the length of the
wing.

The canonical coefficients for the metacarpals of digits III-V (C, G, J)
illustrate the simultaneous nature of the interactive relationships among these
variables. By comparing Fig. 4D with Fig. 6, it will be noted that the negative
end (smallness) of the variable vector for the third metacarpal (C) passes in
proximity to the centroid of pteropodids. This indicates a rather strong tendency
in the direction of small size, especially in the first and second canonical axes.
The variable vectors for the fourth and fifth metacarpals (G, J) orient in nearly
opposite directions from each other, and both orient almost perpendicularly to the
centroid for pteropodids. It will be noted that the latter vector (J) is the stronger
of the two (Table 4) and it is oriented directly toward the rhinopomatids. The
vector for the fourth metacarpal (G) is of a lesser magnitude and is oriented
generally toward the centroid of the Molossidae. The influences of these two
variable vectors on these two centroids will be discussed beyond and are
mentioned here only for orientation by the reader. It is difficult to assess the nature
of the effect these two vectors have on the pteropodid centroid in the first and
second canonical axes; suffice it to say that it is synergistic. In the third canonical
axis (Figs. 4E, 5), the interaction of these three vectors is somewhat clearer. The
vector for the third metacarpal (C) continues in its implication of small size. The
vectors for the fourth and fifth metacarpals (C, J) maintain their opposite orien¬
tation, but their negative (smallness) ends are closer to the centroid of the
Pteropodidae than before. The net effect of all three of these vectors is to carry
the centroid in an upward direction in three-dimensional space and, because it is
the tail end of these vectors that effects this lifting, the implication is small size
for all three metacarpals. An examination of Tables A12, A16, and A19 reveals
that these manal elements of the pteropodid wing contribute the smallest per¬
centage to the overall lengths of digits III to V as compared to other chiropteran
taxa. Norberg (1972) also noted the general shortness of the metacarpals of the
megachiropterans.

A long first phalanx of digits III and V (D, K) is strongly implicated in the
discrimination of pteropodidis in all three canonical axes. The vector for this
phalanx in the fourth digit (H) is most influential in the third canonical axis (Figs.
4E-F, 5,6; Table 4) and here also suggests relatively long length. The vectors for
the second phalanx of digits IV and V (I, L) share a similar orientation as
described above for the fourth and fifth metacarpals (G, J) except that the positive
ends of these variable vectors, rather than the negative ends, carry the centroid
aloft. An examination of the percentages contributed to the discrimination of each
group (Table 6) generally substantiates the characteristically long phalangeal
elements of pteropodids. In addition, Tables A 13, A 17, A 18, A20, and A21 show
the mean percentages contributed by these phalanges to the overall lengths of
digits III to V, respectively, and further support the above interpretations by

BIOLOGY OF THE PHYLLOSTOMATIDAE

251

ranking the pteropodids as the largest, or nearly so, with respect to these wing
elements.

The vector for the second phalanx of digit III (E) presents an interesting paradox
in that it nearly parallels, in both direction and sign, the orientation of the variable
vector for the third metacarpal (C). This seems to suggest short length of this
feature in the first two canonical axes. However, there is a slight elevating quality
by the point of this vector on the centroid in the third dimension of discriminant
space. Those familiar with pteropodid wings should be duly impressed by the
extraordinary length and massive structural nature of this phalanx. However,
though this wing element is outwardly large-sized in appearance, pteropodids
rank fourth largest in terms of the average percentage contributed by this element
to the overall length of digit III, as compared to the Craseonycteridae, Megader-
matidae, and Furipteridae (Table A 14). (The latter group was not included in the
multivariate analyses because the sample size was too small.)

Therefore, pteropodids are characterized by having a relatively short wing as
the combined result of a relatively short forearm and third metacarpal (Fig. 7).
Although the two phalangeal elements of the third digit are long, the shortness
of the metacarpal tends to suppress the overall length of the digit. The total length
of digit III contributes between 61 and 67 per cent to the wingspan, shown by a
mean tip index of 1.78 (2.06-1.57), which ranks in the middle to upper range of
all bats (Table A2). The wings of pteropodids are further characterized by their
generally broad aspect (Fig. 10; Tables A3-A5). Although the shortness of the
fourth and fifth metacarpals would tend to cause a narrow wing, apparently the
combined lengthening of the phalangeal elements of digits III to V maintains the
proportional breadth.

Contrary to Findley et al. (1972), such a wing should have an excellent lift
potential at slow speeds. In addition, the relatively long phalanges of all three
digits, especially those of digits IV and V, should facilitate increased camberability
with relatively little digital flexion and thereby further augment lift potential at
slow speeds. Whereas the nearly equal (isometric) partitioning of the respective
digits may contribute, in a crude sort of way, to the slow-flight characteristics
of pteropodid wings, the fine adjustments necessary for maneuverability in slow
flight, such as hovering, apparently are not possible. In the following accounts
we will show that all other chiropteran families depart from the general isometric
construction of the wing as exhibited by the Pteropodidae.

Rhinopomatidae

As in the pteropodids, the mouse-tailed bats possess a morphometrically unique
and interesting wing (Fig. 7). Whereas the pteropodids are in a generally peripheral
location relative to the variable vector for head and body length (A), the rhinopo-
matids receive nearly the full negative (shortness) force of this vector in the
ordination of their centroid. This vector is discriminatory in all three canonical
axes (Figs. 4D-E, 5, 6). The variable vector for the length of the forearm (B) is
closely aligned with that for head and body length, but the ordinating effect of this
vector is more direct and positive (large-sized) rather than negative, and its

252

SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

influence in discrimination is more important (Table 6). The relative length of
the forearm approaches unity (0.94; Table A8), which may or may not reflect
an interactive relationship between these two variables. It is interesting to note
that this relationship is maintained through the sixth canonical axis (Table 4),
although the position of the centroid shifts slightly into a more peripheral
location.

A much more complex, extra-dimensional interaction exists for the combined
variation of the metacarpal elements of digits III-V (C, G, J) (Figs. 4D-E, 5,
6; Table 4). In the first three canonical axes, and similar to the vector for the
forearm (B), the vector for the fifth metacarpal (J) is a strong positive discriminator
(Table 6). This manal element comprises 68.10 per cent of the total length of
the fifth digit (Table A 19), and this percentage is exceeded only by some ves-
pertilionids and Noctilio. The tail (shortness) of the variable vector for the length
of the fourth metacarpal (G) also is directed toward the centroid for rhinopomatids
in the first two canonical axes. However, in the third canonical axis, the tail of this
vector is directed upward, and, although it seems to interact synergistically with
other vectors to carry some centroids aloft, its effect seems minimal in this regard
to the rhinopomatids. This is interesting in light of the apparent importance of
this variable in the discrimination of the group (Table 6). In Table 4, it will be
noted that the cumulative percentage of the variance contributed by the fourth
metacarpal to the first three canonical axes is low — 20.57 (1.66, 18.52, and 0.39,
respectively), whereas the percentage contributed in the fourth axis of discriminant
space is markedly increased — 51.55 (72.12 cumulative per cent). In addition,
this vector becomes a strong discriminator for shortness of the fourth metacarpal
and is again oriented more directly toward the centroid of the rhinopomatids.
This metacarpal contributes 59.77 per cent to the total length of the fourth digit
(Table A16). The length of the third metacarpal of rhinopomatids is particularly
striking (61 .70 per cent of the length of digit III) compared to that of other bats.
In Table A12, it is exceeded only by the emballonurid Depanycteris (63.21).
However, the influence of this variable on the dispersion in the first three canonical
axes is not readily apparent (Figs. 4D-E, 5, 6). An examination of Table 4 will
show that there exists an extradimensional effect similar to that described for the
fourth metacarpal. Whereas the orientation of this variable vector is oblique
to the rhinopomatid centroid in the first three canonical axes, its point (longness)
directly ordinates this group in the fourth through sixth dimensions of dis¬
criminant space.

The vectors for the lengths of the first and second phalanges of digit III (D, E)
generally indicate small size, although the interaction between these variables
results in vectors that tangentially effect the centroid for rhinopomatids. Again,
this effect becomes more direct in extradimensional space. The variable vectors
for these two phalanges of the fourth digit (H, I) directly indicate shortness in
the first three canonical axes. The effect is strongest for the distalmost member (I)
of this pair of phalanges. Table 6 indicates a rather minor role for the first and
second phalanges of digit V (K, L) in the discrimination of the rhinopomatids.
Nonetheless, the vector for the proximal member of this pair (K) is oriented

BIOLOGY OF THE PHYLLOSTOMATIDAE

253

24 9

20.2 22.0 22.0
6.6 3 6 5 3

196 10. 1 6.3

Fig. 7. — Diagrammatic representation of the wing construct based on the mean lengths of
variables for the Pteropodidae, Emballonuroidea, and Rhinolophoidea. Columns of numbers
associated with each construct are, from left to right: length of forearm, metacarpal, and
phalanges of digit III; length of metacarpal and phalanges of digit IV; and length of meta¬
carpal and phalanges of digit V. Digit IV is projected at the mean alpha angle computed for
each taxon (see Table Al).

positively toward the group centroid in the first two canonical axes (Figs. 4D, 6).
In the third canonical axis, the effect of this variable is reduced. Similarly, the
vector for the terminal phalanx of digit V (L) is oriented toward the centroid for
rhinopomatids, but in the third dimension this vector stands far above the
centroid. This further indicates the complexity of the variation and interactive
associations among variables.

The overall effect of the interplay among the wing elements of rhinopomatids
is to produce a wing with a below average overall aspect ratio of 5.57 (Figs. 7,
10; Table A3). Findley et al. (1972) noted the shortness of the wing tip and
indicated that rhinopomatids had the lowest tip index of all bats examined by
them. We computed an average tip index of 1 .09, which agrees with 1.19 reported
by these authors for Rhinopoma hardwickei. In addition, they commented on
the relatively short wings possessed by these bats. Although these indices and
ratios provide a vague impression of the rhinopomatid wing, they do not clearly
delineate the uniqueness of its shape or the causative aspects of this shape.

The shortness of digits III to V is most greatly effected by short phalangeal
elements and a relatively short fourth metacarpal; the third and fifth metacarpals
are among the longest for all bats (Tables A 16, A 19). As noted above,

254

SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

rhinopomatids possess nearly the longest forearm relative to their head and body
length (the relationship is almost 1:1). Whereas pteropodids, and to a greater
extent molossids, have much higher tip indices, the relative length of digit III
for both is only slightly higher than that shown for rhinopomatids (Tables A2,
A9). The long forearm, in combination with a relatively short fifth digit (0.92
as compared to 1.00 for molossids) produces a plagiopatagium with an extremely
high aspect ratio (2.16) (Figs. 7, 10; Table A5). This mean value is the largest
among bats and is exceeded in range only by molossids and emballonurids.

According to Harrison (1964:62), the flight of Rhinopoma hardwickei is
peculiar and distinctive, consisting of a “series of alternating flutters and glides,
with a rising and falling motion. . . .” Dr. Gamal Madkour, who is familiar
with R. microphylum of Egypt, indicated to us (personal communication) that
these bats are rather swift-flyers that forage in open country. In view of these
apparent conflicting observations, we hesitate to comment on the functionality
of the wing of rhinopomatids other than to say that it should be capable of
producing moderate speed as well as maneuverability. We see little basis for a
close functional relationship between the Rhinopomatidae and the rhinolophoids,
the wings of which are constructed differently. Finally, mouse-tailed bats share
the closest resemblance with the family Emballonuridae — generalized (not
taxonomic) distance 6.94. This resemblance is founded on similarity of variable
vectors for the lengths of the forearm, first phalanx of digit IV, second phalanx
of digits IV and V, and the fifth metacarpal.

Craseonycteridae

Craseonycteris thonglongyai represents the small extreme in the size variation
among the Chiroptera. These bats, recently described as a monotypic family
(Hill, 1974), can truly be thought of as “bumble-bee bats,” as they are scarcely
larger than their hymenopteran namesake. Because of their extreme small size,
we can reemphasize the rather minor effect that general size has on ordination
in discriminant space. In the principal components analysis (Fig. 3), this family
was strongly ordinated along the first component axis by the tail (smallness) of the
variable vector for head and body length (A). As we noted, the pteropodids were
ordinated in the opposite direction and the remaining taxa disperse between
these two extremes. In terms of distance coefficients, this spread (PCA) constitutes
a taxonomic (Euclidean) distance value of 131.18. Similarly large taxonomic
distances were computed between pteropodids and other small-sized taxa such
as the Natalidae and Thyropteridae (105.1 1 and 1 14. 12, respectively). However,
in discriminant space, these general size effects are markedly moderated and the
generalized distances between pteropodids and these three small-sized taxa
(9.23, 6.43, and 6.72, respectively) are suggestive of shape rather than size
differences. Whereas the vectors for nearly all variables effect the ordination of
the pteropodids by pushing them away from the centroid of the craseonycterids,
that for head and body length contributes the least percentage to the group dis¬
crimination vector of the latter (0.36, Table 6). Therefore, again we see that in

Table 6. — Percentage contributed by each direction cosine to the discriminant function of each group. These values may be compared with the
canonical coefficients (Table 4) in an interpretation of the overall effect a variable or set of variables has on the ordination of a group(s).

BIOLOGY OF THE PHYLLOSTOMATIDAE

255

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discriminant space the quantitative effects of general size are much reduced in
lieu of the more complicated synergistic interactions among variables that reflect
the qualitative aspects of size.

The variable vector for the length of the forearm (B) appears to have a mod¬
erately strong influence on the ordination of the craseonycterids (Figs. 4D-E, 5,
6; Tables 4, 6). Although this variable is the shortest, in terms of absolute length,
among all bats, the vector suggests longness of the forearm. In Table A8, the
mean relative length of the forearm (0.82) is somewhat larger than that of all bats
(0.73), further substantiating the interpretation of this vector. The combined
effect of the lengths of digit III and the forearm is the production of a relatively
long wing for the craseonycterids (Tables A2, A7).

Hill (1974), in his detailed comparison of the structure of the wing of
Craseonycteris with those of other bats, noted a rather peculiar variation among
the metacarpal elements. The third metacarpal of Craseonycteris is relatively
short as compared to the fourth and fifth, which are somewhat longer and approxi¬
mately equal in length. The relationship of the vector for the length of the third
metacarpal (C) to the centroid of craseonycterids is similar to that discussed for
pteropodids. The contribution of this element to the length of digit III (43.44,
Table A 12) is below the average of other bats. The qualitative shortness of the
fourth metacarpal is suggested by the vector for this variable (G) in the first
and second canonical axes (Figs. 4D, 6). In the third axis (Figs. 4E, 5), the group
centroid is located somewhat to the side of this variable vector, although the
implication of shortness persists. The relationship of the variable vector for
the fifth metacarpal (J), in all three axes, implies longness. The contribution of
the fourth and fifth metacarpal elements to the lengths of their respective digits
is above average for all bats (Tables A 16, A 19).

Perhaps the most striking feature of the third digit is the relatively long second
phalanx (Fig. 7). This phalanx is nearly equal to the metacarpal in length (Tables
A 12, A 14) and its contribution to the length of the digit is largest among all bats.
Although the percentage contributed to the discrimination vector of the group
(12.42, Table 6) is relatively high, the implication of this variable vector (E) in
the first three canonical axes (Figs. 4D-E, 5, 6) is toward shortness. However,
in the fourth and fifth canonical axes, the positive end (longness) of the vector
is strongly oriented toward the centroid of craseonycterids. Again, this em¬
phasizes the multidimensional and synergistic nature of the interaction among
variables on alar shape.

A similar relationship for the distal phalanx of digit III exists for the
rhinolophids, megadermatids, and, to a lesser extent, nycterids. The actual
structure of the tip portion of the wing in these bats is rather curious and is not
found in any other group. The middle and distal portion of the shaft of the second
phalanx of digit III is arched in such a way as to trap, and maintain taut, a small
section of the alar membrane in much the same fashion as the string of a bow.
The joint between the distal phalanx and the first phalanx of digit III is broad,
and there appears to be a great deal of mobility at this joint, judging from specimens
preserved in alcohol. Although we are not prepared to discuss the functional

BIOLOGY OF THE PHYLLOSTOMATIDAE

257

Fig. 8. — Diagrammatic representation of the wing construct based on the mean lengths of
variables for the Phyllostomatoidea. Columns of numbers associated with each construct are,
from left to right: length of forearm, metacarpal, and phalanges of digit III; length of meta¬
carpal and phalanges of digit IV; and length of metacarpal and phalanges of digit V. Digit IV
is projected at the mean alpha angle computed for each taxon (see Table A1 ).

ramifications of this anatomical configuration, we suggest that the apparent
emphasis in the ordination of these families by this feature implies not only
similarity in shape, but also functional similarity. Perhaps it is employed during
the “flick phase” of the wing beat cycle, or it simply may be a device for furling
this long wing element. Although the phylogenetic sources of shape are not our
primary goal in this paper, we would point out that this feature suggests a close
relationship among these families. The emballonurids possess a slightly different
folding device in this distal region of their wings, and the rhinopomatids, which
lack this feature, might represent the underived (primitive) condition for this
characteristic.

Of all the variables employed in this study, the length of the first phalanx of
digit IV appears to be the most distinctive of Craseonycteris (Table 6). This wing
element is extremely short and constitutes only 10.2 per cent of the total length
of the fourth digit and, in a relative sense, is the shortest observed in all bats (Table
A 17). The shortness of this wing element is emphasized in the discriminant
analysis by the variable vector (H) in the first three canonical axes (Figs. 4D-E,

258

SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

5, 6; Table 4). This vector is involved similarly, but to a slightly lesser degree,
in the ordination of the Rhinolophidae, Nycteridae, Megadermatidae, Noctilioni-
dae, and Natalidae. The second phalanx of digit IV is long and is second in size
only to that of the Noctilionidae (Table A 18). However, the interactive re¬
lationship of this variable is obscured by the synergistic complexity among all
variables.

The variable vectors for both phalangeal elements of the fifth digit (K, L)
also are difficult to interpret, although they indicate longness in the first three
canonical axes. Of the two variables, the length of the first phalanx of this digit
appears to be the most influential in the discrimination of the group (Table 6).
The precentages contributed to the length of digit V by the first and second
phalanges (15.79 and 18.64, respectively) are below the average for all bats
(Tables A20, A21).

The overall aspect ratio of the wing of Craseonycteris thonglongyai is slightly
below the mean for all bats (5.64, Table A3 and Fig. 10). The aspect of the
plagiopatagial portion is not particularly distinctive (1.48) and falls in the middle
to lower range for all bats (Table A5). In addition, the aspect ratio of the tip portion
of the wing is approximately equal to the average for all bats (Table A4 and Fig.
10). On the other hand, the third digit is 1.86 times as long as the forearm, which
is generally high compared to that of other bats (Table A2; Fig. 10).

The overall shape of the craseonycterid wing is the result of a rather unusual
combination of interactions among the various wing elements. The length of
the third digit appears to be most strongly influenced by the length of the distal
phalanx, which tends to offset the shortness of the metacarpal. In the fourth digit,
the relatively long metacarpal and distal phalanx appear to compensate for the
markedly shortened first phalanx. The fifth digit is relatively long, owing to a
generally isometric association with the metacarpal and second phalanx of
digit IV, and tends to offset the length of the third digit. These interactions thereby
contribute to the generally broad aspect of the wing tip.

Prompted by comments made by Findley et al. (1972) concerning an average
or below average aspect ratio coupled with a high tip index, Hill (1974) suggested
a hovering ability for these small bats. We agree that Craseonycteris may possess
this flight potential, but our basis for this assumption lies more with the structural
nature of the third digit, especially the long distal phalanx, rather than with the
relationship between aspect ratio and tip index.

Emballonurid.ae

From the standpoint of wing diversity, the emballonurids represent one of the
most intriguing families of bats. In terms of aspect ratios, they range from slightly
above average (6.05) for the order to extremely high aspect ratios (7.93). Their
forearms may be relatively short to long and, as a consequence, the tip indices
for members of the family also vary from low to high. In these general descriptive
terms, the wings of emballonurids most closely resemble those of bats of the family
Molossidae and, in some respects the Noctilionidae and Mormoopidae. However,

BIOLOGY OF THE PHYLLOSTOMATIDAE

259

this resemblance is merely superficial as these families acquire their extreme
wing shapes through different morphometric modes. To draw attention to this
misleading resemblance, we will draw comparisons between the emballonurids,
molossids, and noctilionids in this account. The group centroids of these three
families are located in separate regions of discriminant space (Figs. 5, 6).

Emballonurids are about average for bats in length of the head and body. The
vector for this variable (A) is a minor force in the overall discrimination of the
group (Table 6). Head and body length has a slightly stronger effect in the
ordination of the Molossidae; this is particularly true in the first and second
canonical axes (Figs. 4D, 6).

Length of forearm appears to be a moderately important variable in the ordi¬
nation of the emballonurid centroid. This appears to be a general feature of
those bats referred to the superfamilies Emballonuroidea and Rhinolophoidea,
which are generally characterized by possessing relatively long forearms (Table
A8). Within the Emballonuridae, the mean, relative length of the forearm ap¬
proaches unity (0.93). Although most species range below this value,
the exceptions are notable: Centronycteris maximiliani (1.14); Cyttarops
alecto (1.10); Emballonura solomonis (1.11); E. beccarii (1.06); and Cormura
brevirostris (1.04). The vector for the length of the forearm (B) contributes a
moderately low percentage (3.15) to the discrimination vector of the emballonurids
(Table 6). By comparison, the ordination of the molossids is more strongly in¬
fluenced by the tail (shortness) end of this vector. This emphasis on short
length of the forearm is reflected in the higher percentage contributed by this
vector (9.28) to the group discrimination vector of molossids (Table 6). There¬
fore, although the absolute length of the forearm in these two groups is outwardly
similar, there is a fundamental difference in their respective contribution to the
shape of the wing (Table A8).

The variation of the dactylopatagial portion depicts even more striking
differences in the wing construction of emballonurids and molossids (Figs. 7, 9).
On the whole, the length of the third digit of emballonurids is not particularly
impressive. The mean tip index (1.61) is well below the average for all bats
(Table A2). Centronycteris, Saccopteryx, and several species of Taphozous,
especially T. peli, have unusually large tip indices (1.70-1.90). On the contrary,
molossids generally are characterized by larger than average tip length (Table
A2).

The vectors for the various elements of digit III (C, D, E) are involved in the
overall complex synergism among variables and their effect is not easily inter¬
preted. In the first canonical axis (Figs. 4D-E, 5, 6), only the vector for the first
phalanx (D) exerts a positive force on the ordination of emballonurids (Tables 4,
5); shortness is emphasized by the other vectors for this digit. The converse of
these actions is implied for the ordination of the Molossidae with respect to the
vectors associated with digit III. Also, the centroid for the Noctilionidae is
closely associated with that of the Molossidae in this canonical axis.

Ordination along the second canonical axis illustrates a somewhat different
picture (Tables 4, 5). Here the vectors for the metacarpal and second phalanx

260

SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

39.2

35.9 35.4 34.2
154 9.3 9.5
20.6 9.4 10.0

34.8

33.7 33. 1 30.0
14.6 9.3 7.9
9.5 6.0 7.5
5.0

42.4

39.6 38 5 36.6
15.0 12.0 9.4
144 10.2 7.6

6.9

42 9

40.1 38.9 35.2
108 9.9 6.0

13.2 13.0 I I. I
I I. I

Fig. 9. — Diagrammatic representation of the wing construct based on the mean lengths of
variables for the Vespertilionoidea. Columns of numbers associated with each construct are,
from left to right: length of forearm, metacarpal, and phalanges of digit III; length of
metacarpal and phalanges of digit IV; and length of metacarpal and phalanges of digit V.
Digit IV is projected at the mean alpha angle computed for each taxon (see Table Al).

(C, E) exert a positive force and that for the first phalanx a negative effect on the
ordination of emballonurids. Again, the molossids are ordinated in an opposite
manner. Interestingly, the centroid for the noctilionids is not carried in association
with the molossids, but is maintained in its same relative position in discriminant
space. As will be noted later, the vectors for elements of the third digit are more
directly involved in the ordination of noctilionids.

In the third canonical axis, the vectors for elements of digit III appear to
be less important in the overall ordination of these three centroids. In this axis,
vectors for the fourth and fifth digits are emphasized in a relative sense.

As stated above, variation of the third digit is difficult to describe because of its
involvement in the complex synergistic interactions among variables. However,
the net effect is a relatively long digit (Table A9). The metacarpal is particularly
important in this regard, judging from the high percentage contributed to the
discrimination vector of emballonurids (22.96, Table 6). The combined effect
of a long digit III and forearm is the production of a relatively long wing as can be
seen in Table A7. In fact, the high extremes in the range of variation are note¬
worthy. The relative length of the wing of Centronycteris is nearly three and a
half times (3.34) longer than the head and body length, which greatly exceeds
that for all bats. Likewise, Cyttarops exhibits an unusually long wing (2.91) as
compared to other chiropterans. These two species also fall at the high extreme
for relative length of digit III (Table A9).

Whereas the length of the third digit is important in the overall length of the
wing, the lengths of the fourth and fifth digits combine to determine the overall
aspect of the dactylopatagium. We have noted that in the rhinopomatids and
craseonycterids the length of digit III is generally offset by a relatively long

BIOLOGY OF THE PHYLLOSTOMATIDAE

261

fourth and fifth digit for the overall production of a short, broad tip. In the for¬
mation of high aspect tips, the trend is toward a relatively long fourth digit and
a shortened fifth digit. The emballonurids, noctilionids, and molossids generally
follow this trend, although the manner in which each responds is somethat dif¬
ferent.

The vectors for elements of the fourth digit (G, H, I) of emballonurids defy
easy interpretation because of their overall interaction with other variables.
In the first two canonical axes, these vectors imply shortness of the fourth digit.
However, in the third axis, a longish fourth metacarpal is suggested. The reader
will recall that the elements of the fourth digit are not particularly strong factors
in the ordination in the first three canonical axes, but that they gain strength in
the extradimensional fourth through sixth axes. In the fourth and fifth axes
(Table 4), the vector for the fourth metacarpal (G) is strong in its effect on the
ordination of the emballonurids and suggests a relatively long length for this
element. The contribution by this vector to the discrimination of the group also
is high (17.86, Table 6). A similar implication applies to the molossids, but to a
lesser extent — 9.90 per cent contributed to the function. This variable appears
to have only a minor role in the discrimination of noctilionids.

The vectors for the respective lengths of the two phalanges of digit IV do not
appear to be important in the overall ordination of the emballonurids. The general
implication is toward small size (Figs. 4D-E, 5, 6). However, the position of the
centroid relative to these two canonical vectors suggests a null effect, or at least
no significant elongation, when compared to the grand centroid for all
bats. The ordination of both the molossids and noctilionids are effected by one or
the other of these vectors. In the case of the molossids, a long first phalanx of
digit IV is emphasized, whereas a long second phalanx, in combination with a
short first phalanx, is suggested for the Noctilionidae.

The length of the fifth digit of emballonurids, as well as that of noctilionids
and molossids, is relatively short as compared to the total length of digits III and
IV, forearm, and head and body (Tables A7-A1 1). In a general sense, molossids
represent the extreme of this variation. The most striking differences among these
three groups is in the composition of this digit and specifically in the relative
length of the metacarpal element (Table A19). The vector for this wing element
(J) is directly involved in the ordination of the emballonurids and molossids,
and, to a lesser extent, noctilionids (Figs. 4D-E, 5, 6). The percentage
contributed by this vector to the discrimination of each of these groups is
9.88, 4.47, and 4.36, respectively (Table 6). This vector implies large size with
respect to this variable for emballonurids and noctilionids, but suggests small
size for molossids. The most important feature of the fifth digit of emballonurids
is a relatively long proximal phalanx (Table 6). This phalanx contributes nearly
a quarter of the total length of digit V (Table A20). Similarly, this phalanx is
distinguished as long in the noctilionids, but the importance in discrimination
of the group is slightly reduced (Table 6). The molossids, more than either of
these two groups, emphasize the length of the first phalanx of digit V (Table 6).
On the average, almost 30 per cent of the total length of the fifth digit is reflected

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by the first phalanx, the largest contribution noted among all bats (Table A20).
Cheiromeles and Otomops (both molossids) represent the high extremes with
39.13 and 35.30 per cent, respectively. The distalmost phalanx of the fifth digit
is markedly shortened in the emballonurids and molossids. The significance of
this reduction, in the overall ordination of these two families, is strongest for the
molossids compared to emballonurids (Table 6). On the other hand, this wing
element is markedly elongated in the Noctilionidae (Fig. 8), and the vector for
this variable (L) contributes 31.67 per cent to the discrimination vector of this
group. It is noteworthy to point out that this is the largest contribution by any
variable vector to the group discrimination vector of any group.

Thus we have seen that emballonurids possess wings that may be characterized
as relatively long and narrow (Fig. 7). An overall aspect ratio generally would
reflect this shape (Table A3), but would reveal little in terms of the composition
and interaction among the variables that produce such a shape. Outwardly, the
short tip index, relatively long wing, and low wing loading tend to confuse any
univariate or bivariate interpretation of this shape (Findley et ai, 1972). The
multivariate approach does help to clarify the issue. The wings of emballonurids
are truly high aspect in nature. However, a functional interpretation of this wing
shape is liable to be confounded if the wings of emballonurids are compared to
the high aspect wings of molossids. In such a comparison, one is likely to be biased
and misled by the apparent high correlation between high aspect ratio and
swiftness of flight, both attributes of molossids. In addition, generally high wing
loading appears to accompany the high aspect ratio of the molossids and not that
of emballonurids (Table A6).

We have shown that the construction of the wings of emballonurids differs
greatly from those of molossids and noctilionids, albeit the end product is vaguely
similar. Emballonurids appear to have modified a fundamentally short tip into a
long, high aspect tip by maintaining relatively long metacarpal elements and
elongating the terminal phalanx of digit III; the distalmost phalanges of digits
IV and V appear to be shortened. The development of a high aspect wing in this
manner may avoid allometric complexities associated with the modification of
more proximal wing elements. In addition, to achieve a high aspect wing, such
modifications might allow greater versatility. The highly maneuverable flight of
emballonurids is suggestive of a wide range of flight potentials. Some species
(notably those of Taphozous, Emballonura, Diclidurinae, and perhaps Centro-
nycteris ) appear to have capitalized on the speed qualities of high aspect wings.

Rhinolophidae

Horseshoe bats possess wings that average the lowest in overall aspect ratio
(5.41) as compared to all other bats (Table A3, Fig. 7). The length of the third
digit averages only slightly longer than the head and body (1.28). Also, the forearm
nearly equals the length of head and body (Table A8). These attributes combine
to produce a wing with next to the lowest average tip index (1.39) for all bats
(Table A2 and Fig. 10); only the rhinopomatids average lower.

BIOLOGY OF THE PHYLLOSTOMATIDAE

263

Fig. 10. — Bivariate graphs that illustrate the relationships between tip index and three
aspect ratios of the wing. Triangles represent phyllostomatid centroids and the grand centroid
for phyllostomatids is indicated by a circle with a plus. See list of specimens examined or
Table 5 for key to alphabetic code and text for discussion.

The synergistic relationships among the raw variables, discussed above for
the rhinopomatids and emballonurids, generally apply to the rhinolophids. A
relatively long forearm is implied by the vector for this variable (B). The ordination
of the group centroid for the rhinolophids, as well as that of the Nycteridae and
Megadermatidae, appear to be more strongly effected by vectors associated with
elements of the third digit. However, the relationships are difficult to characterize
because they are involved in a complex interaction among all variables. In the
first two canonical axes, the implication is toward shortness, whereas in the third
axis there is a general, but weak, expression of large size. Our general impression
is that these vectors describe the shortness of the digit as a whole, but the individual
components are either not affected or show only slight elongation.

With regard to the fourth digit, all variable vectors for the elements of this
digit (G, H, I) imply shortness in the first two canonical axes (Figs. 4D-E, 5, 6).
In the third axis, the vector for the fourth metacarpal (G) further emphasizes
shortness. However, in this third dimension of discriminant space, the vectors
for both phalanges (H, I) of digit IV suggest large size. The percentage of the
variance contributed to the discrimination vector of the group by the proximal
element of this series is exceptionally high (22.92, Table 6).

The vectors for the components of the fifth digit (J, K, L) are somewhat more
influential (Table 6) and all imply large size in the overall ordination of the

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rhinolophids. The vector for the distalmost phalanx (L) tends to elevate the
group centroid in the third dimension of discriminant space, but the combined
effect of the vectors for the metacarpal and the proximal phalanx (J, K) act to
suppress the elevation of the centroid.

The wings of rhinolophids, although perhaps not structurally as striking as
those of the emballonurids, craseonycterids, or rhinopomatids, do agree in
general structure and composition with wings of bats in these families. The
generalized distance between rhinolophids and these other group centroids is
relatively small — Rhinopomatidae (7.24), Craseonycteridae (4.23), and
Emballonuridae (5.92). The most notable difference between the wing of rhino-
lophoids and that of emballonuroids, and a feature that appears to distinguish
the former, is the short tip and generally broad aspect. The composition of the
wing in these two superfamilies appears to be similar and to reveal a relatively
close common ancestry.

Nycteridae and Megadermatidae

Because of their close association in discriminant space, (generalized distance
4.57), we will discuss these two families together. Although the megadermatids
average somewhat larger in general size than do nycterids and rhinolophids, all
three families are similar in general wing shape and composition (Fig. 7). The
ordination of these two families is influenced by vectors of nearly the same direc¬
tion and magnitude as discussed in the preceding account of the Rhinolophidae;
major differences are mostly quantitative rather than qualitative.

The mean aspect ratio of the wings of nycterids and megadermatids is only
slightly higher than that of rhinolophids (Table A3). The relative lengths of digit
III, and consequently the tip indices also, are similar (Tables A9, A2). The agree¬
ment among these values further attests to the qualitative similarity of wing shape
in these three families.

The major differences between the wings of these two families and the
Rhinolophidae appear to involve the two phalanges of digit III. The nature of
these quantitative differences is strong enough to produce a group discrimination
vector capable of consistently classifying the respective members of each family
(Fig. 17).

The first phalanx of the third digit is comparatively longer in nycterids than
in either rhinolophids or megadermatids. The vector for this variable contributes
19.33 per cent to the discrimination of the group (Table 6). The vectors for the
third and fourth metacarpal (C, G) of all three groups ordinate toward small size
as discussed in the account of the Rhinolophidae. The vector for the fifth meta¬
carpal (J) is slightly stronger in the ordination of the Nycteridae than it is in either
the Rhinolophidae or Megadermatidae (Table 6).

In the ordination of the Megadermatidae, the vectors for the third metacarpal
and second phalanx of this digit (C, E) are the strongest relative to these three
families and contribute 9.69 and 8. 17 per cent, respectively, to the discrimination
of the group. The vector of the former implies shortness, whereas the latter in-

BIOLOGY OF THE PHYLLOSTOMATIDAE

265

dicates large size. The combined effect appears to be elements of nearly equal
length. The phalanges of the fourth digit are slightly longer, in a relative sense, and
these vectors, likewise, are strong contributors to the discrimination vector of the
group (27.23 and 13.12 per cent, respectively).

Noctilionidae

Many of the distinguishing features of the wings of Noctilio were discussed
in the account of the Emballonuridae. The wings of both Noctilio albiventris and
N. leporinus are essentially alike in shape even though they differ markedly in the
absolute size of all raw variables. The wing of these two species are nearly two
and a half times the length of the head and body and almost 65 per cent of the span
is composed of the third digit. As a consequence, the tip index for the family is
high for the order (1.92 for N. albiventris and 1.98 for N. leporinus). Although
the overall aspect ratio of the wing is high and similar to that of molossids and
emballonurids, we have noted that the acquisition of this aspect is achieved through
different independent interactions among the elements that comprise the wing
in these three families (Figs. 7, 8, 9).

All vectors relating to features of the third digit (C, D, E, F) weigh heavily in
the ordination of the group. In addition, all but that for the first phalanx indicate
large size. The vectors for the most proximal phalanx of the third digit (D) and
fifth digit (K), as well as those for the fourth and fifth metacarpal (G, J), imply
smallness and tend to suppress the ordination of the group centroid in the third
canonical axis (Figs. 4E, 5).

Although the wings of Noctilio are high in aspect, we again caution comparisons
with the apparent swift flying ability of molossids. We have observed both species
in the field and would note that N. leporinus flies with a constant, but relatively
slow and shallow wing beat. It does not appear to be a particularly fast flier. The
smaller species, N. albiventris , is an insectivorous bat and from our observations
is capable of faster flight judging from the force with which individuals strike a
mist-net. N. albiventris also exhibits a fair amount of maneuverability in close
quarters and is capable of avoiding obstacles.

In our discussion of the Pteropodidae, we suggested that the possession of wing
elements of rather long span allowed for the control of large portions of the cam¬
bered surfaces. Slight flexion of these elements might greatly affect the camber
of the wing, in a manner similar to the downward deflection of the hinged flaps
on an airplane. This would contribute markedly to the lift potential at low speeds.
We further suggested that the nearly equal lengths of the manal elements of
pteropodids might allow for rather crude, yet effective, camber adjustments.
We continue this argument here and suggest that the shortening of a proximal
phalanx, especially in digits III and IV, would allow a greater range of variation
as well as finer dexterous control of the camber of the wing.

With regard to Noctilio , and perhaps mormoopids, the shortened first
phalanx in digits III and IV not only contributes to the high aspect construction,
but might account for the apparent versatility of flight behavior. Furthermore,

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in wings that have three phalangeal segments in the third digit, this means of
differential elongation of elements also may allow an increase in dexterity during
the “flick phase’’ of the wing beat cycle.

Mormoopidae

Bats of this family possess a relatively long wing, 63 per cent of which is
contributed by the third digit (Fig. 8). As we observed in the Emballonuridae, the
relatively long forearm may mask or otherwise offset the length of the tip. The
tip index of mormoopids (1.70) is only slightly higher than that obtained for
emballonurids and both values are well below average for all bats. Our data
suggest that mormoopid wings are well above average in overall aspect ratio
(Table A3) and that the tip can hardly be characterized as short. The mormoopids
appear to be closest, in wing morphology, to the Phyllostomatidae; misclassi-
fication occurred with the least derived species, Pteronotus parnellii, being
assigned to the Phyllonycterinae (Fig. 17).

The length of head and body is a relatively minor feature in the discrimination
of mormoopids (Table 6). Also, the length of forearm appears to have little effect
on the overall discrimination of the group.

The most important variable vectors in the ordination of the mormoopids
appear to be those associated with elements of the third digit (C, D, E, F) — long
metacarpal, short first phalanx, and long third phalanx are emphasized (Figs.
4D-E, 5, 6). The former two components of the mormoopid wing contribute the
most to the discrimination of the group (20.56 and 26.30 per cent, respectively).
Tables A12-A15 generally reflect these features. The percentage contributed to the
length of digit III by the first phalanx is nearly the lowest for all bats (11.18),
whereas that contributed by the distal phalanx is the highest (16.64). This appears
to be a general phyllostomatoid feature.

The effects of the vectors for elements of the fourth digit (G, H, I) are difficult
to interpret because of their apparent involvement in the overall synergistic
interaction among all variables. In the first and second canonical axes (Figs. 4D,
6), the vector for the fourth metacarpal (G) is oriented away from the group
centroid for the mormoopids and thereby implies shortness. However, in the third
axis (Figs. 4E, 5), this vector exerts a more positive force in the ordination of
the centroid. Both vectors for the phalanges of digit IV (H, I) indicate large size,
with emphasis on the distalmost phalanx. This terminal phalanx is not nearly so
long or apparently so important in the discrimination of the group as was observed
in the Noctilionidae (Table 6). The vectors for the corresponding pair of phalanges
in the fifth digit (K, L) also indicate large size with emphasis on the proximal
member. These two phalanges weigh heavily in the discrimination of the
group (Table 6) and appear to cause a lengthening of the fifth digit, which tends
to broaden the wing.

Vaughan and Bateman ( 1 970) presented an excellent discussion of the functional
myology of this group. They noted the remarkable maneuverability of these bats
and their rapid and sustained flight. Mormoops megalophylla is extreme in nearly

BIOLOGY OF THE PHYLLOSTOMATIDAE

267

all aspects of the wing. Mormoops blainvillii is rather curious in that the aspect
ratio of its wing is nearly equal to that of the larger-sized species M. megalophylla
(6.32), whereas its wing loading is a third lower (4.99 Nt/m2). Members of the
genus Pteronotus, and especially P. parnellii, appear to be less specialized in most
features of the wing.

Phyllostomatidae

The New World leaf-nosed bats, along with the noctilionids and mormoopids,
tend to dominate the upper right-hand quadrant of discriminant space (Figs. 5, 6).
Within this portion of space, each of the phyllostomatid subfamilies tends to occupy
a discrete region and group discrimination vectors generally distinguished each
of their centroids. There is a rather high percentage (22.30) of “misclassifica-
tions” (Fig. 17), which reflect a considerable amount of variation within the
family. The majority of these “misclassifications” involves species that occupy
a position near the grand centroid. Misclassifications outside of the family limits,
although fewer in number, also tend to occur in this region. Among phyllosto-
matids, the desmodontines exhibit the most fidelity to their group discrimination
vectors, whereas the carolliines show the least. We will consider the general
nature of phyllostomatid wing morphology before dealing with that of each of the
subfamilies.

As has been the case in previous accounts, the length of head and body of
phyllostomatids is of minor importance in the discrimination of the family
(Table 6). The range of variation of this variable is large and ranges from such
small-sized species as Ametrida centurio to the large-sized Vampyrum
spectrum. This variation nearly encompasses the range of variation observed for
the order.

The vector for the length of the forearm indicates small size with respect to this
variable for all phyllostomatid subfamilies (Figs. 4D-E, 5, 6). The absolute length
of the forearm averages slightly below the mean computed for all bats as does
the relative length of the forearm (Table A8). Table 6 indicates a rather strong
importance of the shortness of the forearm in the discrimination of most sub¬
families. This is strongest for the glossophagines, carolliines, and stenodermines,
but it is rather minor with regard to the phyllonycterines.

Although the dispersion of centroids is caused by the overall interaction among
all variables, the vectors that appear to influence most directly the ordination
of phyllostomatid centroids are those associated with features of the third digit;
most imply large size. The vector for length of the third metacarpal (C) apparently
is a strong factor in the discrimination of all subfamilies (Table 6). The tail end
of the vector for the first phalanx of digit III (D) is oriented toward the phyl¬
lostomatid centroids (Figs. 4D-E, 5, 6) and implies shortness (see also Fig. 8).
This vector is a moderately strong discriminator of the family (Table 6), although
it does not appear to be so important in the discrimination of the Carolliinae. The
proportionately long third phalanx (F) is a strong discriminator of nearly all phyl¬
lostomatid subfamilies (Table 6); phyllonycterines and desmodontines appear to
be less characterized by this variable.

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The overall effect of interplay among the elements of the third digit is the
production of a span that generally averages longer than that of any other group
of bats (Table A9). The length of this digit contributes nearly 67 per cent (range,
61.31-70.10) to the overall length of the wing, which is larger than in any other
chiropteran family. This is of interest in that the third digit of the molossid wing,
which is generally long-tipped and of high aspect, contributes a somewhat lower
average of 66.40 percent (range, 64.36-69.70) to the overall span of the wing. The
combined effect of the relatively short forearm of phyllostomatids and their long
third digits results in the highest tip indices of any group of chiropterans (Fig. 10;
Table A2), as also noted by Findley et al. (1972).

The vectors for elements of the fourth digit (G, H, I) pass tangentially to the
position of phyllostomatid centroids and a precise interpretation of their effect on
alar shape is difficult. The vector for the fourth metacarpal (G) suggests large size
in all three canonical axes (Figs. 4D-E, 5, 6). The vectors for the two phalangeal
elements of digit IV (H, I) appear to exert their greatest force on the ordination of
phyllostomatid centroids in the third canonical axis and here also imply large size.
Although there is variation within the family, as will be discussed below, the
second phalanx of the fourth digit tends to be proportionately longer than the first
(Tables A17, A18). Relative to the span of the wing, the fourth digit of phy-
lostomatids averages longer (60. 1 1 per cent of span) than does that of most other
groups of bats; only the molossids are larger in this respect (60.28 per cent of
span). In addition, the total length of this digit in phyllostomatids averages nearly
one and a half times the length of the forearm (range, 1.23-1.83).

Whereas phyllostomatids and molossids exhibit some similarities relative to
the lengths of digits III and IV, these two families are markedly dissimilar with
regard to the length of digit V. Indicative of the generally low aspect nature of
phyllostomatid wings, the fifth digit is long and averages 1.44 (range, 1.26-
1.68) times the length of the forearm. The vector for the second phalanx of this
digit (L) appears to be an important feature in the discrimination of all subfamilies
of phyllostomatids (Table 6). This variable has its strongest effect on the ordi¬
nation of phyllostomatid centroids in the third canonical axis where it implies large
size (Figs. 4E, 5). The vector for the fifth metacarpal (J), as that of the fourth
metacarpal, is difficult to interpret because it is oriented tangentially to the
phyllostomatid centroids (Figs. 4D-E, 5, 6). In the first and second axes, the
implication is large size, but shortness is emphasized in the third axis. The effect
of the first phalanx of digit V (K) on the ordination of phyllostomatids is some¬
what clearer, and it implies shortness in all three axes. The vector for the second
phalanx of digit V (L) suggests large size. The relative importance of these two
proximal elements in the discrimination of the phyllostomatid subfamilies is
variable but generally high (Table 6).

Finally, the structural, and perhaps phylogenetic, similarity of wing mor¬
phology among phyllostomatids may be summarized by examining the angles
between the discrimination vectors of each subfamily (Table 7). In this table,
the phyllostomatines are nearest the carolliines and glossophagines. The latter
two subfamilies are relatively close to each other as indicated by a 23.08 degree

BIOLOGY OF THE PHYLLOSTOMATIDAE

269

Table 7. — Angles between the group discriminant functions for the subfamilies of the

Phyllostomatidae.

Phyllostomatinae

Glossophaginae

Carolliinae

Stenoderminae

Phyllonycterinae

Desmodontinae

Phyllostomatinae

00.00

28.80

26.30

42.32

43.47

48.41

Glossophaginae

28.80

00.00

23.98

25.46

44.30

37.17

Carolliinae

26.30

23.98

00.00

35.02

48.38

56.18

Stenoderminae

42.32

25.46

35.02

00.00

45.68

42.78

Phyllonycterinae

43.47

44.30

48.38

45.68

00.00

45.48

Desmodontinae

48.41

37.17

56.18

42.78

45.48

00.00

divergence between their respective discrimination vectors. The stenodermines
fall nearest the discrimination vectors of glossophagines, carolliines, and phyl-
lostomatines, respectively. The most divergent angles between group dis¬
crimination vectors occur between phyllonycterines and desmodontines, and all
other subfamilies. The angle between the discrimination vectors of these two sub¬
families also is rather large (45.48 degrees). These relationships suggest that the
phyllostomatines form the nucleus of the family, which is rooted in proximity
to the grand centroid for all bats. The glossophagines and carolliines are positioned
relatively close to the phyllostomatines and these three subfamilies constitute
a core around which the remaining subfamilies are positioned. The stenodermines
appear to be morphologically most similar to the glossophagines, carolliines, and
phyllostomatines, respectively. The phyllonycterines and desmodontines occupy
widely separated positions from each other as well as from the other subfamilies.
The phyllonycterines appear to be morphologically nearer phyllostomatines and
glossophagines, respectively, than to other subfamilies, whereas desmodontines
appear to approach most closely the glossophagines.

Phyllostomatinae

The phyllostomatines are generally the largest bats of the family in terms of
absolute size; Vampyrum spectrum (40), Chrotopterus auritus (39), and
Phyllostomus hastatus (34) far exceed most New World species in overall size.
However, aside from these and several other large-sized species, the phyllosto¬
matines are about average or slightly below average in size. Compared to other
phyllostomatids, their wings are relatively long (Table A7) and the relative length
of the forearm averages longest of all phyllostomatids (Table A8). The relative
length of the third digit is average or slightly above average for the family (Table
A9). As a consequence of the interaction between these two lengths, the tip index
of phyllostomatines is comparatively low for the family (Table A2). In terms of
the overall aspect ratios, the wings of phyllostomatines are in the middle of the
range for the family (Tables A3-A5). Wing loading for this subfamily also is near

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SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

Fig. 11. — Canonical graph of the species of the subfamilies Phyllostomatinae, Carolliinae,
and Desmodontinae plotted on the first and third canonical axes. Stars represent the sub-
familial group centroids: A, Phyllostomatinae; B, Glossophaginae; C, Carolliinae; D,
Stenoderminae; E, Phyllonycterinae; F, Desmodontinae. Genera are encircled as follows:
Phyllostomatinae — Micronycteris (1-12), Macrotus (13-14), Lonchorhina (15-17), Macro-
phyllum (18-19), Tunatia (20-26), Mimon (27-31), Phyllostomus (32-36), Phylloderma
(37), Trachops (38), Chrotopterus (39), Vampyrum (40); Carolliinae — Carollia (41-44),
Rhinophylla (45-47); Desmodontinae — Desmodus (48-49), Diaemus (50), Diphylla (51).
Species are identified by corresponding bold-faced numbers in the list of specimens examined.

the median of the family, although the range of variation within the subfamily is
large (Table A6).

The centroid for the phyllostomatines is located near the grand centroid for
all bats. In the canonical graphs that show positions of individual species (Figs. 1 1,
12), it will be noted that the genus Micronycteris (1-12) encompasses the grand
centroid in the first three canonical axes. It is interesting to note here that the five
classificatory “misses” from this subfamily to the Vespertilionidae (Fig. 17)

BIOLOGY OF THE PHYLLOSTOMATIDAE

271

Fig. 12. — Canonical graph of the species of the subfamilies Phyllostomatinae, Carolliinae,
and Desmodontinae plotted on the first and second canonical axes. See legend of Fig. 1 1 for
key to group centroids (stars) and genera (encircled dots).

involve Micronycteris megalotis (1-2), M. pusilla (8), M. nicefori (9), and M.
behni (11). Most of the other species of phyllostomatines cluster together around
the centroid for the subfamily. However, there are several notable departures
from the group centroid.

Two species, V ampyrum spectrum (40) and Chrotopterus auritus (39), are
most obvious in their departure from the subfamilial centroid, especially along
the third canonical axis. Most of this dispersion appears to be caused by the vector
for length of the head and body. In addition, vectors associated with comparatively
short wings appear to affect these two species. In both, the lengths of forearm
and third digit are short as compared to other members of the subfamily (Tables
A8, A9). The span of the third digit is most influenced by the vector for the third
metacarpal, which implies shortness of this element in these two species (Table

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Fig. 13. — Canonical graph of the species of the subfamily Glossophaginae plotted on the
first and third canonical axes. Stars represent subfamilial group centroids (see legend of Fig.
11 for key). Genera are encircled as follows: Glossophaga (1-4), Monophyllus (5-6),
Leptonycteris (7-9), Lonchophylla (10-14), Lionycteris (15), Anoura ( 16-20), Scleronycteris
(21), Lichonycteris (22-24), Hylonycteris (25), Platalina (26), Choeroniscus (27-31),
Choeronycteris (32), Musonycteris (33). Species are identified by corresponding bold-faced
numbers in the list of specimens examined.

A 12). However, the lengths of the first and third phalanges average the largest
in percentage contributed to the length of digit III (Tables A13, A15). The
metacarpals of the fourth and fifth digit are proportionately short for the sub¬
family (Table A 16, A 19), although the phalangeal elements of these two digits are
generally long. The terminal phalanx of the fifth digit is comparatively longer
than in most other phyllostomatines (Table A21).

For the most part, the genus Phyllostomus (32-36) ordinates with the previous
two species in the first and second canonical axes (Fig. 12). However, Phyl¬
lostomus disassociates from this relationship in the third dimension of dis-

BIOLOGY OF THE PHYLLOSTOMATIDAE

273

Fig. 14. — Canonical graph of the species of the subfamily Glossophaginae plotted on the
first and second canonical axes. See legend of Fig. 1 1 for key to group centroids (stars) and
legend of Fig. 13 for key to genera (encircled dots).

criminant space (Fig. 11). In this axis, Phyllostomus tends to deny the influence
of length of head and body and is aligned by vectors that imply a long third digit.
The vector for the metacarpal (C) is especially important in this regard (Table
A 12). The first phalanx is the shortest among all members of the subfamily and
nearly the family as a whole (Table A13); only the vampire bats have a pro¬
portionately shorter first phalanx in the third digit. Other features that dis¬
tinguish Phyllostomus from most other phyllostomatines are long fourth and fifth
metacarpals (Table A16, A19), and short distal phalanx in digit V (Table A21).
These features also are characteristic of the vampire bats, and it is interesting to
note that all species of Phyllostomus, except P. latifolius (36) and a close associate
Phylloderma stenops (37), “misclassify” as desmodontines. The species latifolius
and stenops “misclassify” as stenodermines (Fig. 17).

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The genera Mimon (27-31) and Tonatia (20-26) portray an interestingly
antagonistic relationship to each other relative to the group centroid. This re¬
lationship is exaggerated by M. crenulatum (29-30) and M. koepckeae (31) on
the one hand and T. silvicola (25) and T. venezuelae (26) on the other. Generally,
Minion has the highest aspect ratio as compared to other phyllostomatines,
whereas Tonatia has the lowest (Tables A3-A5). Mimon has the longest wing,
in a relative sense, of any phyllostomatid, whereas the wing of Tonatia is much
shorter (Table A7). This relationship applies to most features examined in this
study. Incidently, the two extreme species of Mimon “misclassify” as stenoder-
mines (Fig. 17), which generally have longer, narrower wings as compared to
the other phyllostomatids. Other phyllostomatines that are “misclassified” (Fig.
17) are Micronycteris daviesi (12) and Macrophyllum macrophyllum (18-19),
which are aligned with the Glossophaginae.

Glossophaginae

The long-tongued bats tend to form a rather tightly packed cluster (Figs. 13,
14), which nestles close to the clusters of the phyllostomatines and carolliines
(Table 7). As a group, the glossophagines have relatively short wings as com¬
pared to other phyllostomatids (Table A7). The relative length of the forearm
averages a little over half (0.63) the length of head and body (Table A8). Com¬
paratively speaking, the third digit is relatively long, which produces a rather
large average tip index (2.06) for the subfamily (Table A2). The overall aspect
ratio of the wings of glossophagines is highest for the family — notable extremes are
Anoura (16-20) 6.50, Musonycteris ( 33) 6.30, and Scleronycteris { 21) 6.23. This
also applies to the aspect ratio of the tip region (Tables A3, A4).

In view of the tight packing of the group, a precise interpretation of the
variable vectors on the dispersion of glossophagines is difficult. Most of the
differences are small, quantitative shifts in the range of variation. The vectors
that appear to affect most heavily the ordination of the glossophagines are those
for the forearm (B), third metacarpal (C), and second phalanx in the fifth digit
(L). The vector for the forearm (B) implies shortness for most species. However,
Leptonycteris (7-9), Lionycteris (15), Scleronycteris (21), and Choeronycteris
(32) generally have longer forearms than other glossophagines (Table A8).

The vector for the metacarpal of digit III (C) suggests large size and Leptonyc¬
teris and Lionycteris represent the large extremes relative to this feature (Table
A 12). As a group, the glossophagines possess proportionately longer second
phalanges of digit V than do any other bats except the pteropodids (Table A21).

Two species, Hylonycteris underwoodi (25) and Platalina genovensium (26),
“misclassify” to the Stenoderminae and are most closely associated with Sturnira
and Vampyrops. Also, Lichonycteris (23-24) disperses among these stenoder-
mine genera, although its classification is mostly to the Glossophaginae.

Carolliinae

The group discrimination vectors for this subfamily are relatively weak. In
Fig. 17, two species, Rhinophylla pumilio (45) and R. fischerae (47) are “mis-

BIOLOGY OF THE PHYLLOSTOMATIDAE

275

classified” as glossophagines and four other species, Rhinophylla alethina (46),
Carollia subrufa (42), C. brevicauda (43), and C. perspicillata (44) are associated
with the Stenoderminae. This leaves only one species, Carollia castanea (41),
which suggests that, in terms of wing shape, the carolliines are rather indistinct
and may bridge the gap between glossophagines and the stenodermines (Figs. 1 1,
12; Table 7).

As a group, the carolliines have relatively long wings (Table A7). This results
from the combination of a moderately long forearm and an exceptionally long
digit III. These features also characterize the stenodermines. Carolliines further
resemble stenodermines in possessing a comparatively short digit IV; primarily
the result of a proportionately short fourth metacarpal (Table A 16).

Stenoderminae

Stenodermines represent the most diverse of the phyllostomatid subfamilies.
The dispersion of the various species of this subfamily in discriminant space is
comparable to that seen in the Phyllostomatinae, although the group generally
occupies space unfilled by other taxa (Figs. 15, 16). The group, as a whole, is
generally displaced away from the congested area nearer the grand centroid.
However, two small-sized species, Vampyressa pusilla (24) and Sphaeronycteris
toxophyllum (56), approach the grand centroid close enough to be confused with
the Vespertilionidae (Fig. 17). In addition, Phyllops haitiensis (51) and Centurio
senex (57) are “misclassified” as phyllostomatines, and Vampyressa nymphaea
(26), Pygoderma bilabiatum (54), and Ametrida centurio (55) are confused with
glossophagines.

Unlike any other subfamily of phyllostomatids, which tend to orient in uni-
modal directions in discriminant space, the stenodermines appear to ordinate
into two slightly different portions of this space (Figs. 15, 16). The extremes
of this dichotomy are Artibeus (35-47) on one hand and Vampyrops (13-22)
and Sturnira (1-10) on the other. Although the small-sized species of both groups
tend to congregate around the group centroid, the large-sized species of each
group orient away from each other (Fig. 16).

In the first three canonical axes (Figs. 15 and 16), vectors that imply large size
for the forearm (B), fifth metacarpal (J), and second phalanx of digit V (L)
ordinate Artibeus away from Vampyrops and Sturnira (Fig. 16). These vectors
imply shortness of these variables in both Sturnira and Vampyrops. The latter
two taxa are more directly ordinated by vectors associated with the third meta¬
carpal (C), and second and third phalanges of digit III (E, F). All suggest long
length.

The tip index and aspect ratio of the tip are generally higher in Vampyrops and
Sturnira than in Artibeus. As might be expected, Artibeus has a somewhat higher
aspect ratio of the plagiopatagial region, primarily as a result of a proportionately
longer forearm (Table A8). The composition of the third digit is similar in both
groups, although Artibeus tends to have a long metacarpal and generally short
phalangeal elements, whereas in Vampyrops and, to a lesser extent, Sturnira ,
construction of most of the span of this digit results from long phalangeal elements.

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III

I I

Fig. 15. — Canonical graph of the species of the subfamilies Stenoderminae and Phyl-
lonycterinae plotted on the first and third canonical axes. Stars represent subfamilial group
centroids (see legend of Fig. 1 1 for key). Genera are encircled as follows: Stenoderminae —
Sturnira (1-10), Urodermci (1 1-12), Vampyrops (13-22), Vumpyrodes { 23), Vampy ressa (24-
28), Chiroderma (29-33), Ectophylla (34), Artibeus (35-47), Enchisthenes ( 48), Ardops (49),
Phyllops (50-51), Ariteus (52), Stenoderma (53), Pygoderma (54), Ametrida (55), Sphae-
ronycteris (56), Centurio (57); Phyllonycterinae — Brachyphylla (58-59), Erophylla
(60-61), Phyllonycteris (62-63). Species are identified by corresponding bold-faced numbers
in the list of specimens examined.

Sturnira does not quite fit this scheme because the second phalanx is propor¬
tionately short (Table A 14). However, the proportional length of the distalmost
phalanx of the third digit appears to compensate for this (Table A15).

Phyllonycterinae

This subfamily, as well as the desmodontines, is ordinated into a peripheral
position of discriminant space relative to the other phyllostomatid subfamilies

BIOLOGY OF THE PHYLLOSTOMATIDAE

277

Fig. 16. — Canonical graph of the species of the subfamilies Stenoderminae and
Phyllonycterinae plotted on the first and second canonical axes. See legend of Fig. 1 1
for key to group centroids (stars) and legend of Fig. 15 for key to genera (encircled dots).

(Figs. 11-12, 15-16; Table 7). The flower bats have the shortest wings, in a
relative sense, among the Phyllostomatidae (Table A7). They resemble phyl-
lostomatines in possessing relatively long forearms (Table A8). The group has
the shortest relative length of digit III as compared to that of other phyllostomatids.
This is not particularly surprising inasmuch as the vectors for elements of this
digit (C, D, E, F) are oriented away from the group centroid (Figs. 4D-E, 15, 16).
The length of the third digit is composed primarily of the phalangeal elements,
which are equal or subequal in length (Fig. 8). As might be predicted from their
relative position in discriminant space, Erophylla bombifrons (60) and E. seze-
korni (61) “misclassify” as phyllostomatines (Fig. 17).

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Desmodontinae

The vampire bats occupy the most peripheral position in discriminant space
relative to all other phyllostomatid subfamilies. As there is complete fidelity to
their discriminant vectors, there are no instances of “misclassification” of members
of this group (Fig. 17), which suggests the distinctive shape of the desmodontine
wing (Fig. 8). This distinctness also is reflected in the rather large generalized
distance from the other phyllostomatid centroids: Carolliinae, 5.65; Stenoder-
minae, 4.46; Glossophaginae, 4.43; Phyllostomatinae, 4.28; and Phyllonycteri-
nae, 4.28. The most important vectors in the ordination of the group appear to
be those associated with the third metacarpal (C), which imply large size, and
those for the first phalanx of digit III (D) and first and second phalanx of digit V
(K, L), which emphasize shortness (Tables A12, A13, A20, and A21). Because of
the compensating effects of long metacarpal elements in the fourth and fifth
digits, the wing of vampire bats tends to be relatively short and broad and of gen¬
erally low aspect ratio. The vampire wing is the most heavily loaded of all phyl-
lostomatids (Table A6); note that the phyllonycterines follow the desmodontines
in this regard.

Natal idae

An interpretation of the alar shape of natalid wings is difficult. Part of this
results from the rather small sample size for this family as well as for other families
with which the natalids appear to be associated — namely, the Thyropteridae and
Craseonycteridae. Also, these three families appear to be associated with the
Vespertilionidae, for which there was a disproportionately large sample size.
Finally, the centroids of all four families as well as that of the Myzapodidae
lie in proximity to the grand centroid for all bats (Figs. 5, 6), tending to obscure
the precise relationships of one to another.

In the principal component analysis, the natalids, craseonycterids, and
thyropterids dispersed together towards the right-hand portion of Euclidean space
(Fig. 3), which, as we have noted above, indicates their general small size for all
variables. The vespertilionid centroid, although ordinated towards the small¬
sized side of the array, occupies a more central position in the overall dispersion.
On the other hand, the position of these four group centroids in discriminant
space is somewhat different (Figs. 5, 6).

The natalids align most closely with the craseonycterids in the discriminant
analysis. The shared absence of the third phalanx of digit III appears overly to
bias this association. On the basis of this variable alone, the generalized distances
between natalids/craseonycterids, thyropterids, and vespertilionids; craseonycte-
rids/thyropterids and vespertilionids; and thyropterids/vespertilionids are:
0.093, 1.466, and 2.016; 1.466 and 2.016, and 0.550, respectively. The overall
generalized distances between these centroids are 4.050, 3.540, and 4.400;
5.580 and 5.457; and 2.489, respectively. However, the generalized distances
between these four families, on the basis of each variable, tend to indicate a closer
association between natalids, thropterids, and vespertilionids than between
craseonycterids and these three families.

BIOLOGY OF THE PHYLLOSTOMATIDAE

279

Thus, the resemblance of natalids and craseonycterids might be spurious as
a result of the absence of the third phalanx of digit III and a concomitant com¬
pensation in the length of elements of this digit, especially that of the second
phalanx (Fig. 9). In addition, there appears to be a “general” tendency for small¬
sized bats to have similarly constructed wings (that is, long forearm, long digit
III, and generally long digits IV and V). Findley et al. (1972) also noted this
tendency, but we would caution the reader by noting that some relatively large¬
sized bats, such as noctilionids, emballonurids, and nycterids (among others), also
follow this trend (Tables A2, A7, A8-A1 1). Hence, we reiterate our earlier state¬
ment that the relationships between general body size and wing morphometries
are much more complicated than bivariate comparisons would seem to indicate.

In the first three canonical axes (Figs. 4D-E, 5, 6), the ordination of the natalid
centroid appears to be affected by interactions among variables, similar to those
noted above for the craseonycterids. In the previous accounts, we have discussed
the apparent minor role of the length of the head and body in the discrimination
of groups. With regards to the natalids as well as the thyropterids and myzapodids
the influence of this variable, albeit weak, is comparatively stronger than noted
for other families (Table 6). The relative length of the wing of natalids is 2.61 times
the length of the head and body and is among the longest found among all bats
(Table A7). This span is composed of a relatively long forearm (Table A8), and
digit III has a mean relative length (1 .69) that is highest among all bats, Table A9.
Similarly, large values for these relative lengths will be noted for craseonycterids,
thyropterids, and furipterids.

The composition of the third digit of natalids is more like that of thyropterids
and vespertilionids than that of craseonycterids. The vector for the third meta¬
carpal (C) of natalids implies shortness as was the case in the Craseonycteridae.
However, the reader will recall that the second phalanx of digit III offset the
proportional length of the third metacarpal in the craseonycterids. In the
extradimensional fourth and fifth canonical axes, the vector for the third meta¬
carpal more strongly implies longness of this variable for natalids, thyropterids,
and vespertilionids. This also is generally the case for the first and second phalanx
of digit III for these three families.

The combined effect of variable vectors for elements of the fourth digit (G, H,
I) of natalids indicates longness of this digit (Table A 10). The most important
components of length appear to be the phalangeal elements, although these are
generally below the average computed for all bats (Tables A17, A18). The length
of the first phalanx of digit IV contributes markedly to the discrimination of the
group (Table 6). Again, an interpretation of the vectors for this digit is obscured
by the synergistic interaction among all variables. Shortness of the fourth meta¬
carpal (G) is suggested in the first three canonical axes. However, in extradimen¬
sional axes this vector implies longness of this variable in natalids, thyropterids,
and vespertilionids; shortness is indicated for that of craseonycterids.

The relative length of the fifth digit of natalids averages the longest among all
bats (Table All); the mean relative length of this digit for furipterids and thy¬
ropterids also is high. The variable vectors for the lengths of the metacarpal (J)

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and second phalanx (L) strongly suggest longness in the first three canonical axes
(Figs. 4D-E, 5, 6), whereas that for the length of the first phalanx of digit V
implies shortness.

In general appearance (Fig. 9), the wings of natalids are below average in their
overall aspect ratio (Table A3). This low aspect also is reflected in the aspects
of the tip and plagiopatagial portions of the wing (Fig. 10; Tables A4, A5).
Craseonycterids, thyropterids, furipterids, myzapodids, and to a certain extent,
vespertilionids resemble natalids in these respects. It is interesting to note that,
with regard to wing loading, the craseonycterids possess more heavily loaded wings
than do any of the five aforementioned families (Table A6).

Little is known concerning the flight characteristics of natalids. We concur with
Findley et al. (1972) in their suggestion of slow, maneuverable flight potential
for these bats; also, hovering may be well within this potential.

Thyropteridae

The interpretation of wing morphometries of the disc-winged bats is obscured
by the positioning of their group centroid almost exactly on the grand centroid of
all bats (Figs. 5, 6). This, in itself, reflects the average character of the shape of
their wings. However, the confidence circle for the group centroid is com¬
paratively large, possibly reflecting the rather small sample size utilized in this
study.

In the classificatory phase of the discriminant analysis (Fig. 17), both species of
thyropterids “misclassify” as vespertilionids. This could reflect a correct assign¬
ment or it simply might be an artifact of small sample size. The generalized distance
between these two families is comparatively small (2.489) and the generalized
distances, based on each variable, likewise support this close association of the
two.

Myzapodidae

Little can be said concerning the shape of the wing of Myzapoda aurita. The
group centroid is in proximity to the grand centroid for all bats (Figs. 5, 6); the
confidence circle exceeds the limit of the figures and probably reflects the small
sample size of two specimens. In the classificatory phase of the discriminant
analysis, these bats as well as the thyropterids (noted above) were “misclassified”
as vespertilionids.

Vespertilionidae

The members of this family are extremely diverse in the shapes of their wings
and presumably in their flight characteristics. The group centroid is located
near the grand centroid for all bats (Figs. 5, 6), but unlike the previous two groups
the confidence circle is small, and the group discrimination vector appears to be
relatively strong. The one “misclassification” from this family involved Eudiscopus
denticulus, which was confused with the Phyllostomatinae (Fig. 17). Several other
species of vespertilionids were associated with phyllostomatid subfamilies, but

BIOLOGY OF THE PHYLLOSTOMATIDAE

281

only this one was so classified. In previous accounts, it was noted that several
phyllostomatids, as well as thyropterids and myzapodids, were incorrectly assigned
to the Vespertilionidae. Aside from possible errors associated with sample size,
we suspect that these “misclassifications” reflect general similarities among these
species as a result of their proximity to the chiropteran norm (grand centroid).

Generally, the wings of the vespertilionids are moderately long and average
about twice (2.07) the length of head and body (Table A7). The range in variation
is markedly large and extends from Mimetillus moloneyi, with its peculiarly-
shaped wing (barely 1.4 times the length of its head and body), to Otonycteris,
Kerivoula, Miniopterus, and Eudiscopus, wings of which are nearly 2.5 times
the head and body length.

The vector for length of the forearm (B) contributes moderately to the group
discrimination vector of the family (Table 6). The mean relative length of the
forearm is slightly above average for all bats (0.74), but the range within the
family includes nearly the total of variation exhibited by the order (Table A8).

The position of the group centroid relative to the vectors associated with the
elements of digit III (C, D, E, F) generally reflects the emphasis on the long length
of this digit in the composition of the wing (Figs. 4D-E, 5, 6). The mean tip index
for the family (1.81) is slightly below the average for all bats (Table A2), but
the range in variation includes values that are twice the length of the forearm
(for example, Eudiscopus, 2.17; Kerivoula, 2.12; Harpiocephalus, 2.04; and
Lasiurus, 2.00). In the first three canonical axes, the vector for the meta¬
carpal (C) implies large size. The percentage of variation contributed to the group
discrimination vector by this vector is relatively high (18.60, Table 6). Table A12
shows that, on the average, approximately 50 per cent of the length of the third
digit is accounted for by this element. As has been the case for the majority of
the families discussed to this point, the vector for the first phalanx of digit III (D)
implies shortness. On the whole, vespertilionids fall just below the average for all
bats with respect to this feature (Table A13). The vector for the length of the
second phalanx of digit III (E) nearly equals the metacarpal in its influence in
the discrimination of the group centroid (Table 6). The implication of this variable
vector is shortness and the mean percentage contributed to the length of the digit
III (Table A14) tends to support this. The high extremes in the range of variation
of this percentage are noteworthy. The second phalanx constitutes 33.77 per
cent of the total length of digit III in Miniopterus. Similarly, this phalanx is
proportionately long in Lasionycteris, Chalinolobus, and Kerivoula (27.31,
26.77 , and 26.42 per cent, respectively). The vector for the length of the third
phalanx of digit III (F) is moderately important in the discrimination of the family.
However, its precise effect on the dispersion of the group centroid is difficult to
assess because this phalanx is indistinguishable or absent in some species and
markedly elongate in others. In most vespertilionid species, this phalanx com¬
prises 10 per cent or less of the length of digit III (Table A15); 20.13 per cent is
contributed by this element in the wing of Eudiscopus.

The interaction among the elements that compose the fourth digit is complex
and, as will be noted in Tables A16-A18, the range of variation is wide. The

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vector for the metacarpal (G) implies large size. The effect of this vector on the
ordination of the vespertilionids appears to be similar to that exerted in the
Molossidae (Figs. 4D-E, 5, 6), although, in the latter, all vectors associated with
elements of the fourth digit appear to apply a more direct force on the ordination.
The first phalangeal element of the fourth digit is about average in its pro¬
portional length as compared to that of other bats (Table A17). The vector for the
length of the second phalanx of digit IV (I) emphasizes shortness and this is
generally supported in Table A18, although, again, the range of variation is wide.

The length of the fifth digit of vespertilionids appears to be controlled mainly
by the length of the metacarpal element. The vector for this variable (J) is important
in the overall ordination of the vespertilionids as indicated by the relatively high
percentage (16.53) contributed to the discrimination vector of the group (Table
6). The vespertilionids rank second highest with regard to the mean per cent
contributed by the fifth metacarpal to the total length of digit V (Table A 19).
Whereas noctilionids average larger than vespertilionids with regard to the
proportional length of the fifth metacarpal, the high extremes in the range of
variation among vespertilionids far exceed that of any other bats. Notable among
these extremes are Mimetillus (82.35 per cent), Philetor (75.87),
Scotophilus (73.07), Tylonycteris (72.84), and Nyctalus (72.56). The vector for
length of the first phalanx of digit V (K) implies shortness, but this variable is
of minor importance in the discrimination of the group (Table 6). The vector for
the length of the distal phalanx of this digit (L) is somewhat stronger in its influence
on the group discrimination vector (Table 6) and it suggests shortness.

In a general descriptive sense, the wings of vespertilionids are not particularly
striking; they are about average or slightly below average in most respects as
compared to those of other members of the order. However, in terms of internal
composition, wing variation in vespertilionids is the most complex of any family
we have examined. This is particularly true of species that depart from the family
norm, that is, those vespertilionids with wings of higher than average aspect ratio.

To illustrate some of this variation, we can examine the construction of the tip
region in three species — Eudiscopus denticulus, Lasiurus cinereus, and Mimetillus
moloneyi. The aspect ratio for the tip followed by the tip index (in parentheses) for
each of these species is 5.98 (2.16), 5.04 (1.99), and 4.41 (1.59), respectively. In
Eudiscopus , the third metacarpal is proportionately short (41.82 per cent of
digital length), the bulk of the length being contributed by the phalangeal elements,
especially the third phalanx. The fourth and fifth metacarpals are proportionately
longer (61.55 and 64.55 per cent, respectively) than the third, but nearly half
the length of each of these digits is accounted for by the phalanges. In Lasiurus,
the metacarpals are proportionately longer (54.07, 65.01, and 70.63 per cent,
respectively) than in Eudiscopus, and the first and second phalanx of digit III
account for most of the remaining length of this digit; the third phalanx is markedly
shortened. The phalanges of digits IV and V are nearly equal in length. The third
metacarpal of Mimetillus is proportionately longer than that of either of the two
aforementioned species (63.40 per cent of the length). The third phalanx of digit
III is indistinguishable, and the remaining two are about equal in length. The

BIOLOGY OF THE PHYLLOSTOMATIDAE

283

majority of the length of the fourth and fifth digits is contributed by the metacarpal
elements (73.59 and 82.35 per cent, respectively). The second phalanx of digit
IV is much reduced, (comprising less than five per cent of the length of the digit).
Both phalanges of digit V are extremely short and equal or subequal in length.
Together, they comprise 17.65 per cent of the length of this digit.

These three species are only exemplary of the kinds of variation that exist
within the family Vespertilionidae. This would seem to confirm the wide variety
of flight behaviors reported for the family, which range from the swift,
sustained flight of migratory species to the erratic, highly maneuverable flight of
some of the smaller nonmigratory species. Norberg (1972, 1976a, 1976/?) has
clearly demonstrated the hovering ability of Plecotus auritus, and certainly other
species will be shown to possess this flight behavior.

Mystacinidae

The group centroid for this rather unusual, monotypic family ordinates into the
upper right-hand quadrant of discriminant space (Figs. 4D-E, 5, 6). As we have
noted above, this portion of discriminant space is defined generally by a relatively
long and broad chiropatagium and relatively short and broad plagiopatagium.
The centroid of Mystacina tuberculata is most closely associated with that of the
Mormoopidae in the first two canonical axes (Fig. 6). However, interactions among
variable vectors in the third canonical axis (Figs. 4E, 5) cause a rather marked
dissociation of these two centroids, suggesting basic differences in the composition
of the wings of these two families.

The effect of the vector for length of forearm (B), albeit weak as compared to
that of other groups of bats, is somewhat stronger in discrimination of Mystacina
than in mormoopids (Table 6). In both groups and in the first two canonical axes
(Figs. 4D, 6), this variable vector generally suggests longness. In the third canonical
axis (Figs. 4E, 5), the mystacinid centroid appears to be more strongly influenced
by the tail (smallness) end of this vector, whereas the mormoopid centroid is
aligned closer to the positive (longness) end. The relative length of the forearm
of Mystacina ranks slightly below the mean for all bats: mormoopids rank above
this mean (Table A8). This indicates a somewhat greater length of head and body
for Mystacina as compared with that of mormoopids.

Interactions among variable vectors associated with length of digit III (C, D, E,
F) of Mystacina are similar to those discussed for mormoopids. The vector for
the length of the third metacarpal (C) of both these families implies large size
(Figs. 4D-E, 5, 6). The proportional length of this wing element is slightly greater
in Mystacina than in mormoopids and, in both, contributes more than 50 per
cent to the length of digit III (Table A12). As appears to be typical of bats arrayed
in this portion of discriminant space, the vector for the length of the first phalanx
of digit III (D) suggests shortness. This wing element comprises only 14.33 per
cent of the length of the third digit in Mystacina, which is only slightly higher than
that contributed in mormoopids (Table A 13). These two variable vectors appear
to be important in the group discrimination vectors of both Mystacina and mor¬
moopids (Table 6). Although the variable vectors for the two distal phalanges

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(E, F) of both families suggest long length, the contribution of each of these ele¬
ments to the wing of these two groups is somewhat different. The proportional
lengths of all three phalangeal elements are maintained nearly equal or subequal
in the wing of Mystacina (Fig. 9; Tables A13-15). On the other hand, there ap¬
pears to be a definite allometric relationship among these phalangeal elements
in the wing of mormoopids. The relative length of the third digit of the mystacinid
wing lies below the average for all bats (Table A9). Likewise, the tip index of
Mystacina is below the average computed for all bats (Table A2). However, as
noted above, the long forearm tends to mask the length of digit III in these bats
as well as in emballonurids and mormoopids.

The effects of the vectors for elements of the fourth digit (G, H, I) of Mystacina
are similar to those discussed for mormoopids. The vector for the length of the
fourth metacarpal (G) indicates shortness in the first two canonical axes (Figs.
4D, 6), but a slightly positive (longness) influence is suggested in the third axis
(Figs. 4E, 5). The proportional length of this element is well above the average
for all bats (Table A16). The variable vectors for the phalangeal elements of
digit IV (H, I) both imply longness. In terms of the group discrimination vector,
the variable vector for the first phalanx of this digit (H) appears to be important
(Table 6). The proportional length of the second phalanx of digit IV ranks slightly
above the mean for all bats and this element contributes 21.04 per cent of the
length of the digit (Table A 18).

The greatest differences in composition of the wing of Mystacina and that of
mormoopids concern features of the fifth digit. In Mystacina , variable vector for
the length of the fifth metacarpal (J) suggests longness in the first two canonical
axes (Figs. 4D, 6). However, the implication shifts toward smallness in the third
axis (Figs. 4E, 5). Paradoxically, the proportional length of this wing element
(67.42) ranks well above the average for all bats (61.02), whereas that for mor¬
moopids (59.29) falls below the average (Table A19). This variable vector ap¬
pears to be relatively unimportant in the discrimination of the Mystacinidae
(Table 6). The strongest vectors in this regard are those for lengths of the first and
second phalanx of digit V (K L). The vector for the first phalanx (K) strongly
suggests shortness in all three canonical axes (Figs. 4D-E, 5, 6). The proportional
length of this element averages the shortest among all bats (Table A20); mor¬
moopids rank above the overall average with regard to this feature. On the other
hand, the vector for the length of the second phalanx of digit V (L) strongly
implies longness and this element contributes 21.00 per cent to the length of this
digit (Table A21).

The overall aspect ratio of the wing, as well as that of the tip, of Mystacina falls
slightly below the average of all bats (Tables A3, A4). However, the relatively
long forearm and comparatively short fifth digit contribute to the higher than
average aspect ratio of the plagiopatagium (Table A5).

Little is known concerning the flight behavior of Mystacina tuberculata. The
family is endemic to New Zealand where it and Chalinolobus tuberculatus
(Vespertilionidae) comprise the total chiropteran fauna. The phylogenetic re¬
lationships of the family are poorly understood although relationship to the

BIOLOGY OF THE PHYLLOSTOMATIDAE

285

Molossidae has been suggested by various authors (Dobson, 1875, Miller, 1907).
In terms of wing shape, Mystacina most closely resembles mormoopids and
phyllostomatids. This is particularly interesting in view of Daniel’s (1976) recent
report on the food habits of Mystacina in which he included fruit and possibly
nectar along with aerial and terrestrial insects in the feeding regime. If the mor¬
phometric resemblance between Mystacina, mormoopids, and phyllostomatids
is conveyed in functional similarity, the wing of Mystacina should be found to be
relatively versatile.

Molossidae

The shape of the wing in this family is perhaps the most distinctive among all
bats. The molossid wing is extremely narrow and has an unusually long tip region
(Fig. 9). As a consequence, the wing is highest in overall aspect ratio among
bats. We have already discussed some features of molossid wings in the accounts
of emballonurids and noctilionids. Of particular interest is the fact that, even
though the bats of these three families possess wings of high aspect, the mode by
which their wings are constructed is markedly different.

Whereas the forearm is usually long in most other groups of bats, especially those
that possess high aspect wings, the relative length of the forearm of molossids
averages the shortest among all bats (Table A8). The vector for this variable (B)
is oriented almost directly away from the group centroid in the first three canonical
axes and thereby suggests shortness (Figs. 4D-E, 5, 6). The forearm contributes
only 30 to 35 per cent to the total span of the wing. Among molossids, Cheiromeles,
Otomops, and Eumops possess the largest forearms, whereas Sauromys and
Molossus have the shortest. As the orientation and length of the variable vector
indicate, the length of the forearm is an important factor in the group dis¬
crimination vector (Table 6).

The great length of the wing is reflected in the generally positive orientation of
all vectors associated with elements of the third digit toward the molossid centroid
(Figs. 4D-E, 5, and 6). The vectors for the metacarpal, and second and third
phalanges (C, E, F) are not as positively associated with the molossid centroid as
was noted for the noctilionids, mormoopids, phyllostomatids, and vespertilionids.
Nonetheless, these vectors do imply longness of these elements in the Molossidae.
The vector for the first phalanx of digit III (D) strongly suggests longness in the first
two canonical axes and to a certain extent in the third axis. Proportionately, the
length of this phalanx (19 to 26 per cent of the length of digit III) averages among
the largest for all bats (Table A1 3). Although the proportional length of the second
phalanx averages below the mean for all bats (Table A 14), these two phalangeal
elements in consort with the metacarpal produce the major portion of the span
of digit III. It is difficult to interpret the vector for the third phalanx of this digit
because, by comparison, it is rather short. However, this vector appears to be
rather important in the group discrimination vector (Table 6). In this case, the
vector seems to imply simple presence of the phalanx rather than length. Shortness
or absence of the distal phalanx of digit III seems to be the case in other families

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that have proportionately long first and second phalanges (for example, embal-
lonurids, noctilionids, and vespertilionids).

In most of the other groups of bats considered in this study, vectors associated
with elements of the fourth digit (G, H, I) are not easily interpreted, mostly
because of their tangential orientation to centroids and their synergistic inter¬
action with other variables. All three of these vectors are directed toward the
molossid centroid and all imply longness. The most powerful among these are the
vectors for length of metacarpal (G) and second phalanx (I). In addition, these
two vectors are important in the group discrimination vector (Table 6). The
relative length of the fourth digit is not particularly impressive and it averages
below the mean for all bats (Table A 10). However, this value is greatly masked by
the generally long length of head and body of these bats. The fourth digit of
molossids constitutes nearly 60 per cent of the span of the wing, and in these terms,
is the largest among all bats. The metacarpal alone contributes 55 to 69 per cent
of the length of this digit (Table A16). The first phalanx constitutes the bulk of
the remaining length (18 to 28 per cent, Table A17). The length of the second
phalanx of digit IV is variable and can contribute as much as 18.11 per cent
( Sauromys ) or as little as 2.88 and 3.94 ( Tadarida and Promops, respectively)
to the length of this digit. Eumops and Molossus, on the average, possess a rather
short second phalanx on digit IV.

Unusually long third and fourth digits have been discussed in the accounts
of several groups, especially the Phyllostomatidae, which have generally long-
tipped, low aspect wings. Perhaps the most striking feature of the molossid wing
is the markedly short fifth digit, which converts the long tip region into a high
aspect surface. The vector for the length of the fifth metacarpal (J) strongly implies
shortness in the first three canonical axes (Figs. 4D-E, 5, 6). Similarly, the vectors
for the lengths of the two phalangeal elements of this digit (K, L) orient away
from the molossid centroid and thereby imply shortness. All three of these
variable vectors are important factors in the discrimination of the group (Table
6).

In other high aspect wings such as those of emballonurids and noctilionids,
the shortening of the fifth digit is accomplished by shortening the phalangeal
elements while maintaining the metacarpal more or less isometric with the third
and fourth metacarpals. If the apparent versatility in flight behavior of these
bats is any indication, we could assume the formation of a high aspect wing in this
fashion to be a less than total commitment to swift flight. On the other hand, by
shortening the fifth metacarpal, molossids gain dexterous control of a smaller
portion of the camberable surface but at the same time might lose a sizable degree
of flight versatility. In this light, it is interesting to note that the genus Tadarida
(the most diverse, yet least specialized, of the family with some 45 or so species)
exhibits a wide range of variation in the composition of the fifth digit and other
digital elements.

To illustrate the degree of variation in wing composition within the Molossidae,
we have used Cheiromeles , Otomops, Sauromys, and Tadarida. Cheiromeles

BIOLOGY OF THE PHYLLOSTOMATIDAE

287

torquatus is the largest molossid, with a head and body length of 115 to 135
millimeters and a weight of 150 to 170 grams. The proportional lengths of the
metacarpal elements of digits III to V are shortest among the family (43.21,
54.62, and 49.37 per cent contribution to digital length, respectively). On the other
hand, the phalanges of the third and fourth digits are proportionately longer than
those of any other molossid. The first phalanx of the fifth digit is proportionately
longer than that of any other molossid, comprising nearly 40 per cent of the length
of the digit; the second phalanx is about average for the family (1 1.50 per cent of
digital length).

On the average, Otomops possesses the proportionately longest third and
fourth metacarpals of any molossid (54.42 and 68.78 per cent of the digital length,
respectively), although individual species of Tadarida and Eumops possess longer
fourth metacarpal elements (72.52 and 70.18 per cent, respectively). The pro¬
portional lengths of phalangeal elements vary in Otomops. Generally the major
portion of the length of digit III is contributed by the second, first, and third
phalanx (21.18, 20.59, and 4.65 per cent, respectively). Otomops possesses the
shortest first phalangeal element of digit IV of the family (17.82 per cent of the
digital length), and the proportional length of the second phalanx (7.21 per cent)
is well below the average for the family. Whereas the metacarpal of digit V is ex¬
tremely short, the proportional length of the phalanges of this digit are nearly the
largest for the family (35.27 and 12.57 per cent, respectively).

Whereas the two genera discussed above might be considered as among the more
specialized molossids, Sauromys appears to be among the least specialized. The
metacarpal elements of digits III and IV are proportionately average for the fam¬
ily (50.81 and 59.05 per cent, respectively); the fifth metacarpal is unusually long
for the family (63.56 per cent of the digital length). The proportional lengths of
the first and second phalangeal elements of digits III to V vary although they are
generally isometric and range between 22 and 1 5 per cent of the digital length. The
third phalanx of digit III is proportionately long for the family (7.93 per cent of
the digital length).

Finally, Tadarida is perhaps the most variable among the molossids in terms
of wing composition. The proportional lengths of the metacarpals of digits III to
V rank near the family average, but the range is broad (53.97 to 46.02, 72.52 to
57.16, and 67.51 to 53.55 per cent of the digital length, respectively). There is a
general trend of isometry among the proportional lengths of the first and second
phalanx of digit III (20.59 to 19.81 and 23.34 to 18.02 per cent, respectively).
The proportional length of the third phalanx of this digit varies (10.80 to 4.92
per cent). With regard to the phalanges of the fourth digit, patterns of allometry
and isometry vary markedly, especially with respect to proportional length of
the second phalanx (27.10 to 19.94 and 20.49 to 2.88 per cent of digital length,
respectively). Freeman (1977) noted this allometric variation in the composition
of digit IV and interpreted it in terms of zoogeographic distribution. Allometry
is even more pronounced in the proportional lengths of the phalangeal complement
of the fifth digit (36.43 to 21.81 and 15.05 to 7.97 per cent, respectively).

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SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

Generally speaking, the wings of molossids are highly specialized, although
we remind the reader that within this family a degree of variability exists. Wing
loading is normally high (Table A6): Eumops averages highest in the family (28.47
Nt/m2; Eumops auripendulus is highest among all bats with 58.00 Nt/m2); and
Tadarida, although nearly average in this feature (19.54 Nt/m2), exhibits loadings
down to 8.43 Nt/m2. The general composition of the molossid wing suggests a
reduction in the control of camberable surface. No doubt the “automatic” flexing
and extending devices in the elbow and shoulder regions discussed by Vaughan
( 1 959, 1 970a) relate to this alar composition. We suspect that the more generalized
species of the family will be shown to have a greater degree of “manual” control
of their flight surfaces.

Classification

As has been discussed above, in the discriminant analysis a discrimination
vector is computed for each group (in this case families or subfamilies) based on
the synergistic interaction among variables. In the classification phase of the
analysis, each case (species in this analysis) is scrutinized and assigned to that
group to which it is most closely aligned in discriminant space (Fig. 17). Inasmuch
as the discrimination vector for each group is an expression of the complex
qualitative and quantitative aspects of wing shape, species are grouped together
based on similarity of wing shape.

In the overall classificatory analysis, only 14 of 466 species (three per cent)
were incorrectly assigned. The high degree of correct associations appears to
indicate a rather large phylogenetic component in the overall shape of bat wings.
“Misclassifications” may be attributed to several possible sources of error.

The first of these is insufficient sample size, which could have greatly effected
the formulation of an accurate discrimination vector for various groups. We
suspect this might be the case with regard to the Thyropteridae and Myzapodidae,
in which the sample sizes were extremely small. We would not be particularly
surprised if the association of these two families with the Vespertilionidae was
found not to be related to the sample size, because the shape of the wing in these
two families is in fact similar to that of the vespertilionids. Another source of
error involves “leakage” of taxa that ordinate close to the grand centroid for all
bats. This we suspect is the explanation for most of the “misclassifications”
encountered in the Phyllostomatidae.

Yet another source of error might be that of functional similarity. With regard
to the two species assigned to the Megadermatidae, as well as Hypsignathus
monstrosus (Pteropodidae) and Rhinolophus Indus (Rhinolophidae), it is note¬
worthy to point out that each has a relatively long second phalanx in digit III,
which is a major feature of megadermatids. Similarly, the association of Pteronotus
parnellii with the Phyllonycterinae appears to relate to the overall similarity of
alar shape, especially with respect to length of forearm. As noted above, this
association also may reflect some phylogenetic similarity between mormoopids
and phyllostomatids. One molossid species, Tadarida loriae , is classified as a

BIOLOGY OF THE PHYLLOSTOMATIDAE

PTEROP
RHINOP
CRASEO
EMBALL
RHINOL
NYCTER
MEGAD
NOCTIL
MORMO
PHYLOS
GLOSSO
CARO LL
STENOD
PHYNYC
DESMOD
NATAL L
THYROP
MYZAPO
VESPER
MYSTAC
MO LOS

Fig. 17. — Classification graph from the discriminant analysis. Numbers on the diagonal
represent number of species correctly associated by the group discrimination vector for each
group, with their respective taxonomic category. Numbers in rows, and off the diagonal,
represent number of “misclassifications" to other taxonomic groups. This analysis resulted in
97 per cent correct associations. See text for discussion.

vespertilionid. This is not surprising because other generalized species of Tadarida
are ordinated toward the vespertilionid dispersion.

The Phyllostomatidae, as a whole, illustrates a rather high affinity to its various
group discrimination vectors; only 4.32 per cent of its species are assigned outside
the limits of the family. However, within the family there is a relatively high
percentage of “misclassification” (22.30 per cent); this could reflect phylogenetic
infidelity or, again, it simply might be attributable to functional similarities
in wing shape.

The Desmodontinae is the only phyllostomatid subfamily that does not exhibit
a “misclassification.” However, three species of Phyllostomus are confused as

289

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290

SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

desmodontines. Of these, two different samples of P. discolor follow this trend
with 91 to 50 per cent affinity, respectively, to the discrimination vector of
vampires. Phyllostomus latifolius exhibits 44 per cent affinity and P. hastatus,
in two separate analyses, showed 100 per cent affinity with this subfamily. A
tentative explanation of this might be that these large-sized phyllostomatines have
flight requirements similar to those of vampires (high weight-bearing capacity)
and hence wings of similar shape.

The glossophagines illustrate the tightest packing of taxa among the phyl-
lostomatids. Only two species, Hylonycteris underwoodi and Platalina gen-
ovensium, are associated outside of the group. It is difficult to assess their re¬
lationship with the stenodermines other than to say that these two species appear
to be similar to Vampyrops and Sturnira.

The stenodermines, although not so tightly packed, occupy a fairly discrete
portion of discriminant space. The seven “misclassified” species are located in the
congested region near the grand centroid for all bats. The Carolliinae is practically
engulfed in this congestion, and they show little fidelity to their group discrimi¬
nation vector. The fact that this congested area exists and that it is composed
primarily of phyllostomatines would suggest the generalized nature of the wings
of this subfamily. Also this seems generally to support the basal assignment of
this group in terms of phylogenetic relationships within the family.

Conclusions

As stated in our introductory comments, the wide range of variation and
complicated nature of the interactions among the intrinsic wing elements of
chiropteran species makes impossible a precise and definitive explanation of
wing shape. However, the essence of wing shape and the variables that affect
it can be perceived in multidimensional space using such multivariate procedures
as discriminant analysis. This study has been as much an analysis of chiropteran
wings as it has been an example of this morphometric procedure.

The interactions among the variables utilized in this study are summarized
below.

1. Length of head and body appears to have little effect on the shape of
chiropteran wings. Generally speaking, bats tend to possess wings that range
between one and one half and two and one-half times the length of the head and
body. Extremes in excess of three times the length of head and body were noted
among the Emballonuridae. Whereas small-sized bats tend to have longer wings
with lighter loading than do larger bats, there is a great deal of variation and the
picture appears to be more complex than simple bivariate analysis indicates.
We do not believe that the questions of mass, area, and wing shape have been
adequately dealt with, and certainly these considerations were beyond the focus
of our analysis. We suggest that these questions will require further analysis under
free-flight conditions.

2. Lengths of forearm and digit III certainly constitute the majority of the
wing span. However, derived variables that describe their relative proportions

BIOLOGY OF THE PHYLLOSTOMATIDAE

291

(such as tip index) do not adequately represent their influence on wing shape. We
have shown that the forearm can be relatively short or relatively long and in
conjunction with the span of digit III produce a wing of similar or different shape.
Pteropodids, emballonuroids, and rhinolophoids tend to emphasize the long
length of forearm in wing construction. The remaining chiropteran families
generally possess shorter forearms.

3. The composite length of digit III can be relatively short (Rhinopomatidae)
or long (Phyllostomatidae and Molossidae). The interactions among the bony
elements that comprise the length of the third digit are extremely complex.
Chiropteran families are ordinated, rather markedly, into two general groups in
discriminant space by the presence or absence of the third phalanx of this digit.
However, wing tips of nearly equal proportional length are achieved by members
of both groups. Those that possess long wing tips, the phyllostomatoids and
vespertilionoids (except molossids) tend to have a lengthened second phalangeal
element of digit III. The phyllostomatoids generally have a lengthened third
phalanx as well and a shortened first phalanx. Vespertilionoids (except molossids)
tend to possess a lengthened first and second phalanx, in an isometric fashion,
and have a shortened terminal phalanx of digit III. Molossids follow the general
pattern of vespertilionoids, but also have a lengthened metacarpal of this digit.
Those bats with generally short wing tips illustrate an allometric mixture in the
composition of digit III. Most, with the notable exceptions of the Pteropodidae and
Craseonycteridae, possess a moderately long metacarpal. However, most of
the span of digit III is contributed by a relatively long second phalanx or, in some
cases, moderately long first and second phalanges of nearly equal length.

4. The effect of the fourth digit on the shape of the wing is complex and, in
most cases, the influence of its elements are involved in an overall synergism among
variables. In those bats with low aspect tip regions, the length of this digit is
intermediate between digits III and V. The fourth digit is relatively long in the high
aspect wing tips of noctilionids and molossids; in those of emballonurids this
digit is shortened. The composition of the fourth digit also varies. In the phyl¬
lostomatoids, the metacarpal is moderately long and has proportionately length¬
ened first and second phalanges. The terminal phalanx is especially long in
noctilionids. The metacarpal element is lengthened in emballonuroids and
rhinolophoids, and the first phalanx also tends to be proportionately long. In
the Pteropodidae, the metacarpal is markedly shortened, and the length of the
digit is produced by proportionately long first and second phalangeal elements.
The long fourth digit of molossids is comprised of the long metacarpal and first
phalanx.

5. Whereas digit III is important in determining the span of the wing, digit V
determines the chord. The interactions between this digit and other wing com¬
ponents in determining shape are somewhat dualistic in nature. The aspect
ratio of the plagiopatagium can be affected either by a lengthening or shortening
of digit V or the forearm. Thus, in the Rhinopomatidae and Emballonuridae,
a relatively long forearm in combination with a moderately long fifth digit produces

292

SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

a high aspect plagiopatagium. On the other hand, in the Molossidae, shortening
of both elements produces an aspect ratio of similar or higher magnitude. The
interaction of digit III with the fifth digit yields a tip region of high aspect in the
Emballonuridae and Molossidae; by comparison, that of rhinopomatids is low
in aspect. In the Phyllostomatidae, a long fifth digit tends to offset the effects
of the long span of digit III and, in combination with a relatively short forearm,
produces an overall low aspect wing. The composition of the fifth digit, like that
of digits III and IV, varies from group to group. Most bats lengthen or shorten
the fifth digit by differentially lengthening or shortening phalangeal elements;
most taxa, especially molossids, retain a moderately long metacarpal. The
pteropodids, as we have noted, have markedly short metacarpal elements in all
three digits. Of the Microchiroptera, the molossids illustrate the most drastic
proportional shortening of the fifth metacarpal.

6. Finally, we reemphasize that although the overall shape of the wing
(silhouette) may be important from the standpoint of such aerodynamic features
as wetted surface area and wing loading, it is the internal composition of the
wing that determines the camberability and ultimately the dynamics of lifting
potential. Far too little is known concerning the comparative aspects of actual
free-flight behavior of bats to permit meaningful functional interpretation of
wing shape. It is to this end that we suggest future morphometric analyses be
directed, for without this, functional speculations can only be misleading and may
further confound an understanding of mammalian flight.

Literature Cited

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Atchley, W. R„ and D. Anderson. 1978. Ratios and the statistical analysis of biological
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Atchley, W. R., C. T. Gaskins, and D. Anderson. 1976. Statistical properties of ratios.
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Bader, R. S., and J. S. Hall. 1960. Osteometric variation and function in bats. Evolu¬
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Cooley, W. W., and P. R. Lohnes. 1971. Multivariate data analysis. John Wiley and
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Daniel, M. J. 1976. Feeding by the short-tailed bat ( Mystacina tuberculata ) on fruit
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Davis, R. 1969. Wing loading in pallid bats. J. Mamm., 50:140-144.

Dixon, W. J. (ed.). 1973. Biomedical computer programs. Univ. California press,

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Dwyer, P. D. 1965. Flight patterns of some eastern Australian bats. Victoria Nat.,
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Eisentraut, M. 1936. Beitrag zur Mechanik des Fledermausfluges. Zeit. wiss. Zool.,
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BIOLOGY OF THE PHYLLOSTOMATIDAE

295

Appendix

Tables A1-A21 follow and consist of ranked means and statistics for selected
derived variables. Statistics include mean for taxa (range in parentheses), plus
or minus one standard error of the mean, and the coefficient of variation. Vari¬
able means are based on genera within families or subfamilies, or species within
genera. Familial means are ranked from largest to smallest. Within the Phyllosto-
matidae, subfamilial means are similarly ranked as are genera within subfamilies.
The grand mean for all bats is ranked with the familial ranking. These tables were
generated aside from the primary principal component and discriminant analyses,
and are discussed in the text to illuminate the interpretation of these multivariate
procedures.

296

SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

Table Al. — Ranked means and statistics for the alpha angle.

Taxon

Mean

Max-min

± 1 SE

Rhinopomatidae

40.25

Rhinolophidae

39.10

(42.55-37.09)

0.663

4.488

Nycteridae

37.75

Furipteridae

37.74

Megadermatidae

37.52

(38.63-37.10)

0.281

1.672

Natalidae

36.74

Pteropodidae

36.45

(39.23-33.61)

0.206

3.094

Craseonycteridae

35.83

Thyropteridae

35.73

Myzapodidae

35.37

Phyllostomatidae

35.32

(39.29-31.73)

0.239

4.740

Phyllonycterinae

37.71

(38.08-37.24)

0.249

1.145

Phyllonycteris

38.08

(38.49-37.68)

0.403

1.498

Erophylla

37.80

(38.16-37.45)

0.352

1.316

Brachyphylla

37.24

(37.37-37.11)

0.131

0.497

Phyllostomatinae

36.70

(39.29-34.69)

0.470

4.247

Tonal ia

39.29

(40.74-36.98)

0.517

3.481

Vampyrum

38.49

Chrotopterus

38.44

Micronycteris

37.31

(39.21-35.40)

0.395

3.670

Macrotus

36.83

(37.74-35.93)

0.904

3.469

Trachops

36.72

Macrophylluin

36.37

(36.68-36.07)

0.308

1.199

Phylloderma

35.32

Minion

35.15

(37.91-33.04)

1.066

6.782

Lonchorhina

35.12

(36.29-34.37)

0.592

2.918

Phyllostomus

34.69

(35.85-33.31)

0.435

2.802

Desmodontinae

36.27

(39.05-33.86)

1.509

7.204

Desniodus

39.05

(39.37-38.72)

0.329

1.190

Diphylla

35.91

Diaemus

33.86

Carolliinae

35.41

(35.91-34.90)

0.505

2.018

Caro Ilia

35.91

(36.31-35.66)

0.143

0.796

Rhinophylla

34.90

(35.95-33.95)

0.578

2.870

Stenoderminae

35.04

(36.58-34.07)

0.165

1.943

Ainetrida

36.58

(37.16-36.01)

0.578

2.234

Phyllops

35.94

(36.18-35.70)

0.241

0.950

Ar ileus

35.75

Cenlurio

35.42

Ardops

35.41

Pvgoderma

35.37

Artibeas

35.12

(35.71-33.99)

0.141

1.444

Sphaeronvcteris

35.08

Vampyrodes

35.03

Eclophylla

35.02

(35.66-34.38)

0.641

2.589

Enchisthenes

34.89

Stnrnira

34.77

(35.22-33.97)

0.129

1.177

Slenoderma

34.42

Uroderina

34.29

(34.53-34.06)

0.231

0.954

Vampyressa

34.29

(35.39-33.81)

0.279

1.822

Vampyrops

34.16

(34.70-33.46)

0.145

1.346

Chiroderma

34.07

(34.66-33.49)

0.234

1.538

Glossophaginae

33.73

(35.41-31.73)

0.290

3.101

Glossophaga

35.41

(35.63-35.20)

0.089

0.504

Platalina

35.34

Leptonyderis

34.93

(35.43-34.22)

0.365

1.811

Lonchophvlla

34.47

(35.18-33.74)

0.246

1.598

Choeronycteris

33.62

Lionycteris

33.48

Choeroniscus

33.46

(34.25-32.43)

0.331

2.209

Lichonycteris

33.45

(33.85-33.00)

0.246

1.273

Hvlonycteris

33.39

Musonycteris

33.22

Scleronycteris

33.00

Monophylltis

32.97

(33.38-32.57)

0.407

1.747

Anoura

31.73

(32.57-30.79)

0.320

2.255

Mormoopidae

35.17

(37.13-33.21)

1.963

7.894

All bats

153

35.08

(42.55-24.09)

0.237

8.373

Vespertilionidae

35.02

(38.84-31 .41)

0.391

6.220

Mystacinidae

34.78

Emballonuridae

32.95

(35.66-29.07)

0.507

5.330

Noctilionidae

30.38

Molossidae

26.93

(29.82-24.09)

0.516

5.754

BIOLOGY OF THE PHYLLOSTOMATIDAE

297

Table A2. — Ranked means and statistics for the tip index.

Taxon

Mean

Max-min

± 1 SE

Phyllostomatidae

2.04

(2.35-1.59)

0.025

8.546

Carolliinae

2.24

(2.30-2.18)

0.061

3.831

Rhinophylla

2.30

(2.50-2.20)

0.102

7.674

Carol lia

2.18

(2.22-2.15)

0.016

1.457

Stenoderminae

2.15

(2.35-2.04)

0.020

3.902

Pygoderma

2.35

Anieirida

2.27

(2.28-2.25)

0.018

1.1 14

Vampyrops

2.22

(2.34-2.07)

0.024

3.464

Sphaeronycteris

2.22

Ariteus

2.21

Vampyressa

2.20

(2.33-2.08)

0.046

4.699

Chirodernta

2.19

(2.24-2.10)

0.026

2.657

Sturniru

2.17

(2.29-2.10)

0.020

2.898

Stenoderma

2.15

Vampyrodes

2.14

Ardops

2.11

Ectophylla

Uroaerma

2.10

(2.14-2.07)

0.036

2.446

2.09

(2.09-2.08)

0.004

0.286

Phyllops

2.08

(2.11-2.05)

0.030

2.015

Centurio

2.05

Enchisthenes

2.05

Artibeus

2.04

(2.13-1.91)

0.020

3.550

Glossophaginae

2.06

(2.20-1.81)

0.033

5.776

Scleronycteris

2.20

A no lira

2.20

(2.29-2.12)

0.035

3.574

Lichonycteris

2.19

(2.28-2.09)

0.055

4.329

Hylonycteris

2.19

Choeroniscus

2.11

(2.21-2.03)

0.035

3.740

Lonchophylla

2.07

(2.19-1.92)

0.044

4.799

Choeronycteris

2.05

Lionycteris

2.02

Glossophagu

2.00

(2.03-1.95)

0.020

2.021

Monophyllus

1.99

(2.00-1.99)

0.004

0.252

Musonycteris

1.96

Platalina

1.96

Leptonycteris

1.81

(1.85-1.76)

0.024

2.328

Phyllostomatinae

1.92

(2.11-1.68)

0.033

5.696

Mucrophyllum

2.11

(2.13-2.10)

0.017

1.158

Phvlloderma

2.03

Million

1.98

(2.10-1.84)

0.045

5.076

Trachops

1.96

Lonchorhina

1.95

(2.02-1.89)

0.036

3.190

Vampyrum

1.94

Chrotopterus

1.90

(2.16-1.68)

0.038

7.029

Micronycteris

1.89

Tonal ia

1.87

(1.95-1.80)

0.018

2.495

Phyllostomus

1.85

(1.97-1.75)

0.038

4.548

Macrotus

1.68

(1.70-1.66)

0.020

1.643

Desmodontinae

1.86

(2.03-1.59)

0.141

13.083

Diphylla

2.03

Diaemus

1.98

Desmodus

1.59

(1.60-1.57)

0.015

1.348

Phyllonycterinae

1.69

(1.74-1.66)

0.025

2.613

Brachyphylla

1.74

(1.74-1.74)

0.000

0.025

Erophylla

1.66

(1.69-1.64)

0.023

1.955

Phyllonycteris

1.66

(1.68-1.65)

0.016

1.321

Molossidae

1.98

(2.30-1.81)

0.044

6.669

Noctilionidae

1.95

Craseonycteridae

1.86

Natalidae

1.85

All bats

153

1.85

(2.35-1.09)

0.018

12.280

Vespertilionidae

1.81

(2.17-1.60)

0.027

8.396

Thyropteridae

1.81

Myzapodidae

1.79

6.596

Pteropodidae

1.78

(2.06-1.57)

0.021

Nycteridae

1.78

0.017

2.101

Megadermatidae

1.76

(1.83-1.74)

Mystacinidae

1.75

0.123

10.221

Mormoopidae

1.70

(1.82-1.58)

Emballonuridae

1.62

(1.92-1.48)

0.032

6.950

Furipteridae

1.58

0.036

6.764

Rhinolophidae

1.40

(1.52-1.29)

Rhinopomatidae

1.09

298

SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

Table A3. — Ranked means and statistics for the overall aspect ratio.

Taxon

Mean

Max-min

± 1 SE

Molossidae

7.54

(8.05-6.46)

0.174

6.935

Emballonuridae

6.71

(7.93-6.05)

0.147

7.620

Noctilionidae

6.69

Mormoopidae

6.04

(6.39-5.68)

0.356

8.346

All bats

153

5.86

(8.05-4.71)

0.051

10.866

Phyllostomatidae

5.74

(6.50-5.05)

0.046

5.615

Glossophaginae

6.09

(6.50-5.71)

0.061

3.623

Anoura

6.50

(6.74-6.23)

0.087

2.991

Musonycteris

6.30

Scleronycteris

6.23

Lionycteris

6.19

Monophyllus

6.18

(6.29-6.07)

0.109

2.482

Choeroniscus

6.17

(6.40-6.01)

0.073

2.653

Lichonycteris

6.13

(6.30-5.98)

0.091

2.563

Hvlonycteris

6.09

Choeronycteris

6.09

Lonchophylla

5.94

(6.10-5.81)

0.056

2.102

Leplonycteris

5.92

(6.09-5.82)

0.083

2.425

Platalina

5.72

Glossophaga

5.71

(5.80-5.64)

0.033

1.149

Stenoderminae

5.71

(5.96-5.26)

0.050

3.632

Chiroderma

5.96

(6.14-5.78)

0.060

2.240

Uroderma

5.94

(5.97-5.91)

0.027

0.648

Vampyrops

5.93

(6.11-5.78)

0.030

1.606

Stenoderma

5.88

Vampyressa

5.88

(5.96-5.69)

0.049

1.869

Centurio

5.88

Ectophvllu

5.76

(5.92-5.61)

0.151

3.713

Sturnira

5.76

(5.88-5.62)

0.026

1.404

Artibeus

5.75

(6.04-5.57)

0.036

2.231

Vampyrodes

5.71

Ardops

5.66

Pvgoaenna

5.63

Enchisthenes

5.62

Phyllops

5.58

(5.64-5.53)

0.054

1.365

Ariteus

5.49

Sphaeronycteris

Ametrida

5.32

5.26

(5.33-5.19)

0.070

1.874

Carolliinae

5.69

(5.74-5.64)

0.049

1.205

Rhinophylla

5.74

(5.95-5.52)

0.124

3.748

Carollia

5.64

(5.74-5.54)

0.041

1.442

Phyllostomatinae

5.55

(5.92-5.05)

0.094

5.620

Minion

5.92

(6.36-5.41)

0.209

7.892

Phyllostomus

5.91

(6.18-5.54)

0.109

4.107

Lonchorhina

5.89

(6.04-5.60)

0.140

4.135

Phvlloderina

5.78

Macrotus

5.77

(6.06-5.49)

0.284

6.945

Macrophyllum

5.45

(5.48-5.43)

0.029

0.747

Trachops

5.42

Micronycteris

5.41

(5.84-5.02)

0.076

4.867

Chrotopierus

5.25

Vampyrum

5.23

Tonalia

5.05

(5.57-4.74)

0.113

5.913

Desmodontinae

5.50

(5.81-5.17)

0.186

5.874

D iaem us

5.81

D ip h vita

5.51

Desnwdus

5.17

(5.23-5.11)

0.057

1.572

Phyllonycterinae

5.40

(5.44-5.35)

0.027

0.872

Brachvphvlla

5.44

(5.46-5.42)

0.016

0.428

Erophvlla

5.42

(5.52-5.32)

0.098

2.559

Phyllonvcteris

5.35

(5.43-5.27)

0.082

2.169

Vespertilionidae

5.73

(6.90-4.92)

0.085

8.229

Mystacinidae

5.71

Thyropteridae

5.70

Myzapodidae

5.65

Craseonycteridae

5.64

Natalidae

5.60

Rhinopomatidae

Megadermatidae

5.58

5.55

(5.74-5.29)

0.073

2.938

Furipteridae

5.52

Pteropodidae

5.49

(5.97-5.01)

0.035

3.533

Nycteridae

5.48

Rhinolophidae

5.42

(5.99-4.71)

0.157

7.645

BIOLOGY OF THE PHYLLOSTOMATIDAE

299

Table A4. — Ranked means and statistics for the aspect ratio of the wing tip.

Taxon

Mean

Max-min

± 1 SE

Molossidae

5.79

(6.31-4.79)

0.163

8.465

Embalonuridae

5.22

(6.35-4.77)

0.133

8.790

Noctilionidae

5.21

Mormoopidae

4.73

(5.12-4.34)

0.390

11.655

Phyllostomatidae

4.67

(5.41-3.86)

0.048

7.155

Glossophaginae

4.98

(5.41-4.60)

0.066

4.804

Anoura

5.41

(5.68-5.10)

0.105

4.328

Scleronycteris

5.21

Musonyeteris

5.13

Lichanycteris

5.13

(5.32-5.03)

0.096

3.252

Choeroniscus

5.11

(5.29-4.91)

0.076

3.320

Lionycteris

5.06

Hvlonvcteris

5.06

Choeronycteris

4.97

Monophvllus

4.95

(5.04-4.85)

0.097

2.779

Lonchophylla

4.89

(5.11-4.67)

0.088

4.006

Leptonvcteris

4.67

(4.81-4.59)

0.072

2.673

Glossophaga

4.62

(4.67-4.58)

0.025

1.094

Platalina

4.60

Carolliinae

4.78

(4.83-4.74)

0.043

1.279

Rhinophylla

4.83

(5.13-4.58)

0.163

5.865

Carollia

4.74

(4.85-4.62)

0.052

2.215

Stenoderminae

4.70

(4.96-4.22)

0.053

4.628

Vampyrops

4.96

(5.15-4.84)

0.034

2.146

Chiroderma

4.96

(5.16-4.71)

0.073

3.313

Centurio

4.93

Uroderma

4.88

(4.89-4.87)

0.009

0.269

Vampyressa

4.88

(4.99-4.70)

0.057

2.615

Stenoderma

4.87

Pvyoderma

4.76

Sturnira

4.74

(4.88-4.66)

0.020

1.301

Ectophylla

4.73

(4.84-4.63)

0.104

3.107

Vampyrodes

4.70

Artibeus

4.67

(4.87-4.48)

0.037

2.883

Ardops

4.65

Phyllops

4.57

(4.65-4.50)

0.071

2.194

Ariteus

4.52

Enchisthenes

4.47

Ametrida

4.33

(4.37-4.28)

0.045

1.484

Sphaeronvcteris

4.22

Phyllostomatinae

4.48

(4.86-4.04)

0.079

5.878

M ini on

4.86

(5.26-4.38)

0.193

8.892

Lonchorhina

4.79

(4.90-4.60)

0.094

3.413

Phylloderma

4.74

Phyllostomus

4.66

(4.83-4.32)

0.098

4.710

Macrotus

4.59

(4.87-4.32)

0.275

8.475

Macrophyllum

4.47

(4.49-4.45)

0.017

0.535

Trachops

4.33

Micronycteris

4.32

(4.73-3.90)

0.076

6.058

Vampyrum

4.27

Chrotopterus

4.25

Tonatia

4.04

(4.60-3.69)

0.120

7.839

Desmodontinae

4.28

(4.54-3.86)

0.209

8.457

Diaemus

4.54

D ip h vlla

4.42

Desmodus

3.86

(3.88-3.84)

0.019

0.692

Phyllonycterinae

4.16

(4.21-4.10)

0.032

1.320

Brachyphylla

4.21

(4.22-4.20)

0.008

0.254

Erophylla

4.17

(4.24-4.10)

0.069

2.356

Phyllonycteris

4.10

(4.19-4.01)

0.087

2.999

All bats

153

4.58

(6.35-3.44)

0.044

12.004

Natalidae

4.51

Thyropteridae

4.46

Craseonycteridae

4.45

Megadermatidae

4.45

(4.72-4.19)

0.089

4.463

Nycteridae

4.39

Vespertilionidae

4.38

(5.99-3.72)

0.083

10.512

Myzapodidae

4.33

Mystacinidae

4.27

Pteropodidae

4.22

(4.48-3.65)

0.029

3.782

Furipteridae

4.19

Rhinolophidae

3.93

(4.27-3.44)

0.099

6.667

Rhinopomatidae

3.69

300

SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

Table A5. — Ranked means and statistics for the aspect ratio of the plagiopatagium.

Taxon

Mean

Max-min

± 1 SE

Rhinopomatidae

2.17

Molossidae

2.00

(2.21-1.82)

0.038

5.762

Emballonuridae

1.93

(2.20-1.66)

0.054

9.630

Rhinolophidae

1.78

(2.06-1.51)

0.077

1 1 .440

Noctilionidae

1.75

Mormoopidae

1.68

(1.68-1.68)

0.002

0.158

Mystacinidae

1.65

Furipteridae

1.64

Vespertilionidae

1.59

(1.99-1.32)

0.028

9.742

All bats

153

1.58

(2.21-1.20)

0.018

14.364

Myzapodidae

1.57

Thyropteridae

1.54

Pteropodidae

1.52

(1.74-1.34)

0.018

6.619

Craseonycteridae

1.49

Megadermatidae

1.48

(1.52-1.43)

0.018

2.688

Nycteridae

1.46

Natalidae

1.45

Phyllostomatidae

1.40

(1.59-1.19)

0.015

7.265

Phyllonycterinae

1.53

(1.55-1.51)

0.011

1.215

Erophylla

1.55

(1.59-1.51)

0.040

3.670

Phyllonycteris

1.54

(1.54-1.53)

0.007

0.673

Brachyphylla

1.51

(1.52-1.51)

0.006

0.583

Desmodontinae

1.47

(1.56-1.36)

0.060

7.004

Desmudus

1.56

(1.59-1.52)

0.034

3.082

Diaem us

1.51

Diphylla

1.36

Glossophaginae

1.46

(1.58-1.38)

0.019

4.670

Leptonycteris

1.58

(1.62-1.54)

0.023

2.530

Musonycteris

1.55

Monophyllus

1.55

(1.57-1.53)

0.022

2.013

Lionycteris

1.50

Ano lira

1.47

(1.51-1.44)

0.010

1.538

Choeronycteris

1.46

Platalina

1.44

Choeroniscus

1.44

(1.50-1.37)

0.021

3.253

Lonchophylla

1.41

(1.52-1.36)

0.028

4.507

Glossophcigu

1.41

(1.46-1.37)

0.019

2.700

Scleronycteris

1.40

Hvlonycteris

1.38

Lichonycteris

1.38

(1.45-1.31)

0.040

4.986

Phyllostomatinae

1.40

(1.59-1.29)

0.031

7.392

Macrotus

1.59

(1.66-1.52)

0.071

6.292

Phyllostomus

1.57

(1.67-1.44)

0.048

6.789

Lonchorhina

1.46

(1.53-1.35)

0.055

6.526

Mimon

1.44

(1.51-1.35)

0.032

5.007

Micronycteris

1.39

(1.51-1.24)

0.025

6.144

Phylloderma

1.39

Trachops

1.37

Chrotopterus

1.33

Tonatia

1.31

(1.36-1.26)

0.015

2.985

Vanipyrum

1.30

Macrophyllum

1.29

(1.29-1.28)

0.003

0.380

Stenoderminae

1.33

(1.41-1.19)

0.015

4.689

Uroderma

1.41

(1.42-1.40)

0.009

0.949

Enchisthenes

1.40

Artibeus

1.40

(1.50-1.32)

0.016

4.023

Centurio

1.37

Ectophylla

1.36

(1.41-1.30)

0.056

5.803

Stenoderina

1.36

Chiroderma

1.35

(1.38-1.33)

0.009

1.414

Vumpvressa

1.34

(1.42-1.27)

0.026

4.300

Vampyrodes

1.33

Stnrnira

1.33

(1.38-1.25)

0.013

2.999

Ardops

1.33

Phyllops

1.33

(1.34-1.32)

0.007

0.762

Vampyrops

1.33

(1.40-1.26)

0.012

2.823

Sphaeronvcteris

1.28

Ariteus

1.26

Pygoderma

1.20

Ameirida

1.19

(1.21-1.17)

0.015

1.818

Carolliinae

1.26

(1.27-1.25)

0.009

1.006

Carollia

1.27

(1.28-1.25)

0.006

0.870

Rhinophylla

1.25

(1.31-1.19)

0.036

4.936

BIOLOGY OF THE PHYLLOSTOMATIDAE

301

Table A6. — Ranked means and statistics for wing loading in newtons per square meter.

Taxon

Mean

Max-min

±1 SE

Molossidae

21.41

(28.47-15.56)

1.239

17.358

Pteropodidae

19.18

(36.24-11.48)

1.084

30.959

Noctilionidae

17.65

Craseonycteridae

16.70

All bats

153

14.62

(36.08- 3.69)

0.507

42.905

Phyllostomatidae

Desmodontinae

14.50

(28.89- 3.92)

0.686

33.1 19

20.87

(29.23-14.99)

4.293

35.639

Diaemus

32.71

Diphylla

20.34

Desmodus

15.72

(17.26-14.17)

1.548

13.931

Phyllonycterinae

18.40

(21.04-13.75)

2.331

21.940

Phyllonycteris

21.04

(24.56-17.52)

3.522

23.675

Brcichvphvllu

20.41

(22.74-18.08)

2.329

16.134

Erophylla

Stenoaerminae

13.75

(15.46-12.05)

1.708

17.557

15.01

(22.66-10.96)

0.776

21.319

Enchisthenes

22.66

Sturniru

17.85

(28.58-10.68)

2.071

36.690

Ariteus

17.42

Vampyrodes

17.10

Chiroaerma

17.09

(21.86-13.15)

1.424

18.632

Sphaeronycteris

Centurio

16.96

16.76

Vampyrops

16.09

(21.13-10.38)

0.895

17.576

Artibeus

15.94

(23.23-10.56)

1.084

24.515

Uroderma

14.04

(16.59-1 1 .49)

2.549

25.675

Stenodenna

13.39

Ametridu

12.77

(13.85-11.69)

1.079

11.954

Ardops

11.98

Vampyressa

11.50

(13.01- 8.90)

0.840

16.327

Ectophylla

11.47

(12.73-10.22)

1.256

15.485

Phyllops

11.16

(13.29- 9.03)

2.128

26.968

Pygodernia

10.96

Phyilostomatinae

14.04

(19.94- 7.88)

1.205

28.473

Phyllostomus

19.94

(24.11-16.47)

1.242

13.927

Chrotopterus

18.81

Trachons

Phylloaernia

17.36

16.30

Vampyruni

15.48

Tonal ia

14.64

(19.35-10.86)

1.206

21.797

Macrotus

12.96

(14.68-11.25)

1.715

18.708

Mimon

11.23

(13.94- 6.81)

1.295

25.786

Lonchorhina

11.05

(13.22- 9.56)

1.110

17.392

Micronycteris

8.77

(15.43- 5.47)

0.809

31.949

Mucrophyllum

7.88

(10.24- 5.52)

2.361

42.356

Glossophaginae

12.51

(15.58-10.01)

0.421

12.128

Musonycteris

15.58

Leptonycteris

14.17

(15.97-11.77)

1.249

15.270

Choeronycteris

13.75

Hylonycteris

13.50

Lonchophyllu

13.21

(17.03-11.46)

1.005

17.015

Glossophaga

12.54

(14.81-11.32)

0.774

12.343

Anoura

12.31

(17.35- 9.36)

1.415

25.706

Choeroniscus

12.16

(14.01-11.25)

0.510

9.390

Menophyllus

11.93

(12.45-11.41)

0.519

6.151

Lichonycteris

11.63

(12.03-11.27)

0.220

3.276

Lionycteris

10.94

Platalinu

10.94

Scleronycteris

10.01

Carolliinae

10.98

(11.14-10.81)

0.168

2.168

Rhinophylla

11.14

(12.46- 9.70)

0.800

12.428

Caro Ilia

10.81

(12.87- 9.10)

0.857

15.863

Rhinopomatidae

Megadermatidae

11.82

11.33

(15.16- 8.32)

1.428

25.220

Mystacinidae

11.15

1.825

24.406

Mormoopidae

10.57

(12.40- 8.75)

Vespertilionidae

10.54

(19.71- 6.92)

0.544

28.713

Nycteridae

9.82

1.364

48.847

Emballonuridae

9.67

(21.16- 4.73)

Rhinolophidae

8.05

(14.48- 1.84)

1.769

58.147

Myzapodidae

7.41

Thyropteridae

5.91

Natalidae

5.43

Furipteridae

4.20

302

SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

Table A7. — Ranked means and statistics for the relative length of the wing.

Taxon

Mean

Max-min

± 1 SE

Furipteridae

2.62

Natalidae

2.61

Noctilionidae

2.53

Emballonuridae

2.44

(3.34-2.06)

0.110

15.553

Myzapodidae

2.43

Thyropteridae

2.42

Craseonycteridae

2.35

Nycteridae

2.31

Mormoopidae

2.21

(2.33-2.09)

0.118

7.531

Rhinolophidae

2.20

(2.51-2.03)

0.064

7.644

Megadermatidae

2.18

(2.43-1.95)

0.084

8.643

Vespertilionidae

2.07

(2.49-1.44)

0.048

12.907

Phyllostomatidae

2.07

(2.40-1.69)

0.024

8.174

Carolliinae

2.22

(2.24-2.20)

0.018

1.162

Caro Ilia

2.24

(2.46-2.04)

0.090

8.029

Rhinophylla

2.20

(2.39-2.03)

0.105

8.251

Phyllostomatinae

2.18

(2.40-2.00)

0.034

5.103

Minion

2.40

(2.75-2.22)

0.096

8.969

Lonchorhina

2.31

(2.49-2.18)

0.094

7.087

Macrophvlhwi

2.25

(2.36-2.14)

0.106

6.656

Micronycteris

2.22

(2.56-1.82)

0.058

9.078

Phylloderma

2.19

Macrotus

2.18

(2.29-2.07)

0.111

7.204

Trachops

2.13

Vampyrum

2.12

Chrotopterus

2.11

Phyllostomus

2.08

(2.18-1.99)

0.036

3.845

Tonal ia

2.00

(2.22-1.60)

0.081

10.764

Stenoderminae

2.14

(2.36-1.89)

0.035

6.840

Ardops

2.36

Stenodenna

2.34

Phyllops

2.27

(2.49-2.06)

0.212

13.188

Centurio

2.24

Vampyrodes

2.22

Pygoderma

2.21

Vampyrops

2.20

(2.46-1.99)

0.042

5.999

Artibeus

2.19

(2.44-2.00)

0.039

6.343

Ectophylla

2.19

(2.34-2.03)

0.150

9.731

Chiroderma

2.16

(2.23-2.10)

0.024

2.528

Vampyressa

2.15

(2.41-2.03)

0.074

7.713

Sturnira

2.05

(2.22-1.83)

0.043

6.606

Uroderina

2.03

(2.06-2.00)

0.029

2.034

Ariteus

1.96

Ametrida

1.96

(2.04-1.87)

0.083

5.968

Enchisthenes

1.90

Sphaeronycteris

1.89

Glossophaginae

1.93

(2.16-1.75)

0.033

6.112

Scleronycteris

2.16

Lionycteris

2.10

Choeronycteris

2.01

Anoura

1.98

(2.26-1.75)

0.102

11.532

Lichonycteris

1.98

(2.03-1.93)

0.029

2.571

Monophyllus

1.96

(2.04-1.87)

0.085

6.174

Choeroniscus

1.92

(2.00-1.83)

0.029

3.312

Platalina

1.91

Glossophaga

1.87

(1.91-1.84)

0.016

1.683

Leptonycteris

1.84

(1.97-1.75)

0.068

6.354

Lonchophylla

1.84

(1.90-1.79)

0.021

2.516

Hylonycteris

1.79

Musonycteris

1.75

Desmodontinae

1.93

(2.05-1.69)

0.118

10.617

Desmodus

2.05

(2.06-2.05)

0.005

0.314

D ip h vlla

2.05

Diaemus

1.69

Phyllonycterinae

1.92

(2.05-1.78)

0.077

6.951

Erophylla

2.05

(2.14-1.96)

0.093

6.384

Brachyphylla

1.94

(2.00-1.87)

0.068

4.933

Phy lionycteris

1.78

(1.89-1.68)

0.106

8.400

All bats

153

2.06

(3.34-1.44)

0.023

13.586

Mystacinidae

1.97

Rhinopomatidae

1.97

Molossidae

1.86

(2.10-1.63)

0.053

8.476

Pteropodidae

1.80

(2.13-1.59)

0.027

8.108

BIOLOGY OF THE PHYLLOSTOMATIDAE

303

Table A8. — Ranked means and statistics for the relative length of the forearm.

Taxon

Mean

Max-min

± 1 SE

Furipteridae

1.02

Rhinopomatidae

0.94

Emballonuridae

0.93

(1.14-0.76)

0.034

12.771

Rhinolophidae

0.92

(1.01-0.86)

0.022

6.479

Natalidae

0.92

Myzapodidae

0.87

Thyropteridae

0.86

Noctilionidae

0.86

Nycteridae

0.83

Craseonyctridae

0.82

Mormoopidae

0.82

(0.82-0.81)

0.007

1.130

Megadermatidae

0.79

(0.88-0.71)

0.028

7.967

Vespertilionidae

0.74

(0.92-0.56)

0.014

10.850

All bats

153

0.73

(1.14-0.52)

0.010

16.238

Mystacinidae

0.72

Phyllostomatidae

0.68

(0.81-0.56)

0.009

9.145

Phyllostomatinae

0.75

(0.81-0.69)

0.012

5.232

Macrotus

0.81

(0.85-0.78)

0.035

6.162

Mimon

0.80

(0.91-0.71)

0.034

9.502

Lonchorhinu

0.78

(0.85-0.74)

0.032

7.171

Micronycteris

0.77

(0.90-0.64)

0.021

9.585

Phyllostomus

0.73

(0.77-0.70)

0.013

3.857

Chrotopterus

0.73

Macrophyllum

0.72

(0.75-0.69)

0.030

5.940

Phylloaerma

0.72

Vampyrum

Trachops

0.72

Tonatia

0.69

(0.77-0.57)

0.026

9.746

Phyllonycterinae

0.72

(0.77-0.67)

0.029

6.996

Erophylla

0.77

(0.81-0.73)

0.042

7.640

Brachyphytla

0.71

(0.73-0.68)

0.025

4.913

Phyllonycteris

0.67

(0.71-0.63)

0.044

9.255

Caroiliinae

0.69

(0.70-0.67)

0.018

3.807

Carollia

0.70

(0.77-0.65)

0.026

7.338

Rhinophylla

0.67

(0.69-0.63)

0.018

4.555

Desmodontinae

0.68

(0.79-0.57)

0.065

16.603

Desmodus

0.79

(0.80-0.79)

0.003

0.523

Diphylla

0.67

Diaemus

0.57

Stenoderminae

0.68

(0.76-0.59)

0.013

7.756

Ardops

0.76

Sienoderma

0.74

Phyllops

0.74

(0.80-0.68)

0.062

11.844

Centurio

0.73

Artibeus

0.72

(0.80-0.66)

0.011

5.679

Vampyrodes

0.71

Ectophylla

0.70

(0.76-0.65)

0.057

11.414

Vumpyrops

0.68

(0.76-0.61)

0.013

5.890

Chiroderma

0.68

(0.69-0.66)

0.006

2.041

Vumpyressa

0.67

(0.74-0.64)

0.017

5.645

Pygoderma

0.66

Uruderma

0.66

(0.67-0.65)

0.010

2.229

Sturnira

0.65

(0.70-0.56)

0.014

6.773

Enchisihenes

0.62

Ariteus

0.61

Ainetrida

0.60

(0.63-0.57)

0.028

6.712

Sphaeronycteris

0.59

Glossophaginae

0.63

(0.70-0.56)

0.010

5.819

Lionycteris

0.70

Scleronycteris

0.68

Choerortycteris

0.66

Leptonycteris

0.66

(0.71-0.62)

0.030

7.848

Monophyllus

0.65

(0.68-0.63)

0.028

6.059

Platalinci

0.65

Glossophaga

0.62

(0.65-0.61)

0.008

2.675

Lichortycteris

0.62

(0.64-0.59)

0.016

4.520

Choeroniscus

0.62

(0.65-0.59)

0.010

3.751

Anuura

0.62

(0.69-0.55)

0.027

9.645

Lonchophylla

0.60

(0.63-0.56)

0.013

4.816

Musonycteris

0.59

Hvlonycteris

0.56

Pteropodidae

0.65

(0.80-0.52)

0.012

9.730

Molossidae

0.63

(0.72-0.56)

0.018

8.610

304

SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

Table A9. — Ranked means and statistics for the relative length of digit III.

Taxon

Mean

Max-min

± 1 SE

Natalidae

1.69

Noctilionidae

1.67

Furipteridae

1.61

Myzapodidae

1.56

Thyropteridae

1.56

Craseonycteridae

1.53

Emballonuridae

1.51

(2.19-1.28)

0.077

17.721

Nycteridae

1.48

Mormoopidae

1.39

(1.50-1.28)

0.111

1 1.291

Megadermatidae

1.39

(1.55-1.24)

0.057

9.091

Phyllostomatidae

1.39

(1.60-1.11)

0.018

9.206

Carolliinae

1.54

(1.54-1.54)

0.000

0.018

Rhinophvlla

1.54

(1.71-1.39)

0.092

10.325

Caro Ilia

1.54

(1.69-1.40)

0.064

8.370

Stenoderminae

1.46

(1.60-1.28)

0.024

6.728

Ardops

1.60

Stenoderma

1.59

Pvgodenna

1.55

Phyllops

1.54

(1.69-1.39)

0.150

13.834

Vampyrops

1.51

(1.70-1.36)

0.030

6.345

Vainpyrodes

1.51

Cenlurio

1.51

Chirodenna

1.48

(1.54-1.42)

0.021

3.098

Ectophvlla

1.48

(1.57-1.39)

0.094

8.931

Vumpyressa

1.48

(1.68-1.38)

0.059

8.884

Anioeus

1.47

(1.66-1.31)

0.028

6.936

Sturnira

1.40

(1.54-1.27)

0.030

6.720

Uroderma

1.37

(1.39-1.35)

0.019

1.941

Ametrida

1.36

(1.41-1.30)

0.054

5.639

Arireus

1.35

Sphaeronvcteris

1.30

Enchisthenes

1.28

Phyllostomatinae

1.43

(1.59-1.30)

0.026

6.095

Mimon

1.59

(1.84-1.48)

0.065

9.137

Macrophvlluni

1.53

(1.60-1.45)

0.076

6.995

Lonchorhina

1.52

(1.65-1.43)

0.064

7.265

Phvlloderma

1.47

Micronycteris

1.45

(1.66-1.18)

0.040

9.658

Trachops

1.41

Vampvrum

1.40

Chrotopterus

1.38

Macrotus

1.37

(1.44-1.29)

0.076

7.823

Phyllostomus

1.35

(1.41-1.27)

0.028

4.658

Tonal ia

1.30

(1.46-1.03)

0.056

11.356

Glossophaginae

1.30

(1.49-1.16)

0.026

7.090

Scleronycteris

1.49

Lionycteris

1.41

Anoura

1.36

(1.57-1.20)

0.076

12.464

Lichonycteris

1.36

(1.40-1.34)

0.020

2.487

Choeronycteris

1.35

Choeroniscus

1.30

(1.38-1.24)

0.022

3.759

Monophyllus

1.30

(1.36-1.25)

0.057

6.231

Platalina

1.27

Glossophaga

1.25

(1.27-1.23)

0.009

1.423

Lonchophylla

1.24

(1.28-1.21)

0.013

2.432

Hvlonycteris

1.23

Leptonycteris

1.19

(1.26-1.14)

0.038

5.529

Musonycteris

1.16

Desmodontinae

1.25

(1.37-1.13)

0.071

9.809

Diphylla

1.37

Desmodus

1.26

(1.27-1.25)

0.007

0.841

Diaemus

1.13

Phyllonycterinae

1.21

(1.28-1.11)

0.050

7.107

Erophylla

1.28

(1.33-1.23

0.051

5.629

Brachyphvlla

1.23

(1.27-1.19)

0.043

4.945

Phyllonycteris

1.11

(1.17-1.05)

0.062

7.885

Vespertilionidae

1.34

(1.69-0.89)

0.035

14.739

All bats

153

1.33

(2.19-0.89)

0.015

14.209

Rhinolophidae

1.28

(1.52-1.15)

0.046

9.478

Mystacinidae

1.26

Molossidae

1.24

(1.41-1.07)

0.038

9.294

Pteropodidae

1.15

(1.33-0.99)

0.017

8.043

Rhinopomatidae

1.03

BIOLOGY OF THE PHYLLOSTOMATIDAE

305

Table A10. — Ranked means and statistics for the relative length of digit IV.

Taxon

Mean

Max-min

± 1 SE

Noctilionidae

1.27

Natalidae

1.25

Furipteridae

1.25

Myzapodidae

1.25

Thyropteridae

1.20

Craseonycteridae

1.18

Nycteridae

1.10

Emballonuridae

1.07

(1.43-0.85)

0.049

15.900

Vespertilionidae

1.07

(1.27-0.74)

0.022

11.381

Rhinolophidae

1.04

(1.20-0.89)

0.038

9.718

Mystacinidae

1.03

Mormoopidae

1.03

(1.07-0.99)

0.044

5.998

Phyllostomatidae

1.03

(1.19-0.82)

0.012

8.208

Carolliinae

1.11

(1.11-1.10)

0.004

0.516

Rhinophvlla

1.11

(1.19-1.04)

0.045

6.930

Carollia

1.10

(1.19-1.01)

0.038

6.933

Stenoderminae

1.08

(1.19-1.00)

0.013

4.954

Ardops

1.19

Stenoderma

1.16

Phvl lops

1.14

(1.24-1.04)

0.101

12.438

Pvgoderma

1.13

Vampyrodes

1.12

Artibeus

1.10

(1.22-1.00)

0.016

5.277

Ectophvlla

1.09

(1.15-1.03)

0.062

8.082

Vampyrops

1.09

(1.19-0.98)

0.019

5.659

Vampvressa

1.08

(1.21-1.02)

0.037

7.644

Sphaeronvcteris

1.07

Chiroderma

1.07

(1.09-1.05)

0.008

1.605

Centurio

1.06

Ametrida

1.05

(1.09-1.01)

0.039

5.234

Sturnira

1.04

(1.16-0.93)

0.023

6.938

Ariteus

1.02

Uroderma

1.00

(1.01-0.99)

0.010

1.403

Enchisthenes

1.00

Phyllostomatinae

1 1

1.07

(1.15-1.00)

0.015

4.726

Macrophtlhnn

1.15

(1.21-1.09)

0.061

7.502

Minton

1.14

(1.30-1.05)

0.046

9.065

Lonchorhina

1.11

(1.20-1.02)

0.050

7.846

Micronycteris

1.11

(1.23-0.92)

0.026

8.110

Trachops

Phvlloaerma

1.09

1.07

Vampyrunt

1.05

Chrotopterus

1.05

0.035

9.170

Tonatia

1.02

(1.16-0.86)

Phyllostomus

1.02

( 1 .10-0.97)

0.028

6.068

Macrotus

1.00

(1.09-0.90)

0.094

13.335

Desmodontinae

0.99

(1.06-0.89)

0.052

9.075

Diphylla

1.06

Desntodus

1.04

(1.04-1.03)

0.004

0.520

Diaemus

0.89

Phyllonycterinae

0.95

(1.00-0.88)

0.036

6.533

Erophylla

1.00

(1.03-0.97)

0.031

4.418

Brachyphylla

0.97

(1.00-0.93)

0.035

5.093

Phvllonycteris

0.88

(0.94-0.82)

0.060

9.631

Glossophaginae

0.94

(1.05-0.82)

0.016

6.247

Scleronveteris

1.05

Lionycteris

1.01

Choeronvcreris

0.98

Monophvlhts

0.97

(1.00-0.93)

0.034

5.003

Lichonycteris

0.96

(0.97-0.96)

0.003

0.515

A no lira

0.96

(1.08-0.83)

0.047

10.987

Plcilalina

0.95

Glossophaga

0.93

(0.94-0.92)

0.007

1.420

Choeroniscus

0.93

(0.98-0.88)

0.017

4.156

Lonchophvlla

0.90

(0.93-0.86)

0.014

3.618

Leptonycteris

0.89

(0.94-0.86)

0.028

5.496

Hvlonycieris

0.88

Mtisonvcteris

0.82

Megadermatidae

1.03

(1.09-0.95)

0.029

6.294

All bats

153

1.02

(1.43-0.74)

0.010

11.544

Molossidae

0.95

(1.10-0.85)

0.026

8.256

Pteropodidae

0.92

(1.04U80)

0.013

7.458

Rhinopomatidae

0.86

306

SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

Table A 1 1. — Ranked means and statistics for the relative length of digit V.

Taxon

Mean

Max-min

± 1 SE

Natalidae

1.26

Furipteridae

1.24

Nycteridae

1.14

Thyropteridae

1.12

Myzapodidae

1.11

Craseonycteridae

1.10

Megadermatidae

1.07

(1.18-0.99)

0.038

7.920

Rhinolophidae

1.04

(1.21-0.86)

0.046

11.774

Phyllostomatidae

0.98

(1.14-0.76)

0.015

10.554

Carolliinae

1.09

(1.11-1.07)

0.021

2.713

Caro Ilia

1.11

(1.21-1.01)

0.043

7.690

Rhinoph ylla

1.07

(1.15-1.01)

0.041

6.683

Phyllostomatinae

1.07

(1.12-0.93)

0.017

5.211

Maerophyllum

1.12

(1.17-1.08)

0.044

5.498

Minion

1.12

(1.24-1.00)

0.052

10.289

Vampyrum

1.11

Micronycteris

1.10

(1.19-0.96)

0.021

6.469

Chrotopterus

1.10

Lonchorhinu

1.07

(1.13-0.99)

0.042

6.774

Tonatia

1.06

(1.21-0.89)

0.040

10.035

Trachons

PhyUoaermu

1.05

1.04

Macrotus

1.03

(1.12-0.94)

0.090

12.441

Phyllostonius

0.93

(1.01-0.88)

0.025

5.971

Stenoderminae

1.02

(1.14-0.89)

0.017

6.888

Ardops

1.14

Phyllops

1.11

(1.21-1.01)

0.099

12.597

Pvgoderma

1.10

Stenodermu

1.09

Cenlurio

1.07

Vamp yrodes

1.06

Ectophylla

Art in e us

1.04

(1.08-0.99)

0.042

5.669

1.04

(1.17-0.93)

0.019

6.750

Vampyrops

1.03

(1.13-0.94)

0.016

4.962

Ametrida

1.01

(1.07-0.95)

0.061

8.608

Vumpyressu

1.01

(1.12-0.93)

0.035

7.779

Chiroderma

1.00

(1.02-0.98)

0.007

1.554

Sturnira

0.98

(1.08-0.85)

0.024

7.917

Ariteus

0.97

Uroderma

0.94

(0.96-0.92)

0.021

3.186

Sphaeronycieris

Enchisthenes

0.91

0.89

Phyllonycterinae

0.93

(0.99-0.87)

0.034

6.389

Erophylla

0.99

(1.02-0.96)

0.027

3.915

Brachyphylla

0.93

(0.96-0.91)

0.028

4.249

Phyllonycteris

0.87

(0.93-0.81)

0.061

9.876

Desmodontinae

0.92

(1.02-0.76)

0.085

15.868

Desmodus

1.02

(1.04-1.00)

0.018

2.472

Diphvlla

0.99

Diaemus

0.76

Glossophaginae

0.87

(0.97-0.76)

0.015

6.188

Scleronvcteris

0.97

Lionycteris

0.93

Lichonycteris

0.90

(0.91-0.89)

0.006

1.213

Plalalina

0.90

Choeronycteris

0.90

Gtossophaga

0.89

(0.90-0.87)

0.007

1.554

Choeroniscus

0.86

(0.91-0.79)

0.021

5.493

Lonchophylla

0.85

(0.90-0.82)

0.016

4.147

Monophyllus

0.84

(0.87-0.82)

0.024

4.045

Anoura

0.84

(0.93-0.75)

0.037

9.846

Leptonycteris

0.83

(0.90-0.79)

0.034

7.116

Hvlonycteris

0.81

Musonycteris

0.76

Noctilionidae

0.98

Emballonuridae

0.98

(1.38-0.72)

0.052

18.283

Mormoopidae

0.98

(0.98-0.97)

0.009

1.250

All bats

153

0.94

(1.38-0.57)

0.012

16.083

Vespertilionidae

0.94

(1.22-0.57)

0.025

15.026

Mystacinidae

0.87

Rhinopomatidae

0.87

Pteropodidae

0.85

(1.01-0.72)

0.014

8.678

Molossidae

0.63

(0.72-0.57)

0.015

7.133

BIOLOGY OF THE PHYLLOSTOMATIDAE

307

Table A12. — Ranked means and statistics for the percentage contributed to length of digit III

by the metacarpal.

Taxon

Mean

Max-min

± 1 SE

Rhinopomatidae

61.70

Emballonuridae

57.67

(63.21-53.06)

0.920

5.524

Furipteridae

54.80

Thyropteridae

53.85

Mystacinidae

53.37

Mormoopidae

52.87

(56.37-49.36)

3.505

9.375

Vespertilionidae

51.74

(63.40-41.82)

0.708

7.621

Rhinolophidae

51.57

(56.03-46.57)

1.246

6.391

Molossidae

50.70

(54.42-43.21)

1.015

6.004

Natalidae

49.92

All bats

153

47.04

(63.40-35.39)

0.533

14.017

Phyllostomatidae

Desmodontinae

45.12

50.53

(54.67-36.15)

(54.67-48.44)

0.456

2.068

7.081

7.088

Desmodus

54.67

(55.89-53.44)

1.227

3.175

Diaemus

48.48

Diphvlla

48.44

Phyllonycterinae

48.71

(49.70-47.69)

0.581

2.065

Phyllonycteris

49.70

(50.43-48.98)

0.727

2.069

Brachyphvlta

48.73

(49.53-47.92)

0.805

2.336

Erophylla

47.69

(47.85-47.54)

0.156

0.462

Glossophaginae

46.99

(49.43-44.66)

0.413

3.167

Leptonycteris

49.43

(50.21-48.95)

0.393

1.377

Lionycteris

48.71

Plaialina

48.37

Glossophaga

48.02

(48.33-47.18)

0.283

1.177

Lonchophylla

47.72

(49.55-46.75)

0.534

2.503

Monophyllus

47.13

(47.47-46.79)

0.342

1.026

Musonycteris

47.12

Hvlonycteris

46.98

Choeroniscus

46.80

(47.75-45.32)

0.435

2.077

Choeronycteris

45.84

Lichonycteris

45.13

(45.65-44.22)

0.460

1.765

Scleronycteris

45.02

Anoura

44.66

(46.46-43.17)

0.542

2.714

Phyliostomatinae

43.61

(48.03-36.15)

1.178

8.960

Phyllostomus

48.03

(49.21-46.62)

0.500

2.330

Lonchorhinu

47.34

(48.55-45.80)

0.810

2.963

Macrophylluni

46.66

(46.82-46.50)

0.158

0.480

Phylloderma

46.23

Micronvcteris

46.17

(48.80-43.72)

0.588

4.415

Macrotus

44.35

(44.39-44.32)

0.035

0.111

Minion

43.70

(45.18-42.78)

0.473

2.419

Tonal ia

42.21

(43.08-41.27)

0.276

1.729

Trachops

40.82

Chrotopterus

38.08

Vampyrum

36. 1 5

Stenoderminae

43.41

(45.49-40.92)

0.273

2.594

Uroderma

45.49

(45.52-45.46)

0.028

0.088

Artibeus

44.76

(46.66-43.21)

0.279

2.251

Enchisthenes

44.75

Ariteus

44.34

Vampyrodes

43.97

Ectophylla

43.68

(43.69-43.68)

0.005

0.015

Sphaeronycteris

Ardops

43.67

43.58

Chiroderma

43.57

(45.49-42.29)

0.676

3.467

Stenodenna

43.42

Sturnira

43.11

(44.51-41.67)

0.323

2.370

Vampyressa

43.11

(44.34-40.57)

0.659

3.416

Phyllops

42.97

(44.08-41.86)

1.115

3.669

Centurio

42.37

Vampyrops

42.25

(44.06-40.46)

0.426

3.187

Amelrida

42.00

(42.35-41.66)

0.345

1.161

Pygoderma

40.92

1.634

Carolliinae

42.34

(42.83-41.85)

0.489

Carollia

42.83

(43.39-42.37)

0.241

1.126

Rhinophyllu

41.85

(42.81-40.06)

0.896

3.708

Myzapodiaae

44.82

Noctilionidae

44.77

Craseonycteridae

43.44

Nycteridae

41.44

Megadermatidae

39.43

(40.53-37.48)

0.566

3.210

Pteropodidae

38.71

(42.49-35.39)

0.311

4.396

308

SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

Table A13. — Ranked means and statistics for the percentage contributed to length of digit III

by the first phalanx.

Taxon

Mean

Max-min

± 1 SE

Nycteridae

28.50

Pteropodidae

27.16

(29.52-25.32)

0.186

3.743

Thyropteridae

Megaaermatidae

23.32

22.65

(25.68-21.22)

0.788

7.776

Molossidae

22.30

(26.36-19.15)

0.701

9.425

Natalidae

21.38

Rhinolophidae

20.39

(23.54-12.46)

1.390

18.039

Myzapodidae

20.26

All bats

153

20.14

(29.52- 9.96)

0.379

23.292

Vespertilionidae

19.94

(24.70-11.77)

0.373

10.417

Emballonuridae

17.33

(21.77-10.76)

1.086

21.711

Phyllostomatidae

16.89

(21.00-10.21)

0.329

13.638

Carolliinae

18.93

(19.00-18. 87)

0.065

0.488

Caroltia

19.00

(19.23-18.55)

0. 1 54

1.619

Rhinophylla

18.87

(19.34-18.37)

0.279

2.560

Phyllonycterinae

17.77

(20.14-15.60)

1.312

12.791

Erophvlla

20.14

(20.45-19.83)

0.309

2.170

Phyllonvcteris

17.57

(18.13-17.01)

0.562

4.525

Bracliyphvlla

15.60

(16.06-15.15)

0.453

4.108

Phyllostomatinae

17.73

(21.00-13.03)

0.667

12.472

Vampyruni

21.00

Chrotopterus

19.99

Macrotus

18.84

(19.15-18.54)

0.305

2.292

Micronycteris

18.80

(21.17-16.87)

0.437

8.047

Tonal in

18.77

(20.16-17.72)

0.320

4.511

Macrophvllum

18.46

(18.90-18.02)

0.439

3.366

Trachops

17.55

Million

16.29

(19.37-14.51)

1.053

14.445

Lonchorhina

16.28

(17.61-15.50)

0.668

7.105

Phvlloderma

16.04

Phyllostomus

13.03

(14.25-12.07)

0.426

7.304

Glossophaginae

17.08

(18.76-14.98)

0.365

7.706

Platalina

18.76

Choeronycieris

18.53

Scleronycteris

18.11

Lichonycteris

18.09

(18.61-17.48)

0.329

3.150

Hvlonycteris

17.95

Glossophaga

17.67

(18.09-16.89)

0.267

3.021

Musonycteris

17.47

Choeroniscus

17.15

(17.81-16.41)

0.225

2.937

Lonchophyllu

16.90

(19.41-16.05)

0.632

8.354

Monophyllus

15.99

(16.30-15.68)

0.308

2.721

Anoura

15.23

(16.09-14.65)

0.265

3.886

Lionvcteris

15.22

Leptonvcteris

14.98

(15.48-14.40)

0.312

3.609

Stenoderminae

16.94

(19.58-14.97)

0.350

8.513

Centurio

19.58

Pvgoderma

19.44

Phyllops

19.13

(19.48-18.77)

0.356

2.632

Vampyressa

17.75

(19.57-16.58)

0.664

8.365

Chiroderina

17.25

(18.45-15.81)

0.488

6.329

Sphaeronvcteris

17.25

Ectophvlla

17.20

(19.08-15.33)

1.876

15.423

Sturnira

17.19

(18.55-16.55)

0.182

3.344

Enchisthenes

17.01

Vampyrops

Uroaerma

16.78

(17.69-15.31)

0.237

4.465

16.52

(16.72-16.32)

0.200

1.716

Vampvrodes

16.37

Arlibeus

15.80

(17.66-14.11)

0.316

7.220

Ainetrida

15.44

(15.57-15.31)

0.131

1.204

Slenoderina

15.25

Ardops

15.08

Ariteus

14.97

Desmodontinae

10.51

(10.99-10.21)

0.244

4.015

Desmodus

10.99

(11.03-10.95)

0.041

0.525

Diphvlla

10.33

Diaemus

10.21

Mystacinidae

14.33

Craseonycteridae

14.32

Rhinopomatidae

13.42

Noctilionidae

12.30

Mormoopidae

11.18

(11.36-11.00)

0.182

2.302

Furipteridae

9.96

BIOLOGY OF THE PHYLLOSTOMATIDAE

309

Table A14. — Ranked means and statistics for the percentage contributed to length of digit III

by the second phalanx.

Taxon

Mean

Max-min

± 1 SE

Craseonycteridae

42.23

Megadermatidae

37.92

(40.32-35.44)

0.782

4.612

Furipteridae

35.23

Pteropodidae

34.13

(37.82-29.33)

0.347

5.566

Noctilionidae

30.95

Nycteridae

30.05

Natalidae

28.70

Rhinolophidae

28.03

(39.31-23.55)

2.245

21.185

All bats

153

25.47

(42.23-14.21)

0.528

25.647

Emballonuridae

24.99

(28.71-21.06)

0.691

9.574

Rhinopomatidae

24.88

Phyllostomatidae

23.62

(28.68-18.04)

0.297

8.810

Carolliinae

24.68

(25.03-24.33)

0.354

2.030

Rhinophylla

25.03

(25.93-24.17)

0.507

3.508

Carollia

24.33

(25.30-23.44)

0.396

3.257

Stenoderminae

24.39

(28.68-21.11)

0.405

6.840

Pvgoderma

28.68

Vampyrodes

25.84

Ariteus

25.37

Stenoderma

25.18

Centurio

25.00

Ardops

24.89

Sphaeronycteris

24.57

Vampyrops

24.51

(26.17-23.57)

0.280

3.610

Chiroderma

24.49

(25.47-23.68)

0.340

3.105

Vampyressa

24.43

(25.66-22.83)

0.469

4.294

Artibeus

24.28

(25.43-23.52)

0.164

2.440

Ametrida

24.17

(24.50-23.84)

0.330

1.930

Uroderma

23.77

(24.14-23.41)

0.362

2.153

Ectophylla

23.67

(24.51-22.84)

0.835

4.989

Phyllops

23.23

(23.27-23.19)

0.039

0.236

Sturnira

21.44

(22.84-20.41)

0.237

3.494

Enchisthends

21.11

Glossophaginae

23.73

(26.05-21.92)

0.296

4.504

Lionyeteris

26.05

Anoura

24.79

(25.53-24.30)

0.248

2.234

Lichonycteris

24.60

(25.07-23.91)

0.350

2.467

Scleronycieris

24.06

Hvlonycteris

23.98

Choeroniscus

23.93

(24.54-23.18)

0.233

2.178

Leptonycteris

23.87

(24.54-23.14)

0.404

2.934

Lonchophylla

23.44

(24.38-22.69)

0.272

2.593

Monophyllus

23.27

(24.09-22.45)

0.820

4.982

Musonycteris

23.11

Choeronycteris

23.03

Glossophaga

22.41

(23.25-21.54)

0.358

3.198

Platalina

21.92

Desmodontinae

23.54

(26.32-18.80)

2.385

17.547

Diaemus

26.32

Diphvlla

25.52

1.445

Desmodus

18.80

(18.99-18.60)

0.192

Phyllostomatinae

23.45

(26.41-20.27)

0.529

7.484

True hops

26.41

Lonchorhina

25.77

(26.26-24.97)

0.402

2.704

Mimon

24.42

(25.29-23.17)

0.346

3.168

Vampyrum

23.84

Phyllostomus

23.75

(24.31-22.50)

0.323

3.041

Macrophyllum

23.42

(23.43-23.42)

0.003

0.019

Chrotopterus

23.34

0.630

9.612

Micronvcteris

22.69

(27.07-20.16)

Phylloderma

22.55

(22.41-20.36)

0.302

3.724

Tonal ia

21.45

Macrotus

20.27

(20.28-20.27)

0.005

0.036

Phyllonycterinae

18.87

(20.47-18.04)

0.803

7.371

Brachyphylla

20.47

(20.50-20.44)

0.031

0.213

Erophylla

18.08

(18.72-17.45)

0.636

4.976

Phyllonycteris

18.04

(18.57-17.52)

0.523

4.103

Myzapodidae

21.59

0.272

4.023

Molossidae

20.31

(21.51-18.76)

Mormoopidae

19.31

(23.16-15.46)

3.850

28.203

Vespertilionidae

19.30

(33.77-14.21)

0.790

22.797

Mystacinidae

17.60

Thyropteridae

14.86

310

SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

Table A15. — Ranked means and statistics for the percentage contributed to length of digit III

by the third phalanx.

Taxon

Mean

Max-min

± 1 SE

Mormoopidae

16.64

(16.81-16.48)

0.163

1.386

Mystacinidae

14.70

Phyllostomatidae

Desmodontinae

14.36

(19.01-10.02)

0.318

15.510

15.42

(15.71-14.99)

0.218

2.449

D ip h Vila

15.71

Desmodus

15.55

(16.62-14.47)

1.076

9.789

Diaemus

14.99

Stenoderminae

15.26

(18.39-10.96)

0.446

12.044

Anietrida

18.39

(19.19-17.58)

0.806

6.200

Sturnira

18.26

(20.31-16.48)

0.359

6.222

Enchisthenes

17.13

Vampyrops

16.46

(18.17-14.15)

0.384

7.374

Ardops

16.45

Stenodenna

16.15

Ectaphylla

15.44

(16.49-14.40)

1.045

9.574

Ariteus

15.32

Artiheus

15.16

(16.79-13.92)

0.240

5.708

Vampyressa

14.71

(15.72-14.08)

0.326

4.953

Chiroderma

14.69

(15.40-13.98)

0.241

3.674

Phvllops

14.67

(16.18-13.16)

1.510

14.551

Sphaeronycteris

14.52

Urodermu

14.22

(14.41-14.03)

0.190

1.887

Vampyrodes

13.82

Centurio

13.05

Pvgoderma

10.96

Phyilostomatinae

15.21

(19.01-10.62)

0.835

18.209

Vamp yrum

19.01

Chrotopterus

18.59

Tonal ia

17.57

(19.17-15.18)

0.467

7.037

Macrotus

16.53

(16.80-16.27)

0.265

2.270

Mimon

15.59

(17.89-12.56)

0.988

14.167

Trachops

15.23

Phyllostonius

15.20

(16.03-14.18)

0.335

4.925

Phylloderma

15.18

Micronycteris

12.35

(15.57- 7.56)

0.671

18.828

Macrophyllum

11.46

<11.74-1 1.18)

0.278

3.429

Lonchorhina

10.62

(11.62- 9.69)

0.560

9. 1 30

Phyllonycterinae

14.66

(15.20-14.09)

0.321

3.791

Brachyphvlla

15.20

(15.52-14.88)

0.321

2.984

Phyllonycteris

14.68

(16.50-12.87)

1.813

17.458

Erophylta

14.09

(14.26-13.92)

0.172

1.722

Carolliinae

14.05

(14.25-13.85)

0.200

2.013

Rhino phyllu

14.25

(17.40-12.37)

1.584

19.259

Carollia

13.85

(14.98-13.10)

0.430

6.217

Glossophaginae

12.20

(15.32-10.02)

0.360

10.640

Anoura

15.32

(17.46-11.92)

1.011

14.753

Monophyllus

13.61

(14.47-12.76)

0.854

8.868

Scleronycteris

12.81

Choeronycteris

12.60

Musonycteris

12.31

Lichonyeteris

12.18

(12.36-11.92)

0.133

1.885

Choeroniscus

12.13

(13.02-1 1.55)

0.250

4.601

Lonchophylla

11.94

(14.28- 9.91)

0.808

15.128

Glossophaya

11. 90

(12.71-10.99)

0.382

6.415

Leptonvcteri s

1 1.73

(12.72-10.21)

0.773

11.415

Hvlonycteris

1 1.08

Piatalina

10.96

Lionyceris

10.02

Myzapodidae

13.33

All bats

12.14

(20.13- 4.64)

0.378

30.027

Noctilionidae

11.98

Vespertilionidae

9.02

(20.13- 0.00)

0.672

41.474

Thyropteridae

7.97

Molossidae

6.68

( 8.92- 4.64)

0.506

22.711

BIOLOGY OF THE PHYLLOSTOMATIDAE

311

Table A16. — Ranked means and statistics for the percentage contributed to length of digit IV

by the metacarpal.

Taxon

Mean

Max-min

± 1 SE

Thyropteridae

68.56

Emballonuridae

66.37

(70.34-63.19)

0.690

3.600

Mormoopidae

65.88

(66.33-65.44)

0.442

0.949

Nataiidae

64.78

Furipteridae

64.38

Molossidae

63.63

(68.78-54.62)

1.557

7.342

Mystacinidae

62.95

Rhinolophidae

62.92

(65.78-59.65)

0.864

3.633

Vespertilionidae

62.67

(73.59-58.48)

0.577

5.128

Craseonycteridae

61.46

Noctilionidae

60.13

Nycteridae

59.88

Rhinopomatidae

59.77

Phyllostomatidae

Desmodontinae

58.95

(67.11-47.84)

0.425

5.045

63.63

(67.11-61.36)

1.765

4.805

Desmodus

67.11

(68.09-66.13)

0.980

2.066

Diphvtla

62.43

Diaem us

61.36

Glossophaginae

60.72

(63.09-59.66)

0.263

1.562

Hvlonycteris

63.09

Lionycleris

61.70

Scleronycteris

61.28

Anoura

61.17

(62.12-58.96)

0.574

2.100

Lichonycteris

60.85

(62.43-59.77)

0.808

2.299

Monophyllus

60.83

(61.49-60.18)

0.656

1.525

Choeroniscus

60.46

(61.05-59.61)

0.277

1.024

Musonycteris

60.43

0.483

Glossophaga

60.12

(61.07-58.97)

1.606

Lonchophylla

60.06

(61.59-58.75)

0.499

1.857

Choeronycteris

59.84

Leptonycteris

59.79

(60.33-58.78)

0.507

1.470

Platalina

59.66

Phyllonycterinae

59.16

(59.54-58.72)

0.237

0.695

Phyllonycieris

59.54

(60.32-58.77)

0.775

1.841

Brachyphyllu

59.21

(59.77-58.64)

0.568

1.357

Eraphylla

58.72

(58.88-58.57)

0.152

0.366

Phyllostomatinae

58.48

(63.24-52.65)

1.060

6.013

Phyllodermu

63.24

Phyllosromus

63.01

(64.30-61.83)

0.485

1.723

Lonchorhina

61.41

(63.66-59.56)

1.201

3.388

Minion

60.75

(64.94-56.94)

1.495

5.502

Micronycteris

59.79

(63.15-56.99)

0.578

3.347

Macrophvlluni

59.14

(60.04-58.24)

0.898

2.147

Macrotus

56.79

(57.44-56-13)

0.659

1.641

Trachops

55.96

4.957

Tonal ia

55.93

(60.05-53.09)

1.048

Chrotopterus

54.63

Vampyrum

52.65

0.628

4.519

Stenoderminae

57.29

(59.08-47.84)

Uroderma

59.08

(59.11-59.05)

0.034

0.081

Srenoderma

58.75

Chiroderma

58.67

(60.47-57.08)

0.735

2.802

Artibeus

58.56

(59.93-56.96)

0.229

1.410

Phyllops

58.53

(59.46-57.60)

0.929

2.244

Ardops

58.38

0.173

Ectophylla

58.16

(58.33-57.99)

0.422

Vampyressu

58.12

(58.83-56.63)

0.395

1.521

Arireus

58.04

Vampyrodes

57.98

Siurnira

57.65

(59.91-55.77)

0.368

2.020

Enchisthenes

57.63

Vampvrops

57.60

(60.27-56.03)

0.426

2.339

Sphaeronvcteris

56.84

Centurio

56.08

Pvgoderma

55.92

1.241

Ametrida

47.84

(48.26-47.42)

0.420

Carolliinae

56.97

(57.16-56.77)

0.195

0.484

Caro lli a

57.16

(58.11-56.20)

0.459

1.607

Rhinophylla

56.77

(58.04-55.50)

0.734

2.239

Megadermatidae

58.66

(60.84-56.58)

0.712

2.715

All bats

153

58.59

(73.59-41.96)

0.567

11.978

Myzapodidae

57.18

0.451

5.306

Pteropodidae

46.52

(53.21-41.96)

312

SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

Table A17. — Ranked means and statistics for the percentage contributed to length of digit IV

by the first phalanx.

Taxon

Mean

Max-min

± 1 SE

Pteropodidae

26.23

(28.69-22.90)

0.291

6.069

Molossidae

23.62

(27.95-17.82)

1.128

14.327

Rhinopomatidae

22.57

Nycteridae

21.62

Myzapodidae

20.88

All bats

153

20.49

(28.69-10.23)

0.327

19.719

Vespertilionidae

20.25

(25.71-13.30)

0.448

12.318

Rhinolophidae

19.41

(21.07-15.88)

0.687

9.358

Thyropteridae

18.93

Emballonuridae

18.88

(22.35-14.01)

0.684

12.559

Phyllostomatidae

18.10

(22.93-11.92)

0.362

14.008

Carolliinae

20.60

(20.99-20.20)

0.394

2.708

Caro Ilia

20.99

(21.21-20.63)

0. 1 36

1.295

Rhino phylla

20.20

(21.15-19.38)

0.514

4.407

Phyllostomatinae

19.23

(22.93-14.33)

0.818

14.113

Mucrotus

22.93

(22.98-22.88)

0.048

0.298

Vampyrum

22.00

Tonal ia

21.84

(23.10-20.50)

0.429

5.197

Chrotopterus

21.04

Micronycteris

19.48

(22.83-17.35)

0.556

9.888

Trachops

19.15

Macrophyllum

18.95

(19.44-18.46)

0.488

3.640

Lonehorhina

18.49

(19.91-17.42)

0.738

6.914

Mimon

18.21

(20.75-15.92)

0.886

10.877

Phylloderma

15.17

Phvllostomus

14.33

(16.93-12.10)

0.910

14.207

Stenoderminae

19.08

(22.12-16.84)

0.353

7.619

Centurio

22.12

Ametrida

21.31

(21.35-21.27)

0.039

0.260

Phyllops

20.42

(20.47-20.38)

0.044

0.306

Ectophylla

20.07

(21.53-18.60)

1.465

10.328

Chiroderma

19.91

(21.13-18.80)

0.416

4.673

Vampyressa

19.55

(21.48-17.70)

0.751

8.592

Vampyrops

19.45

(20.51-18.04)

0.256

4.161

Enchisihenes

19.26

Sturnira

19.03

(19.89-18.37)

0.156

2.590

Uroderma

18.90

(18.93-18.87)

0.029

0.217

Vampyrodes

18.76

Pvgodernta

18.34

Stenoderma

18.34

Artiheus

17.99

(19.85-16.12)

0.349

6.992

Ardops

17.09

Ariteus

16.99

Sphaeronvcteris

16.84

Phyllonycterinae

18.64

(20.44-17.25)

0.944

8.770

Erophylla

20.44

(20.56-20.32)

0.121

0.834

Phyllonycteris

18.22

(18.67-17.77)

0.453

3.517

Brachyphvlla

17.25

(17.47-17.03)

0.222

1.824

Glossophaginae

16.75

(18.58-14.31)

0.340

7.314

Glossophaga

18.58

(19.15-18.03)

0.260

2.794

Platalina

18.35

Choeronycteris

17.62

Scleronycteris

17.25

Lonchophylla

17.22

(17.99-16.08)

0.316

4.102

Lichonycteris

17.1 1

(17.89-16.51)

0.408

4.136

Choeroniscus

16.99

(17.71-15.70)

0.365

4.807

Musonyeteris

16.88

Leptonycteris

16.45

(16.75-16.06)

0.202

2.123

Anoura

16.17

(17.15-15.34)

0.320

4.420

Hylonycteris

16.02

Monophyllus

14.81

(14.92-14.70)

0.112

1.074

Lionycteris

14.31

Desmodontinae

12.05

(12.14-11.92)

0.065

0.931

Diphylla

12.14

Desmodus

12.09

(12.63-11.54)

0.548

6.412

Diaemus

11.92

Megadermatidae

17.86

(19.85-16.18)

0.746

9.338

Natalidae

17.28

Mormoopidae

16.64

(18.28-15.01)

1.634

13.885

Mystacinidae

16.01

Furipteridae

15.98

Noctilionidae

10.61

Craseonycteridae

10.23

BIOLOGY OF THE PHYLLOSTOMATIDAE

313

Table A18. — Ranked means and statistics for the percentage contributed to length of digit IV

by the second phalanx.

Taxon

Mean

Max-min

± 1 SE

Noctilionidae

29.27

Craseonycteridae

28.31

Pteropodidae

27.25

(29.77-22.67)

0.335

6.742

Megadermatidae

23.48

(24.93-19.30)

1.054

10.039

Phyllostomatidae

22.95

(30.84-20.10)

0.280

8.528

Desmodontinae

24.32

(26.71-20.80)

0.795

12.788

Diaemus

26.71

Diphylla

25.44

Desmodus

20.80

(21.24-20.37)

0.432

2.940

Stenoderminae

23.63

(30.84-21.05)

0.577

10.061

Ametrida

30.84

(31.22-30.46)

0.380

1.745

Sphaeronycteris

26.32

Pygoderma

25.74

Ariteus

24.97

Ardops

24.53

Artibeus

23.45

(25.33-21.41)

0.257

3.949

Sturnira

23.32

(25.59-21.69)

0.349

4.730

Vampyrodes

23.26

Enchisthenes

23.11

Vampyrops

22.95

(23.94-20.97)

0.285

3.923

Sienoderma

22.91

Vampyressa

Uroaerma

22.33

(23.71-20.43)

0.557

5.579

22.02

(22.08-21.95)

0.063

0.405

Centurio

21.80

Ectophvlla

21.77

(23.06-20.48)

1.292

8.393

Chiroderma

21.42

(23.08-19.05)

0.712

7.430

Phyllops

21.05

(21.93-20.16)

0.885

5.944

Glossophaginae

22.53

(24.35-20.90)

0.287

4.599

Monophyllus

24.35

(24.90-23.81)

0.544

3.157

Lionycteris

23.99

Leptonvcteris

23.76

(24.47-23.14)

0.387

2.822

Lonchophylla

22.72

(25.17-21.29)

0.658

6.474

Musonycteris

22.70

Anoura

22.66

(23.89-22.09)

0.321

3.169

Choeroniscus

22.55

(23.25-22.13)

0.212

2.106

Choeronycteris

22.54

Lichonycteris

22.04

(23.30-21.06)

0.662

5.199

Platalina

21.99

Scleronycteris

21.47

4.239

Glossophaga

21.29

(22.03-20.05)

0.451

Hvlonycteris

20.90

Carolliiriae

22.43

(23.02-21.85)

0.589

3.716

Rhinophylla

23.02

(25.11-21.88)

1.046

7.872

Carollia

21.85

(22.60-21.25)

0.334

3.061

Phyllostomatinae

22.28

(25.35-20.10)

0.555

8.257

Vampyrum

25.35

Trachops

24.89

Chrotopterus

24.33

0.692

6.824

Phyllostomus

22.66

(24.06-20.15)

Tonal ia

22.23

(23.96-18.74)

0.773

9.204

Macro phy llum

21.91

(23.30-20.52)

1.386

8.945

Phylloaerma

21.58

0.632

6.713

Mimon

21.05

(22.44-19.13)

Micronycteris

20.73

(22.77-18.64)

0.359

6.005

Macrotus

20.29

(20.99-19.58)

0.707

4.931

Lonchorhina

20.10

(20.84-18.92)

0.596

5.135

Phyllonycterinae

22.21

(23.54-20.84)

0.781

6.091

Brachyphylla

23.54

(23.89-23.20)

0.346

2.077

Phyllonycteris

22.24

(22.56-21.92)

0.322

2.048

Erophvlla

20.84

(21.11-20.57)

0.272

1.849

Myzapodicfae

21.94

Mystacinidae

21.04

26.054

All bats

153

20.92

(31.22- 5.27)

0.441

Furipteridae

19.63

Nycteridae

18.51

Natalidae

17.94

Rhinolophidae

17.67

(24.48-14.13)

1.458

21.842

Rhinopomatidae

17.66

1.192

9.646

Mormoopidae

17.47

(18.67-16.28)

Vespertilionidae

17.08

(26.56- 5.31)

0.771

25.135

Emballonuridae

14.75

(19.47-10.33)

0.851

19.977

Molossidae

12.75

(18.11- 5.27)

1.683

39.616

Thyropteridae

12.51

314

SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

Table A19. — Ranked means and statistics for the percentage contributed to length of digit V

by the metacarpal.

Taxon

Mean

Max-Min

± 1 SE

Noctilionidae

74.69

Vespertilionidae

68.57

(82.35-59.25)

0.808

6.557

Rhinopomatidae

68.18

Mystacinidae

67.42

Thyropteridae

65.80

Craseonycteridae

65.57

Furipteridae

63.91

Natalidae

63.86

Myzapodidae

63.85

Emballonuridae

63.57

(67.72-57.70)

0.811

4.417

Phyllostomatidae

Desmodontinae

62.12

(67.99-52.13)

0.429

4.838

66.34

(67.76-64.29)

1.049

2.740

Diaemus

67.76

Desmodus

66.96

(67.54-66.38)

0.578

1.221

Diphrlla

64.29

Phyllonycterinae

62.93

(65.09-61.36)

1.113

3.063

Brachyphylla

65.09

(65.97-64.20)

0.889

1.932

Phyllonycteris

62.36

(62.76-61.96)

0.401

0.910

Eronhyllu

Stenoaerminae

61.36

(62.06-60.66)

0.697

1.606

62.53

(64.80-52.13)

0.785

5.178

XJroderma

64.80

(65.16-64.44)

0.362

0.789

Stenoderma

64.67

Arliheus

64.58

(67.64-62.21)

0.522

2.916

Sturnira

64.52

(66.05-62.75)

0.439

2.153

Vampvrodes

64.48

Chiroderma

63.89

(65.51-62.95)

0.432

1.514

Vampyressa

63.69

(66.00-61.89)

0.707

2.482

Enchisthenes

63.67

Sphaeronvcteris

63.60

Vampyrops

63.50

(65.47-61.98)

0.372

1.854

Ariteus

63.48

Ectophvlla

63.42

(64.68-62.16)

1.258

2.806

Ardops

63.00

Phyllops

62.14

(62.72-61.56)

0.581

1.323

Pvgoderma

59.15

Centurio

58.32

Ametrida

52.13

(52.49-51.76)

0.366

0.993

Phyllostomatinae

1 1

61.51

(67.99-55.66)

1.213

6.542

Phvllostonuts

67.99

(70.39-66.08)

0.880

2.894

Phvllodenmi

67.07

Minion

64.20

(69.16-59.32)

1.886

6.570

Lonchorhina

63.38

(65.11-60.19)

1.600

4.372

Macrophyllum

62.22

(62.30-62.15)

0.074

0.169

Micronvcteris

61.67

(65.46-58.16)

0.718

4.030

Macrotus

60.48

(61.61-59.35)

1.130

2.643

True hops

59.89

Chrotopterus

57.18

Tonal in

56.91

(60.78-54.38)

0.928

4.316

Vunipvruni

55.66

Glossophaginae

61.22

(62.70-59.21)

0.254

1.495

Choeronycteris

62.70

Leptonycteris

62.40

(62.96-61.91)

0.306

0.848

Lonchophylla

62.01

(63.12-61.52)

0.290

1.046

Musonycteris

61.87

Glossophaya

61.35

(61.96-60.62)

0.291

0.950

Choeroniscus

61.32

(62.51-59.84)

0.488

1.780

Hvlonycteris

61.28

Lionycteris

60.99

Lichonycteris

60.83

(61.27-60.22)

0.315

0.896

Platalina

60.77

Scleronvcteris

60.76

Anoura

60.36

(61.06-59.07)

0.379

1.404

Monophvllns

59.21

(59.65-58.78)

0.435

1.040

Carolliinae

60.26

(60.48-60.04)

0.222

0.522

Rhinophylla

60.48

(61.79-59.66)

0.660

1.890

Carol 1 in

60.04

(60.28-59.59)

0.154

0.515

All bats

153

61.02

(82.35-46.71)

0.551

11.164

Megadermatidae

59.85

(61.13-58.53)

0.542

2.027

Nycteridae

59.39

Mormoopidae

Molossidae

59.29

(64.15-54.43)

4.861

11.593

58.88

(63.56-49.37)

1.679

8.557

Rhinolophidae

57. 1 3

(60.70-52.96)

1.091

5.052

Pteropodidae

50.78

(53.41-46.71)

0.295

3.184

BIOLOGY OF THE PHYLLOSTOMATIDAE

315

Table A20. — Ranked means and statistics for the percentage contributed to length of digit V

by the first phalanx.

Taxon

Mean

Max-min

±1 SE

Molossidae

29.46

(39.13-20.66)

1.837

18.703

Pteropodidae

24.16

(26.06-22.91)

0.160

3.636

Emballonuridae

23.42

(29.75-18.26)

1.038

15.348

Furipteridae

23.22

Rhinolophidae

23.12

(27.43-17.74)

1.464

16.759

Mormoopidae

22.22

(26.70-17.74)

4.480

28.512

Nycteridae

20.58

All bats

153

20.13

(39.13- 8.83)

0.374

22.977

Megadermatidae

20.05

(25.43-17.91)

1.395

15.555

Myzapodidae

18.92

Thyropteridae

17.75

Natalidae

17.61

Vespertilionidae

17.44

(20.61- 8.83)

0.395

12.610

Rhinopomatidae

17.20

Noctilionidae

17.15

Phyllostomatidae

16.88

(22.86-12.74)

0.303

12.565

Carolliinae

18.61

(19.00-18.23)

0.389

2.952

Caro Ilia

19.00

(19.62-18.63)

0.224

2.359

Rhinophylla

18.23

(19.52-17.06)

0.715

6.799

Phyllostomatinae

18.25

(21.50-13.93)

0.708

12.858

Tonal ia

21.50

(23.1 1-20.52)

0.337

4.143

Macrotus

20.67

(21.28-20.06)

0.613

4.196

Vampyrum

19.99

Chrotopterus

19.66

Micronycteris

19.60

(21.83-17.53)

0.478

8.448

Trachops

18.66

Lonchorhina

17.47

(18.81-16.80)

0.668

6.617

Macrophyllum

17.39

(17.60-17.18)

0.211

1.715

Mimon

16.74

(20.05-14.74)

1.106

14.780

Phylloderma

15.19

Phyllostomus

13.93

(15.96-12.05)

0.701

11.264

Phyllonycterinae

16.89

(19.00-15.58)

1.067

10.940

Erophylla

19.00

(19.19-18.81)

0.190

1.413

Brachyphylla

16.09

(16.33-15.84)

0.247

2.170

Phyllonycteris

15.58

(16.35-14.81)

0.769

6.986

Stenoderminae

16.71

(22.86-14.88)

0.547

13.491

Centurio

22.86

Ameirida

21.07

(21.35-20.79)

0.279

1.872

Sphaeronycteris

18.90

Phyllops

17.26

(17.42-17.10)

0.161

1.317

Ectophylla

16.98

(17.89-16.08)

0.904

7.524

Pyyoderma

16.64

Vampyre ssa

16.32

(17.70-14.64)

0.515

7.063

Chiroderma

16.14

(16.84-15.08)

0.320

4.426

Vumpyrops

16.09

(17.30-15.05)

0.241

4.744

Stenoderina

15.71

Vainnyrodes

15.36

Uroaerma

15.27

(16.09-14.45)

0.818

7.576

Sturnira

15.24

(16.57-14.08)

0.259

5.366

Arileu.s

15.21

Ardops

15.10

Artibeus

14.94

(16.89-13.48)

0.317

7.652

Enchisthenes

14.88

Glossophaginae

16.38

(17.83-14.89)

0.246

5.422

Platalina

17.83

Glossophaga

17.69

(18.31-16.83)

0.327

3.697

Monophyllus

17.15

(17.31-16.99)

0.162

1.338

Scleronycteris

16.93

Lonchophylla

16.66

(18.16-15.52)

0.437

5.862

Choeroniscus

16.38

(17.21-15.19)

0.332

4.537

Lionycteris

16.34

Leptonycieris

16.24

(16.25-16.23)

0.006

0.059

Lichonycteris

16.21

(16.57-15.73)

0.250

2.670

Musonycteris

15.77

Choeronycieris

15.76

Hylonycteris

15.11

Ano ura

14.89

(15.58-14.18)

0.279

4.196

Desmodontinae

13.75

(15.13-12.74)

0.712

8.970

Diphylla

15.13

Diaeinus

13.39

Desmodus

12.74

(12.85-12.64)

0.102

1.129

Craseonycteridae

15.79

Mystacinidae

11.59

316

SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

Table A21. — Ranked means and statistics for the percentage contributed to length of digit V

by the second phalanx.

Taxon

Mean

Max-min

±1 SE

Pteropodidae

25.06

(28.54-22.08)

0.291

6.357

Phyllostomatidae

21.00

(26.80-17.49)

0.275

9.161

Glossophaginae

22.40

(24.75-20.95)

0.310

4.990

Anoura

24.75

(25.35-24.47)

0.163

1.473

Monophyllus

23.64

(23.91-23.36)

0.273

1.634

Hvlonycteris

23.62

Lichonycteris

22.96

(23.44-22.42)

0.295

2.229

Lionycteris

22.67

Musonycteris

22.36

Scleronycteris

22.31

Choeroniscus

22.30

(24.97-20.80)

0.711

7.129

Choeronycleris

21.54

Platalina

21.40

Leptonycteris

21.36

(21.84-20.80)

0.302

2.448

Lonchophylla

21.33

(22.51-19.98)

0.464

4.860

Glossophaga

20.95

(21.82-19.73)

0.465

4.442

Carolliinae

21.13

(21.29-20.96)

0.166

1.112

Rhinophylla

21.29

(21.90-20.81)

0.322

2.620

Caro Ilia

20.96

(21.17-20.78)

0.102

0.970

Stenoderminae

20.76

(26.80-17.49)

0.509

10.101

Ametrida

26.80

(26.89-26.72)

0.087

0.459

Pvgodernia

24.21

Ardops

21.89

Enchisthenes

21.45

Art ileus

21.31

Phvt lops

20.60

(21.02-20.18)

0.421

2.889

Ariibeus

20.47

(22.13-18.87)

0.282

4.964

Vampyrops

20.40

(21.39-19.16)

0.277

4.288

Sturnira

20.24

(22.43-18.28)

0.367

5.730

Vampyrodes

20.16

Vampyressa

19.99

(22.58-17.42)

1.073

11.997

Chirodenna

19.97

(21.06-17.72)

0.578

6.476

Uroderma

19.93

(20.38-19.47)

0.456

3.238

Stenoderma

19.61

Ectophylla

19.60

(19.95-19.24)

0.355

2.559

Centurio

18.82

Sphaeronycteris

17.49

Phyllostomatinae

20.23

(24.35-17.75)

0.649

10.646

Vampyrum

24.35

Chrotopterus

23.16

Tonal ia

21.60

(23.92-18.62)

0.680

8.334

Trachops

21.45

Macro ph yll uni

20.39

(20.52-20.25)

0.137

0.948

Lonchorhina

19.14

(21.00-18.07

0.934

8.451

Minion

19.06

(20.93-16.02)

0.880

10.327

Macrotus

18.85

(19.37-18.33)

0.517

3.878

Micronycteris

18.73

(21.43-16.26)

0.441

8.151

Phyllostomus

18.08

(18.90-17.51)

0.269

3.323

Phylloderma

17.75

Phyllonycterinae

20.18

(22.07-18.83)

0.973

8.351

Phyllonycteris

22.07

(22.43-21.70)

0.368

2.358

Erophylla

19.64

(20.15-19.13)

0.507

3.650

Brachyphylla

18.83

(19.47-18.19)

0.642

4.826

Desmodontinae

19.91

(20.58-18.85)

0.536

4.663

Diphylla

20.58

Desmodus

20.30

(20.78-19.82)

0.476

3.319

Diaeni us

18.85

Mystacinidae

21.00

Megadermatidae

20.09

(21.41-16.04)

1.017

11.315

Nycteridae

20.03

Rhinolophidae

19.75

(22.56-18.18)

0.637

8.532

All bats

153

18.84

(28.54- 8.16)

0.406

26.629

Craseonycteridae

18.64

Natalidae

18.54

Mormoopidae

Myzapodidae

18.48

17.23

(18.87-18.10)

0.380

2.910

Thyropteridae

16.46

Rhinopomatidae

14.62

Vespertilionidae

13.98

(21.09- 8.83)

0.559

22.239

Emballonuridae

13.01

(16.23- 9.55)

0.580

15.451

Furipteridae

12.87

Molossidae

1 1.66

(15.78-10.16)

0.570

14.663

Noctilionidae

8.16

REPRODUCTIVE PATTERNS

Don E. Wilson

“It follows, then, that an ecologist setting out to learn the workings of some part
of the natural world must study the strategies of individual species. The question
he must ask himself is: What are the tricks used to turn resources into babies?”
This quotation from Colinvaux (1973) may be appropriate to describe the fol¬
lowing approach to the problem of reproductive strategies in phyllostomatid bats.

One function of a review paper is to provide the reader with a summary of
available data and pertinent references with which to pursue the subject. I have
attempted to do this at the species level, although nothing is known concerning
reproduction in some species of leaf-nosed bats.

Knowledge of bat reproductive patterns has undergone a spurt in growth in
recent years, as has been the case in many other fields. Enough information is now
available to make speculation tantalizing, but not enough to make generalization
rewarding. Nevertheless, some general patterns seem to be widespread within
the primarily tropical phyllostomatids.

Early knowledge of bat reproduction was based mainly on temperate-zone
species. Reproductive cycles in temperate regions usually are forced into a tightly
controlled and relatively short time-span owing to rigors of the climate. Thus,
most temperate-zone bats produce only one young per year, and populations are
highly synchronized.

Some members of the family Phyllostomatidae, such as Macrotus and Lep-
tonycteris, extend northward into subtropical and temperate zones. Macrotus
californicus probably has the most distinctive reproductive pattern in the family.
Not only is this species monestrous and monotocous, but it has a rather unique
system of delayed development that allows the embryo to stay dormant during
unfavorable times of the year (Bradshaw, 1961).

A variation of this pattern is seen in Artibeus jamaicensis in Panama (Fleming,
1971). These animals have young in March or April, followed by a postpartum
estrus and a second pregnancy. This results in the second young being born in
July or August; another postpartum estrus follows. The embryos from the second
postpartum estrus implant in the uterus, but undergo delayed development until
November, when they resume the normal pace of development and finish the
cycle with parturition in March or April.

A far more common pattern is one of bimodal or seasonal polyestry similar
to that of Artibeus jamaicensis, but without delayed development. Fleming
(1973) and Wilson (1973) have discussed this pattern for Panamanian and
Costa Rican bats. Members of the genera Glossophaga, Carollia, Uroderma, and
Artibeus commonly have this bimodal pattern. In Panama, this pattern involves
birth peaks in March-April and July-August. In Costa Rica, the peaks may be
shifted to February-March and June-July (Fleming et al., 1972). In Colombia,

317

318

SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

the pattern seems to have shifted still more with birth peaks occurring in January-
February and May-June for some species. A recent paper by Taddei (1976) has
confirmed this pattern for many species in Brazil as well.

In many of the polyestrous species there is so much asynchrony within a given
population that it is difficult to determine if the individual animals are producing
more than one young per year or if they simply are out of phase with each other.
The presence of females that are both pregnant and lactating is one simple indica¬
tion of polyestry.

It may be possible for some polyestrous individuals to produce three young
per year, as is the case with Myotis nigricans, a neotropical vespertilionid (Wilson
and Findley, 1970). Variation in age of first reproductive activity, copulation
time, fertilization, gestation period, and timing of postpartum estrus all tend to
cause asynchrony in a population. If an individual bat became pregnant at the
onset of copulatory activity and proceeded through the first two pregnancies with
little or no delay, there would be sufficient time for a third pregnancy in many
cases. I suspect that two per year is a more common occurrence.

At the other extreme from synchronized monestrous cycles are year-round
continuous reproductive cycles as exemplified by the vampire Desmodus rotun-
dus. Even here, it is likely that individual bats produce only two young per year
on the average, and asynchrony within populations gives the appearance of
continuous activity. In Colombia, for example, pregnant, lactating, and in¬
active Artibeus lituratus can be taken in any month of the year. Nevertheless,
there are peak periods of pregnancies in the months of December and May.

All of these patterns may be viewed as variations on a single theme. Given a
year’s time, what is the most efficient way to produce offspring? For animals
limited by the rigorous climates of the temperate zones, this results in a single,
population-wide effort at the time of maximum food availability. For tropical
species, it often might be possible to produce two litters during the favorable
period of food abundance, which is usually extended in tropical areas. Most
reproductive patterns in tropical areas seem to be correlated with seasonal rain¬
fall patterns. The dry season is probably the most stressful time of year for many
species, and reproductive strategies seem geared to avoid the weaning of young
during this season. In polyestrous species, the weaning of young from the first
birth peak is usually timed to coincide with the beginning of the rainy season, a
period of maximum food abundance. Desmodus rotundus has probably been
allowed by natural selection to adopt a year-round, asynchronous cycle due to the
year-round availability of its food source, blood from domestic cattle.

Species Accounts

The following accounts are arranged in the same order as the list of species
given by Jones and Carter (1976). Each account consists of a short summary or
discussion, to be used in conjunction with the listing of available data from the
literature presented in tabular form. Within the tables, localities, listed by state or

BIOLOGY OF THE PHYLLOSTOMATIDAE

319

country, are arranged from North to South, insofar as possible. Dates, listed by
month, are arranged chronologically from January to December, although varying
dates in a single reference necessitate some departure from the chronological
scheme. Numbers under the columns labeled “Pregnant”, “Lactating”, and “In¬
active,” refer to number of specimens. An “X” in these columns means the num¬
ber was not given. In the references column, “USNM” refers to records from the
National Museum of Natural History that have not been published previously.

The following list includes those species for which no data are available. I
emphasize these species here and in the species accounts in hopes that it will spur
efforts to gather such data: Micronycteris pusilla, Micronycteris behni, Lonchor-
hina orinocensis, Tonatia brasiliense, Tonatia carrikeri, Tonatia venezuelae,
Mimon bennettii, Mimon koepckeae, Phyllostomus latifolius, Lonchophylla
hesperia, Lonchophylla thomasi , Anoura werckleae , Scleronycteris ega, Licho-
nycteris degener, Platalina genovensium, Sturnira nana, Vampyrops aurarius,
Vampyrops nigellus, Vampyrops recifinus, Chiroderma doriae, Phyllops
falcatus, Phyllops haitiensis, Ariteus flavescens, Ametrida centurio, Sphae-
ronycteris toxophyllum, Brachyphylla nana , Phyllonycteris major.

Micronycteris megalotis

Data are insufficient from any one locality to speculate effectively on the sea¬
sonal reproductive pattern of M. megalotis. The data are not inconsistent with a
pattern of seasonal breeding in harmony with the rainfall pattern. In the northern
part of the range, females are pregnant during the beginning of the rainy season.
In the southern part of the range, however, they become pregnant earlier in the
year and the rainy or breeding season may last longer, possibly including two
breeding cycles per female per year. This may be due to an earlier and longer
lasting rainy season in the southern portions of the range. See Table 1 .

Micronycteris schmidtorum

The only reference to this species appears to be that of Mares and Wilson
(1971), who reported a male with nonscrotal testes taken in February in Costa
Rica.

Micronycteris minuta

Data available (Table 1) fit a pattern of breeding initiated at the beginning of
the rainy season. Confirmation of this pattern must await information relating to
other seasons. See Table 1.

Micronycteris hirsuta

Trinidad dates (Table 1) are from the appropriate times of year suggesting at
least a bimodal reproductive pattern. Lack of data from later in the year pre¬
cludes further speculation.

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Table 1. — Reproductive data for the genus Micronycteris.

Place

Date

Pregnant Lactating

Inactive

Reference

Micronycteris megalotis

Veracruz

Feb

Hall and Dalquest, 1963

Dec

Jun

Lackey, 1970

Yucatan

Apr

Jones et al., 1973

May

Birney et aL, 1974

Michoacan

May

Villa-R., 1966

Oaxaca

May

El Salvador

Mar

Burt and Stirton, 1961

Nicaragua

Mar

Jones et al., 1971a

Apr

Jun

1 2

Aug

Costa Rica

Feb

Gardner et al., 1970

Panama

May

Enders, 1935

Trinidad

Feb

Goodwin and Greenhall, 1961

Mar

Jun

Colombia

Jun

Thomas, 1972

Venezuela

Jul

USNM

Aug

Peru

Aug

Tuttle, 1970

Brazil

Jun

Peracchi and Albuquerque, 1971

Micronycteris minuta

Costa Rica

Mar

Gardner et al., 1970

Trinidad

Mar

Goodwin and Greenhall, 1961

May

1 4

Peru

Jul

Tuttle, 1970

Micronycteris hirsuta

Trinidad

Mar

Goodwin and Greenhall, 1961

May

Peru

Jul

Tuttle, 1970

Micronycteris sylvestris

Nayarit

Mar

Jones, 1964 ft

Veracruz

Dec

Hall and Dalquest, 1963

French Guiana

Feb

Brosset and Dubost, 1967

Mar

Micronycteris brachyotis

Goodwin and Greenhall (1961) reported a “breeding male” in May and three
others in June from Trinidad. Rick (1968) found one pregnant and six lactating
females in July in Guatemala.

Micronycteris pusilla

Nothing is known about reproduction in M. pusilla.

BIOLOGY OF THE PHYLLOSTOMATIDAE

321

Micronycteris nicefori

The only records are those of Goodwin and Greenhall (1961), reporting two
“breeding males” from Trinidad in October, and Baker and Jones (1975), re¬
porting a lactating female from Nicaragua in July.

Micronycteris sylvestris

In the northern part of the range, known records are from late in the rainy
season, whereas from the southern portion they are from early in the rainy season.
Data from other times of the year are necessary before speculating further. See
Table 1.

Micronycteris behni

Nothing is known about the reproductive pattern of M. behni.

Micronycteris daviesi

Tuttle (1970) collected a pregnant female in August in Peru. This is ap¬
parently the only record of reproductive activity for this species.

Macrotus waterhousii

In Mexico, M. waterhousii probably has a single young per year (Table 2).
The available evidence suggests May as the most likely period for parturition.
Additional study may reveal a delayed development system such as that de¬
scribed for the congeneric M. californicus in the following account. Data from the
Caribbean populations are insufficient for any meaningful analysis.

Macrotus californicus

In addition to the information in Table 2, Bradshaw (1961, 1962) has de¬
scribed the reproductive strategy of M. californicus in southern Arizona. A good
summary of the reproductive pattern also may be found in Anderson (1969).
Males undergo spermatogenesis in summer and autumn and inseminate females in
autumn; ovulation and fertilization occur immediately following copulation. The
single embryo undergoes slow growth during winter until March, when develop¬
ment proceeds at a more rapid rate resulting in a gestation period of about eight
months. Bradshaw (1961) coined the term “delayed development” to describe
the reproductive pattern. Parturition occurs in June and young are foraging by
August. Young-of-the-year females apparently breed during the first autumn,
but males are not reproductively mature until the following year.

Lonchorhina aurita

The little evidence available points to a breeding season that is correlated with
the beginning of the rainy season (Table 2). Panamanian pregnancies are during
the dry season and should result in the young being born at the beginning of the
rainy season.

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Table 2. — Reproductive data for the genera Macrotus, Lonchorhina, Macrophyllum,

Tonatia, andMimon.

Place Date Pregnant Lactating Inactive Reference

Macrotus waterhousii

Sinaloa

Jul

Jones et al., 1972

Jalisco

Feb

Watkins et al., 1972

Mar

May

X X

Jul

Sep

Oct

Tres Marias Is.

May

Merriam, 1898

Durango

Jun

2 1

Jones, 1964 c

Jamaica

Dec

7 2

Osburn, 1865

Dec

Goodwin, 1970

Crooked Is.

Apr

Buden, 1975

Cuba

Mar

Anderson, 1969

Caicos Is.

Feb

Buden, 1975

Apr

Macrotus californicus

Miller, 1904

California

Mar

Cockrum, 1955

Apr

USNM

Apr

Grinnell, 1918

May

Huey, 1925

Baja Calif.

Jul

Jones et al., 1965

Sonora

Apr

Burt, 1938

May

Jul

Aug**

Mar

Cockrum and Bradshaw, 1963

Apr

Lonchorina aurita

Quintana Roo

Aug

Jones et al., 1973

Oaxaca

Feb

Walker, 1975

Mar

Schaldach, 1965

Guatemala

Jan

Jones, 1966

Panama

Feb

Bloedel, 1955

Mar

Feb

Fleming et al., 1972

Mar

Nov

Trinidad

Apr

Goodwin and Greenhall, 1961

Peru

Jul

Tuttle, 1970

Aug

BIOLOGY OF THE PHYLLOSTOMATIDAE

323

Table 2. — Continued.

Macrophyllum macrophyllum

El Salvador

Oct

Harrison, 1975

Costa Rica

Mar

LaVal, 1977

May

Aug**

French Guiana

Oct

Brosset and Dubost, 1967

Nov

Tonatia bidens

Guatemala

Feb

Carter et aL, 1966

Honduras

Aug**

Valdez and LaVal, 1971

Costa Rica

Jan

Gardner et aL, 1970

Aug**

LaVal, 1977

Trinidad

May

Goodwin and Greenhall, 1961

Peru

Apr

Gardner, 1976

Jul

Tonatia minuta

Honduras

Aug

Valdez and LaVal, 1971

Nicaragua

Jul

Costa Rica

Feb

LaVal, 1977

Apr

Panama

Feb

Tonatia silvicola

Davis et al., 1964

Panama

Mar

Fleming et al., 1972

Oct

Nov

Dec

Colombia

Jan

Thomas, 1972

Peru

Jul

Tuttle, 1970

Aug

Mimon cozumelae

Veracruz

Apr

Hall and Dalquest, 1963

Yucatan

Apr

Jones et al., 1973

Jul

Campeche

May

Guatemala

Mar

Rick, 1968

Aug

1 1

Honduras

Jul

Valdez and LaVal, 1971

Costa Rica

Apr

LaVal, 1977

Aug

Mimon crenulatum

Campeche

Feb

Jones, 19646

Costa Rica

Apr

LaVal, 1977

Venezuela

Mar

Goodwin and Greenhall, 1961

Peru

Jul

Tuttle, 1970

•One with twins.
••Young taken.

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SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

Lonchorhina orinocensis

Nothing is known about reproduction in L. orinocensis.

Macrophyllum macrophyllum

Felten (1956a) postulated that this species breeds in the dry season. The
finding of pregnant animals during the late rainy season in French Guiana is
unusual when compared with the cycles in other members of the subfamily. See
Table 2.

Tonatia bidens

Although the records are scattered, I suspect that this species breeds more than
once a year (Table 2). Records from Honduras suggest a bimodal pattern with
subadult animals representing the earlier breeding cycle.

Tonatia brasiliense

Nothing is known about the reproductive pattern of this species.

Tonatia carrikeri

Nothing is known about the reproductive pattern of T. carrikeri, although
Gardner (1976) reported two reproductively inactive females from Peru in July.

Tonatia minuta

This species also appears to fit the bimodal pattern, although additional data
are obviously necessary to confirm this hypothesis. See Table 2.

Tonatia silvicola

Females appear to give birth during the early half of the rainy season; there is
thus far no evidence of more than one young per year. See Table 2.

Tonatia venezuelae

No information is available on reproduction in this species.

Minion bennettii

Nothing is known about the reproductive pattern of this bat.

Mimon cozumelae

This species (Table 2) apparently produces young at the beginning of the rainy
season and the available data suggest only a single young per year.

BIOLOGY OF THE PHYLLOSTOMATIDAE

325

Mimon crenulatum

Records from Campeche and Venezuela are from the dry season, whereas
Peruvian records are from the rainy season. The single record from Costa Rica
was taken in the period of transition between dry and rainy seasons. See Table 2.

Mimon koepckeae

No data are available on the reproductive pattern of this species.

Phyllostomus discolor

In addition to records listed in Table 3, Mares and Wilson (1971) found 80
per cent of 43 animals in 1968 and 51 per cent of 69 animals in 1970 to be re-
productively active during February and March in Costa Rica. Tamsitt (1966)
stated that in Colombia this species is acyclic or continuous in its breeding habits.
Most of the above data suggest this pattern for other areas as well; however, the
lack of reproductive activity as noted by Fleming et al. (1972) for Costa Rica
seems unusual. Heithaus et al. (1975) suggested that P. discolor may be mon-
estrous in Costa Rica.

Phyllostomus hastatus

Starrett and de la Torre (1964) reported that one of two July-taken males
from Costa Rica had small, inguinal testes, and the other had large, scrotal testes;
both were in an early stage of spermatogenesis with no mature sperm in the testes.

The available data could support either a monestrous (in Nicaragua, Panama,
and Trinidad) or polyestrous (in Colombia) pattern. In fact, this may be a
species in which the reproductive strategy varies geographically. See Table 3.

Phyllostomus elongatus

Additional data from times of the year other than those listed in Table 3 are
needed to elucidate the pattern of this species. The above data show that these
animals breed during the middle part of the rainy season.

Phyllostomus latifolius

Nothing is known about the reproductive pattern of this species.

Phylloderma stenops

The only report of reproductive activity for this rare species is that of LaVal
(1977), who reported a pregnant female in February (embryo length, 33 mm.)
from Costa Rica. Gardner (1976) reported a reproductively inactive female from
Peru that was collected in May.

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SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

Table 3. — Reproductive data for the genera Phyllostomus, Trachops, Chrotopterus, and

Vampyrum.

Place

Date

Pregnant Lactating

Inactive

Reference

Phyllostomus discolor

Guatemala

Mar

Jones, 1966

El Salvador

Feb

Felten, 1956 h

Jun

Aug

Sep

Nov

Dec

Burt and Stirton, 1961

Nicaragua

Mar

Jones, 1964a

Costa Rica

Jan

Fleming et al., 1972

Mar

Apr

May

Jul

Dec

Jul

Tamsitt and Valdivieso, 1961

Trinidad

Feb

Goodwin and Greenhall, 1961

Mar

Jun

Aug

Sep

Oct

Colombia

Feb*

Tamsitt and Valdivieso, 1964

Mar

May

Sep

Oct*

Venezuela

Jul

Smith and Genoways, 1974

Brazil

Jul

Walker, 1975

Phyllostomus hastatus

Nicaragua

Mar

Jones et al., 1971a

Jun

Jul

Aug

Panama

Apr

Fleming et al., 1972

May

Jun

Oct

Trinidad

Mar

Goodwin and Greenhall, 1961

Apr

Jun

Sep

Nov

Venezuela

Aug

USNM

Colombia

Mar

Thomas, 1972

May

Aug

BIOLOGY OF THE PHYLLOSTOMATIDAE

327

• Table 3. — Continued.

Colombia

Sep

2 1

Thomas, 1972

Oct

Nov

Dec

1 5

Jul

Arata and Vaughn, 1970

Peru

Jun

Tuttle, 1970

Aug

Brazil

Aug

Phyllostomus elongatus

Peracchi and Albuquerque, 1971

Colombia

Jun

Thomas, 1972

Peru

Jul

Tuttle, 1970

Aug

Trachops cirrhosus

Veracruz

Apr

Hall and Dalquest, 1963

Campeche

Feb

Jones et al., 1973

Oaxaca

Mar

Villa- R, 1966

Chiapas

Dec

Mar

Carter et al., 1966

Guatemala

Mar

Jones, 1966

Apr

El Salvador

Feb

Burt and Stirton, 1961

Honduras

Aug

Valdez and LaVal, 1971

Nicaragua

May

Carter et al., 1966

Costa Rica

Mar

Aug

1 6

Armstrong, 1969

Panama

Aug

Fleming et aL, 1972

Oct

Nov

Trinidad

Mar

Goodwin and Greenhall, 1961

Peru

Jul

Chrotopterus auritus

Tuttle, 1970

Veracruz

Apr

Hall and Dalquest, 1963

Yucatan

Apr

Jones et al., 1973

Jul

1 1

Argentina

Jul

Vampyrum spectrum

Villa-R. and Villa-C., 1969

Costa Rica

Aug

Gardner et aL, 1970

Trinidad

May

Goodwin and Greenhall, 1961

‘Pregnant and Iactating.

Trachops cirrhosus

Felten (1956a) stated that T. cirrhosus breeds in the dry season in El Salvador,
and the data of Burt and Stirton (1961) support this. This species may have an
extended season, or may be geographically variable with regard to the reproduc¬
tive cycle. Additional data on other seasons from any of the above localities would
be useful. See Table 3.

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SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

Chrotopterus auritus

Data are insufficient (Table 3) to allow speculation on the possible reproduc¬
tive pattern of C. auritus except to note that this species produces young during
the early part of the rainy season.

Vampyrum spectrum

The only information available other than that in Table 3 seems to be Green-
hall’s (1968) report on a birth in captivity. Again, other than the fact that V.
spectrum produces young during the rainy season, little can be said about its re¬
productive cycle.

Glossophaga soricina

Cockrum (1955) believed G. soricina to be polyestrous, with the young born
at any time of the year in Mexico. Fleming (1973) felt that this species is sea¬
sonally polyestrous in Panama, with bimodal birth peaks occurring in March-
April and July- August. Tamsitt (1966) indicated that G. soricina is acyclic or
continuously breeding in Colombia. Felten (1956a) noted that this species breeds
throughout the year in El Salvador. Heithaus et al. (1975) suggested bimodal
polyestry for Costa Rican animals.

This is one of the few species of phyllostomatid bats for which a fair amount of
reproductive data are available from a variety of localities (Table 4). The data
suggest G. soricina is polyestrous in most areas. Reproduction may be somewhat
geographically variable inasmuch as data from Panama indicate no pregnancies
during the period August-December. Also, in some of the areas where these bats
appear to breed continuously, there may well be a bimodal pattern for individuals
but enough asynchrony within the population to allow for individuals in all stages
of the reproductive cycle to be collected at any given time.

Rasweiler (1972) demonstrated this species to be polyestrous with approxi¬
mately a 24-day cycle in captivity. He described the preimplantation development
and histology of the oviduct in some detail.

Glossophaga alticola

I can find no reproductive information for G. alticola in the literature. The
National Museum of Natural History has two specimens taken in Oaxaca in
April, one of which was pregnant and the other inactive.

Glossophaga commissarisi

The data available (Table 4) are not inconsistent with a pattern of bimodal
polyestry. Although the data are scanty, the information from Jalisco supports
this hypothesis.

Glossophaga longirostris

This species appears to breed during the rainy season, but the data are in¬
conclusive (Table 4).

BIOLOGY OF THE PHYLLOSTOMATIDAE

329

Table 4. — Reproductive data for the gem/s Glossophaga.

Place

Date

Pregnant

Lactating

Inactive

Reference

Glossophaga soricina

Sonora

May

Cockrum, 1955

Dec

Cockrum and Bradshaw, 1963

Chihuahua

Jul

Anderson, 1972

Durango

Jun

Jones, 1964 c

San L. Potosi

Jun

Dalquest, 1953

Sinaloa

Jan

Jones et ai, 1972

Mar

May

Aug

Sep

Oct

Nov

Dec

Nayarit

Jan

Cockrum, _1 955

Feb

Aug

Jalisco

Feb

Watkins et al„ 1972

Mar

Apr

Sep

Oct

Tres Marias Is.

May

Merriam, 1898

Colima

Nov

Villa-R., 1966

Dec

Queretaro

Jan

Schmidly and Martin, 1973

Dec

Puebla

Jan

LaVal, 1972

Veracruz

Mar

Hall and Dalquest, 1963

Apr

Sep

Nov

Jun

Lackey, 1970

Jul

Tabasco

May

Villa-R., 1966

Jun

Jul

Yucatan

Feb

X**

x**

Jones et al., 1973

Apr

X* *

X**

Jul

X**

Apr

Birney et al., 1974

Aug

Pearse and Kellogg, 1938

Oaxaca

Mar

USNM

Apr

Sep

Cockrum, 1955

Chiapas

Feb

Villa-R., 1966

Aug

X***

Barlow and Tamsitt, 1968

Guatemala

Mar

Jones, 1966

Aug

330

SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

Table 4. — Continued.

El Salvador

Jan

Felten, 1956ft

Feb

Mar

Apr

Jun

Jul

Aug

Sep

Oct

Nov

Dec

Sep

Burt and Stirton, 1961

Nov

Jul

Starrett and de la Torre, 1964

Honduras

Aug

Costa Rica

Jul

Aug

Jul

Tamsitt and Valdivieso, 1961

Aug

Panama

Jan

Fleming et at., 1972

Feb

Mar

Apr

May

Jun

Jul

Aug

Sep

Oct

Nov

Dec

Feb

Bloedel, 1955

Jamaica

Jan

Goodwin, 1970

Trinidad

Jan

Goodwin and Greenhall, 1961

Feb

Mar

Apr

May

Jun

Dec

Venezuela

Aug

USNM

Colombia

Jan

Thomas, 1972

Feb

Mar

Apr

May

Jun

Jul

Aug

Sep

Oct

Nov

BIOLOGY OF THE PHYLLOSTOMATIDAE

331

Table 4. — Continued.

Colombia

Dec

Thomas, 1972

Jan

Tamsitt and Valdivieso, 1964

Mar

Apr

Jun

Jul

Aug

Sep

Oct

Nov

Dec

Jul

Arata and Vaughn, 1970

Aug

French Guiana

Feb

Brosset and Dubost, 1967

Mar

Oct

Peru

Jun

Tuttle, 1970

Jul

Aug

Brazil

Nov

Hamlett, 1935

Dec

Jan

Peracchi and Albuquerque, 1971

Paraguay

Oct

USNM

Glossophaga commissarisi

Durango

Jul

Baker and Greer, 1962

Sinaloa

Jan

Jones et ai, 1972

Jul

Jalisco

Feb

Watkins et al„ 1972

Apr

May

Jul

Sep

Nov

Dec

Guatemala

Feb

Jones, 1966

Nicaragua

Feb

Jones, 1964a

Glossophaga longirostris

Trinidad

Feb

Goodwin and Greenhall, 1961

Mar

Apr

Aug

Sep

Jun

Goodwin, 1958

Venezuela

Jul

USNM

Jul

Smith and Genoways, 1974

•Parturition.
••Pregnant or lactating.
•••Twins.

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SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

Monophyllus redmani

Goodwin (1970) felt that the high percentage of pregnancies in his sample
suggested a discrete breeding season for this species. Additional data from other
times of the year are needed in order to verify his opinion. See Table 5.

Monophyllus plethodon

Schwartz and Jones (1967) reported pregnant females from Dominica in
March and April, perhaps indicating a distinct breeding season for this species.

Leptonycteris nivalis

Davis (1966) reported that the breeding season is restricted to April, May,
and June. Easterla (1972) felt that young probably are born in Mexico, possibly
in June, prior to the time bats arrive in the Big Bend area of Texas. Records from
Veracruz (Table 5) indicate a second pregnancy of the year for this migratory
species.

Leptonycteris sanborni

Cockrum and Ordway (1959) and Hayward and Cockrum (1971) have re¬
ported on reproduction in L. sanborni in Arizona. They found that pregnant fe¬
males arrive in southern Arizona in early May and the young are bom shortly
thereafter. By August there are subadult females containing embryos 10 mm. in
crown-rump length; however, all bats have left for Mexico by the early part of
October. They hypothesized another birth peak in Mexico in early November.
In January females have small embryos and in February they begin to move to
the northern part of their range where the young will be bom. Hayward and Cock¬
rum (1971) suggested, as an alternative hypothesis, that delayed development as
described for Macrotus waterhousii might be involved. See Table 5.

Leptonycteris curasoae

Smith and Genoways (1974) found a large colony of this species on Margarita
Island, Venezuela, which in July was estimated to contain 4000 females nursing
nearly full-grown young. In November, seven of 34 females examined were preg¬
nant, and no juveniles were present. In addition, adult males with large (6 to 8
mm.) testes were present in November, whereas males had been absent in July.
This appears to be the only record of reproduction for this species.

Lonchophylla Hesperia

Nothing is known about the reproductive pattern of this species.

Lonchophylla mordax

Thomas (1972) collected a reproductively inactive female of this species in
Colombia in January. I can find no other literature records relating to repro¬
duction in L. mordax.

BIOLOGY OF THE PHYLLOSTOMATIDAE

333

Lonchophylla concava

Although there are few reports giving reproductive condition (Table 5), these
are from sufficiently distinct times of the year as to suggest the possibility of more
than one birth peak per year.

Lonchophylla robusta

Apparently, reproductively active individuals of this species have not been
recorded. The capture of inactive females in several months of the year (Table 5)
suggests an asynchronous reproductive cycle.

Lonchophylla thomasi

No data are available concerning the reproductive pattern of this species.

Lionycteris spurrelli

The only published record of reproductive activity in L. spurrelli is that of
Tuttle (1970), who reported a pregnant female taken in August in Peru.

Anoura geoffroyi

Alvarez and Ramirez-Pulido (1972) netted 12 males but no females at the
mouth of a cave in Michoacan and suggested that this species may form sexually
segregated colonies. The data of Goodwin and Greenhall (1961) from Trinidad
support this notion for certain times of the year. They reported 20 males and 25
females in June; 29 males and one female in October; and 32 males and 56 fe¬
males in November — all from the same caves. Their data also recommend a
discrete breeding season occurring late in the rainy season, a rather unusual pat¬
tern for phyllostomatids. See Table 5.

Anoura caudifer

Pregnancy records (Table 5) from several months throughout the year suggest
an asynchronous reproductive cycle. Additional data from other months of the
year would be useful in discerning the true reproductive patterns.

Anoura cultrata

Gardner et al. (1970) reported a pregnant female from Costa Rica taken in
August. They also collected males in February, May, and July and gave testicular
measurements.

Anoura werckleae

Nothing is known about the reproductive pattern of this species.

Anoura brevirostrum

When Carter (1968) described A. brevirostrum from Peru, he included data
from a lactating female and two inactive females taken in August.

334

SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

Table 5. — Reproductive data for the genera Monophyllus, Leptonycteris, Lonchophylla,
Anoura, Lichonycteris, Hylonycteris, Choeroniscus, and Choeronycteris.

Place

Date

Pregnant Lactating Inactive

Reference

Jamaica

Feb

Monophyllus redmani

Osburn, 1865

Feb

6 4

McNab, 1976

Jan

Goodwin, 1970

Dec

15 4

Caicos Is.

Jan

Homan and Jones, 1975

Hispaniola

Feb

Dec

Puerto Rico

Feb

Texas

Jun

Leptonycteris nivalis

Easterla, 1972

Coahuila

Jul

Baker, 1956

Tamaulipas

Aug

Alvarez, 1963

Veracruz

Sep

Hall and Dalquest, 1963

Arizona

Aug

Leptonycteris sanborni

Hoffmeister and Goodpaster, 1954

Sonora

Mar

Cockrum and Bradshaw, 1963

Apr

Sinaloa

Feb

Jones et ai, 1972

Jul

Nov

Jalisco

Jan

Watkins et al., 1972

Jul

Oct

Morelos

Sep

1 2

Villa-R., 1966

Mexico

Nov

1 2

Costa Rica

Mar

Lonchophylla concava

1 1

Davis et al., 1964

Aug

Gardner et al., 1970

Costa Rica

Mar

Lonchophylla robusta

Mares and Wilson, 1971

Colombia

Mar

Thomas, 1972

Apr

Jul

Sep

Peru

Aug

Tuttle, 1970

Zacatecas

Jun

Anoura geoffroyi

Matson and Patten, 1975

Sinaloa

Jul

Jones et al., 1972

Colima

Nov

Villa-R., 1966

Dec

Guerrero

Sep

Oaxaca

Jul

Baker and Womochel, 1966

Nov

Schaldach, 1966

Dec

BIOLOGY OF THE PHYLLOSTOMATIDAE

335

Nicaragua
Costa Rica
Trinidad
Peru

Colombia

French Guiana
Brazil

Guatemala
Costa Rica

Jalisco

Tabasco

Oaxaca

Guatemala
Costa Rica

Sinaloa

Oaxaca

Honduras

Nicaragua

Costa Rica

Trinidad

Peru

Table 5. — Continued.

Jul

Jones et aL, 1971 a

Mar

Mares and Wilson, 1971

Nov

Goodwin and Greenhall, 1961

Jun

Tuttle, 1970

Anoura caudifer

Mar

Thomas, 1972

May

Nov

Jun

Tamsitt and Valdivieso, 1966a

Jan

Brosset and Dubost, 1967

Feb

Kuhlhorn, 1953

Jan

Lichonycteris obscura

Peracchi and Albuquerque, 1971

Feb

Carter et al„ 1966

Jan

Gardner et aL, 1970

Mar

Hylonycteris underwood i

Jul

Phillips and Jones, 1971

Sep

May

Villa-R., 1966

Nov

1 1

Jul

Baker and Womochel, 1966

Mar

Carter et al., 1966

Jan

LaVal, 1977

Feb

Mar

Apr

Gardner et aL, 1970

May

Jun

Jul

Aug

LaVal, 1972

Oct

Nov

Choeroniscus godmani

Jul

Jones, 19646

May

Schaldach, 1965

Jul

Valdez and LaVal, 1971

Mar

Jones et al., 1971a

Apr

Mar

Mares and Wilson, 1971

Choeroniscus intermedius

Aug

Goodwin and Greenhall, 1961

Jul

Tuttle, 1970

336

SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

Table 5. — Continued.

Choeronycteris mexicana

Arizona

Jun

Campbell, 1934

Jun

Barbour and Davis, 1969

Jun

Walker, 1975

Jul

Aug

Hoffmeister and Goodpaster, 1954

New Mexico

Jun

Mumford and Zimmerman, 1962

Jun

Mumford et al., 1964

Coahuila

Mar

Baker, 1956

Jun

Aug

Sep

Jun

Axtell, 1962

Tamaulipas

Aug

Alvarez, 1963

Sonora

Jul

Villa-R., 1966

Sinaloa

Feb

Jones et al., 1972

Jalisco

Jan

Watkins et al., 1972

Feb

Mar

Sep

Oct

Guerrero

Feb

Villa-R., 1966

Scleronycteris ega

Nothing has been recorded about reproduction in this species.

Lichonycteris degener

Nothing is known about the reproductive pattern of L. degener.

Lichonycteris obscura

This species is reproductively active during the dry season in Middle America
(Table 5), but until data are available from other months of the year, little can
be said of the overall pattern.

Hylonycteris underwoodi

The data from Costa Rica (Table 5) fit the bimodal pattern common to many
other species. The second birth peak appears to be later in the rainy season than
for some other species.

Platalina genovensium

Nothing is known about reproduction in this species.

BIOLOGY OF THE PHYLLOSTOMATIDAE

337

Choeroniscus godmani

Choeroniscus godmani (Table 5) seems to fit the usual pattern of weaning
young during the early part of the rainy season, but the lack of data from later
in the year makes this conclusion tentative.

Choeroniscus minor

The only apparent report of reproductive activity for this species is that of
Tamsitt et al. (1965), who reported a lactating female from Colombia in Decem¬
ber. Tuttle (1970) collected a juvenile in August in Pern.

Choeroniscus intermedius

The data in Table 5 are too few to provide much insight into the reproductive
pattern of this species.

Choeroniscus inca

Goodwin and Greenhall (1961) noted a pregnant female taken in February in
Trinidad.

Choeroniscus periosus

The only record is that of Thomas (1972), who captured two lactating females
in Colombia in January.

Choeronycteris mexicana

These animals are pregnant in the early spring in Mexico (Table 5), and those
that migrate to Arizona and New Mexico give birth in June. The possibility of a
second period of parturition, as suggested for Leptonycteris sanborni, is supported
by the pregnancy record in September from Jalisco.

Musonycteris harrisoni

There are no published records of reproductive activity for this species, but
Alfred L. Gardner has kindly made available to me his unpublished field notes,
which record one inactive and two pregnant females taken in September in
Colima.

Carollia castanea

Pine (1972) suggested that C. castanea is polyestrous, but cautioned that in any
one locality there may be one or two more or less fixed seasons. This caveat is
supported by Fleming (1973), who suggested that in Panama C. castanea is
bimodally polyestrous, with birth peaks occuring in March-April and July-August.
Thomas’ (1972) data show that females are pregnant during the period September-
November in Colombia corresponding to a period of reproductive quiescence in

338

SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

Panama (Fleming, 1973). These differences probably reflect contrasts in the sea¬
sonality of the rainfall patterns at the different localities. See Table 6.

Carollia subrufa

Felten (1956a) suggested that C. subrufa breeds both in the dry and wet
seasons in El Salvador. Pine (1972) felt that they either breed throughout the
year or that possibly there is a period of inactivity in the early winter months, at
least in some areas. The data from El Salvador, the most extensive for any one
area, would seem to fit a bimodal pattern (Table 6).

Carollia brevicauda

Pine (1972) suggested that C. brevicauda breeds from midwinter to early
spring. Three records of females both pregnant and lactating (Table 6) attest to
the presence of polyestry in Central America. This species may exhibit the bi¬
modal type of breeding season seen for other Central American phyllostomatids;
however, data from late in the year are needed for clarification of this pattern.

Carollia perspicillata

Fleming (1973) and Heithaus et al. (1975) have shown that C. perspicillata
fits the model of bimodal polyestry, and the data summarized here support this
contention (Table 6). Birth peaks occur in the periods February-May and June-
August in Panama, and somewhat earlier in other areas, depending on seasonal
rainfall patterns in the various localities. Several of the data sets from various
localities show a distinct drop in reproductive activity during the latter part of
the rainy season, usually in the period from October to December, but earlier in
Colombia. Fleming et al. (1972) correlated testis size with spermatogenic
activity and found that males had large testes just preceding those times when
females were likely to be sexually active.

Rhinophylla pumilio

The data in Table 6 are too few to warrant speculation on the reproductive
pattern of R. pumilio.

Rhinophylla alethina

Although the sample (Table 6) is admittedly small, the timing of the repro¬
ductive events recorded here suggests an extended or possibly asynchronous
breeding season. Data from August-November would be useful for clarifying the
pattern.

Rhinophylla fischerae

The lack of reproductive activity for animals taken in July and August seems
striking when compared against what is known for other phyllostomatids. See
Table 6.

BIOLOGY OF THE PHYLLOSTOMATIDAE

339

Table 6. — Reproductive data for the genera Carollia and Rhinophylla.

Place

Date

Pregnant

Lactating Inactive

Reference

Carollia castanea

Honduras

May

Pine, 1972

Jul

Nicaragua

Feb

Jones et al., 1971a

Mar

Apr

Jun

Jul

Aug

Costa Rica

Feb

Mar*

Aug

Panama

Jan

Feb

Mar

Jan

Fleming et al., 1972

Mar

Apr

Jun

Jul

Aug

Oct

Nov

Dec

Colombia

Jan

2 5

Thomas, 1972

Feb

Mar

Apr

1 7

May

Jun

Jul

Sep

Oct

Nov

Dec

French Guiana

Jan

Brosset and Dubost, 1967

Feb

Mar

Peru

Jul

Tuttle, 1970

Aug

Carollia subrufa

Puebla

Jun

LaVal, 1972

Guerrero

May

Pine, 1972

Dec

Oaxaca

May

Villa-R., 1966

Chiapas

Feb

May

Pine, 1972

Jul

340

SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

Table 6. — Continued.

Chiapas

Aug

Oct

Nov

Guatemala

Feb

Nov

El Salvador

Jan

Feb

Mar

Sep

Oct

Nov

Dec

Honduras

Jul

Nicaragua

Jul

Aug

Panama

Jan

Feb

Mar

Dec

Carollia

brevicauda

San L. Potosi

Apr

Aug

Veracruz

Feb

Mar

Dec

Tabasco

Apr

May

Campeche

Jan

Quintana Roo

Apr

Aug

Oaxaca

Feb

Mar

Chiapas

Jun

Jul

Nov

Guatemala

Feb

Mar

Aug

Feb

Honduras

Apr

May*

Jun

Nicaragua

Jul*

Costa Rica

Mar

Apr*

Panama

Jan

Feb

Mar

Ecuador

Mar

Peru

Aug

Oct

Pine, 1972

Felten, 1956c

Pine, 1972

Walker, 1975

Pine, 1972

Jones, 1966
Rick, 1968
Pine, 1972

BIOLOGY OF THE PHYLLOSTOMATIDAE

341

Table 6. — Continued.

Carollia perspicillata

Puebla

Jan

LaVal, 1972

Veracruz

May

Villa-R., 1966

Jun

Lackey, 1970

Jul

Campeche

May

Jones et al., 1973

Jul

Pine, 1972

Quintana Roo

Jul

Jones et al., 1973

Aug

Oaxaca

Apr

Hahn, 1907

Chiapas

Aug

Pine, 1972

Guatemala

Mar

Jones, 1966

El Salvador

Mar

Burt and Stirton, 1961

Apr

Felten, 1956c

Oct

Nov

Dec

Mar

Pine, 1972

Honduras

May

Jun

Jul

Nicaragua

Feb

Apr

May

Jun

Jul

Aug

Costa Rica

Feb

Mar

Apr

Jul

Aug

Jan

Fleming et al., 1972

Feb

Mar

Apr

May

Jun

Jul

Aug

Sep

Oct

Nov

Panama

Jan

Feb

Mar

Apr

May

Jun*

Jul

342

SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

Table 6. — Continued.

Panama

Trinidad

Venezuela
French Guiana

Colombia

Ecuador

Aug

Sep

Oct

Nov

Dec

Mar

May

Feb

Mar

Apr

Jun

Feb

Mar

Apr

May

Jun

Jul

Aug

Sep

Oct

Jun

Jul

Aug

Sep

Oct

Nov

Jul

Aug

Sep

Jan

Mar

Apr

Oct

Jan

Feb

Mar

Apr

May*

Jun

Jul

Aug

Sep

Oct

Nov

Dec

Jun

Jul

Mar

Jul

Aug

Sep

Fleming et al., 1972

Enders, 1935

Hall and Jackson, 1953

Pine, 1972

Hahn, 1907

Goodwin and Greenhall, 1961

Pirlot, 1963

Brosset and Dubost, 1967

Arata and Vaughn, 1970

Tamsitt and Valdivieso, 1964

Thomas, 1972

Pine, 1972

Bolivia

BIOLOGY OF THE PHYLLOSTOMATIDAE

343

Table 6.— Continued.

Peru

Aug

Pine, 1972

Jun

Tuttle, 1970

Jul

Aug

Brazil

Jan

Peracchi and Albuquerque, 1971

Sep

Oct

Rhinophylla pumilio

Venezuela

Dec

1 1

Walker, 1975

Peru

Jun

Tuttle, 1970

Jul

Colombia

Jan

Thomas, 1972

Apr

May

Jun

1 2

Jul

Dec

Rhinophylla fischerae

Peru

Jul

Tuttle, 1970

Aug

Carter, 1966

*Pregnant and lactating.

Sturnira lilium

Jones (1966) and Jones et al. (1973) suggested that S. lilium probably breeds
through the year. Actually, the data presented in Table 7 support the model of
bimodal polyestry as suggested by Fleming et al. (1972). Support for this model
is provided by the data from Costa Rica and Colombia. In Costa Rica, birth peaks
occur in February-March and in June-July (Heithaus et al., 1975); in Colombia,
there appears to be much less synchrony in the cycle.

Sturnira thomasi

Genoways and Jones (1975) reported two lactating females, a subadult, and
a juvenile from Guadeloupe in July. This seems to be the only record of repro¬
duction available for this species.

Sturnira tildae

The records listed in Table 7 provide no basis for speculation on the repro¬
ductive habits of S. tildae.

Sturnira magna

Tuttle (1970) provided testicular measurements on three Peruvian males
taken in July. Gardner (1976) took one inactive and one lactating female in
May and another inactive female in July in Peru.

344

SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

Table 7. — Reproductive data for the gcmrsSturnira.

Place

Date

Pregnant

Lactating

Inactive

Reference

Sturnira lilium

Sonora

Sep

Findley and Jones, 1965

Sinaloa

Apr

Cockrum and Bradshaw, 1963

May

Jones et al., 1972

Jun

Aug

Durango

Jun

Jones, 1964 c

Jul

Baker and Greer, 1962

Jalisco

Jan

Watkins et al., 1972

Mar

Apr

Jun

Jul

Aug

Sep

Oct

Nov

Queretaro

Jan

Spenrath and LaVal, 1970

Puebla

Jan

LaVal, 1972

Veracruz

Jun

Lackey, 1970

Jul

Campeche

Jan

Jones et al., 1973

Jul

Quintana Roo

Aug

Apr

Birney et al., 1974

Oaxaca

Apr

USNM

Jul

Baker and Womochel, 1966

Dec

Schaldach, 1966

Chiapas

May

Villa-R., 1966

Jun

Guatemala

Feb

Jones, 1966

Mar

Jun

Jul

Aug

May

Rick, 1968

El Salvador

Jul

Starrett and de la Torre, 1964

Nicaragua

Jul

Costa Rica

Jan

Fleming et aL, 1972

Feb

Mar

Apr

May

Jun

Jul

Aug

Dec

BIOLOGY OF THE PHYLLOSTOMATIDAE

345

Dominica

Martinique
St. Lucia
St. Vincent
Colombia

French Guiana

Peru

Brazil

Trinidad

Peru

Brazil

Costa Rica

Jalisco

Colima

Queretaro

Puebla

Table 7. — Continued.

Mar

Apr

Aug

1 4

Mar

Aug

Jul

3 4

Aug

1 13

Jan

Feb

Mar

Apr

1 2

May

2 3

Jun

1 3

Jul

2 2

Aug

2 1

Sep

4 4

Oct

3 2

Nov

Dec

1 3

Jun

Jul

Aug

Jun

Jul

Aug

Sturnira tildae

Mar

Jul

Jun

Jul

Sturnira mordax

Feb

May

Aug

Sturnira ludovici

Apr

May

Jul

Aug

1 13

Sep

Nov

Dec

Nov

Sep

Nov

Jan

Jones and Phillips, 1976

Arata and Vaughn, 1970

Thomas, 1972

Brosset and Dubost, 1967

Tuttle, 1970

USNM

Peracchi and Albuquerque, 1971

Goodwin and Greenhall, 1961

Tuttle, 1970

USNM

Gardner et al., 1970

LaVal, 1977
Armstrong, 1969

Watkins et al., 1972

Jones and Phillips, 1964
Villa-R., 1966

Schmidly and Martin, 1973
LaVal, 1972

346

SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

Table 7. — Continued.

Oaxaca

Jul

Baker and Womochel, 1966

Chiapas

Aug

Villa-R., 1966

Costa Rica

Jul

Starrett and de la Torre, 1964

Colombia

Jan

Thomas, 1972

Feb

Mar

Apr

May*

9 6

Jun

Jul

1 1

Aug

Sep

Oct

Nov

Dec

Peru

Jun

Tuttle, 1970

Sturnira erythromos

Colombia

May

Thomas, 1972

Dec

Peru

Jun

Gardner and O’Neill, 1969

Aug

*Pregnant and lactating.

Sturnira mordax

The presence of pregnant females both in February and in August suggests
polyestry for S. mordax (Table 7).

Sturnira bidens

The only published record of reproductive activity for this species is that of
Gardner and O’Neill (1969), who reported three pregnant females and one in¬
active female from Peru in August.

Sturnira nana

Nothing is known about the reproductive pattern of S. nana.

Sturnira aratathomasi

Thomas and McMurray (1974) reported pregnant females from Colombia in
February and August. These pregnancy dates are not inconsistent with those for
other, more common species of the genus Sturnira and may represent the familiar
bimodal pattern.

Sturnira ludovici

Sturnira ludovici appears to me to be another species with a bimodal polyes-
trous pattern (Table 7). The data from Colombia are strikingly similar to those

BIOLOGY OF THE PHYLLOSTOMATIDAE

347

presented for S. lilium. Starrett and de la Torre (1964) presented data on testis
size and spermatogenesis from two males from Costa Rica.

Sturnira erythromos

Speculation on the reproductive pattern of S. erythromos must await further
data. See Table 7.

Uroderma bilobatum

Davis (1968) suggested that U. bilobatum seemingly lacks a restricted breed¬
ing season based on his examination of 58 females from a variety of localities
from Oaxaca to Venezuela. Of these, three were pregnant in January, five in
February, and one each in May, July, and November (Table 8). Fleming (1973)
pointed out that in Panama this species is another example of bimodal polyestry
and much of the above data are in agreement with that conclusion. Again, the in¬
formation from Colombia shows that the timing of reproductive peaks is quite
different from that in Panama, with the second major pregnancy period in Colom¬
bia occurring in the late rainy season. Fleming et al. (1972) presented data on
testis size and spermatogenesis, showing that males undergo active spermato¬
genesis in a bimodal fashion also.

Uroderma magnirostrum

Although the data are few and from widely scattered localities (Table 8), I
suspect U. magnirostrum will prove to have a polyestrous pattern like that of its
congener, U. bilobatum.

Vampyrops infuscus

The only report of reproduction in this species appears to be that of Marinkelle
(1970), who collected one pregnant female and three lactating females in
Colombia in March.

Vampyrops vittatus

Pregnancies occur in the early part of the rainy season in Costa Rica (Table 8),
but data from other seasons are lacking.

Vampyrops dorsalis

The Colombian data (Table 8) show V. dorsalis to fit the pattern of bimodal
polyestry common to several other species of Colombian phyllostomatids.

Vampyrops aurarius

No data are available about reproduction in this species.

Vampyrops nigellus

Nothing is known about the reproductive pattern of V. nigellus.

348

SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

Table 8. — Reproductive data for the genera Uroderma, Vampyrops, and Vampyrodes.

Place

Date

Pregnant

Lactating

Inactive

Reference

Uroderma bilobatum

Veracruz

Jun

Lackey, 1970

Jul

Chiapas

May

Villa-R., 1966

Aug

Guatemala

Feb

Jones, 1966

El Salvador

Jan

Felten, 1956a

Jan

Burt and Stirton, 1961

May

Honduras

Jul

Baker et aL, 1975

Nicaragua

Feb

Jones, 1964 a

Aug

Davis et aL, 1964

Panama

Jan

Davis, 1968

Mar

Bloedel, 1955

Jan

Fleming et al., 1972

Feb

Mar

Feb

Walker, 1975

Mar

Apr

Fleming et al., 1972

May

Jun

Jul

Aug

Sep

Oct

Nov

Dec

Trinidad

Feb

Goodwin and Greenhall, 1961

May

Colombia

Jan

Tamsitt and Valdivieso, 1964

Mar

Jul

Sep

Nov

Thomas, 1972

Peru

Aug

Tuttle, 1970

Brazil

Jul

USNM

Uroderma magnirostrum

El Salvador

Jun

Davis, 1968

Nicaragua

Mar

Jones et al., 1971a

Jul

Davis, 1968

Bolivia

Sep

Brazil

Jun

USNM

BIOLOGY OF THE PHYLLOSTOMATIDAE

349

Table 8. — Continued.

Vampyrops vittatus

Costa Rica

Mar

LaVal, 1977

Apr

Davis et al., 1964

Jan

Gardner et aL, 1970

May

Jun

Jul

Tamsitt and Valdivieso, 1961

Colombia

May

Thomas, 1972

Oct

Dec

Peru

Jun

Tuttle, 1970

Aug

Vampyrops dorsalis

Colombia

Aug

Arata and Vaughn, 1970

Jan

Thomas, 1972

Feb

Mar*

Apr

May

Jun

Jul*

Aug

Sep

Oct

Nov

Dec

Vampyrops brachycephalis

Venezuela

Feb

Rouk and Carter, 1972

Jul

Oct

1 3

Colombia

Jul

Peru

Aug

Vampyrops helleri

Tabasco

May

Villa-R., 1966

Chiapas

May

Jul

Davis et al., 1964

Guatemala

May

Rick, 1968

El Salvador

Jun

LaVal, 1969

Honduras

Aug

Nicaragua

Mar

Jones et al., 1971 a

Apr

Jun

Jul

Aug

Costa Rica

Mar

Mares and Wilson, 1971

Aug

Starrett and de la Torre, 1964

Panama

Jan

Fleming et al., 1972

350

SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

Table 8. — Continued.

Panama

Apr

Fleming et al., 1972

Jul

Sep

Oct

Nov

Dec

Colombia

Aug

Arata and Vaughn, 1970

Jan*

Thomas, 1972

Feb

Mar

Apr*

May

Jun*

Jul

Aug

Sep

Oct

Nov

Dec

French Guiana

Aug

Sep

Peru

Jul

Tuttle, 1970

Aug

Vampyrodes caraccioli

Veracruz

Apr

Villa-R., 1966

Chiapas

Jun

Jones, 1964 b

Jul i

Davis et al., 1964

Honduras

May

Aug

Valdez and LaVal, 1971

Nicaragua

Jul

Jones et al., 1971a

Aug

Panama

Jan

Fleming et al., 1972

Apr

Tobago

Sep

Goodwin and Greenhall, 1961

Colombia

Jan

Thomas, 1972

Feb

Mar*

Apr

May

Jun

Jul

Aug

Thomas, 1972

Sep

Oct

Nov

Peru

Jun

Tuttle, 1970

Jul

Pregnant and lactating.

BIOLOGY OF THE PHYLLOSTOMATIDAE

351

Vampyrops brachycephalis

It is fruitless to speculate on the reproductive pattern of V. brachycephalis
on the basis of the few known records (Table 8).

Vampyrops heller i

Jones et al. (1971 a) suggested that Nicaraguan V. helleri probably breed
throughout much of the year. Fleming et al. (1972) thought that this species
might be bimodally polyestrous based on their evidence from Panama. Thomas’
(1972) work in Colombia, by far the most extensive, indicated a single period
of nonpregnancies from July through September. This is also suggestive of a bi-
modal polyestrous pattern. See Table 8.

Vampyrops lineatus

Peracchi and Albuquerque (1971) reported pregnant females in January,
March, and December in Brazil.

Vampyrops recifinus

Nothing is known about the reproductive pattern of V. recifinus.

Vampyrodes caraccioli

The data from Colombia (Table 8) suggest the familiar pattern of two sequen¬
tial breeding periods followed by a quiescent period, as indicated here by fewer
pregnancies during the July-September period.

Vampyressa pusilla

Although the data are not complete (Table 9), they suggest a pattern of bi-
modal polyestry. Panamanian females have been recorded as preg¬
nant and lactating during the early part of the rainy season, whereas records
from Colombia indicate the mid-rainy season break seen in other species in this
area.

Vampyressa melissa

Nothing is known about the reproductive pattern of this species, although
Gardner (1976) reported three reproductively inactive females from Peru in
May.

Vampyressa nymphaea

Colombian samples (Table 9) are substantial, and indicate the familiar pattern
of two periods of activity followed by a quiescent period. The time of inactivity
seems to be slightly later in V. nymphaea than in other species.

352

SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

Table 9. — Reproductive data for the genera Vampyressa, Chiroderma, and Ectophylla.

Place Date Pregnant Lactating Inactive Reference

Vampyressa pusilla

Campeche

Feb

Jones et al., 1973

Chiapas

Jul

Davis et aL, 1964

Guatemala

Jul

Rick, 1968

Honduras

Aug

Valdez and LaVal, 1971

Nicaragua

Mar

Jones et al., 1971a

Jul

Starrett and de la Torre, 1964

Costa Rica

Feb

Mares and Wilson, 1971

Mar

Jul

Armstrong, 1969

Aug

Panama

Jan

Fleming et al., 1972

Mar

Apr

Hall and Jackson, 1953

Colombia

Mar

Thomas, 1972

Apr

May

rr •

Jul

Aug

Nov

Aug

Arata and Vaughn, 1970

Vampyressa nymphaea

Nicaragua

Feb

Jones et al., 1971a

Costa Rica

Apr

Gardner et al., 1970

Panama

May

Hall and Jackson, 1953

Colombia

Jan

Thomas, 1972

Feb

Mar*

Apr*

May

Jun

Jul*

Aug*

Sep

Oct

Nov

Dec

Chiroderma villosum

Chiapas

May

Davis et al,, 1964

Jul

Dec

Nicaragua

Mar

Jones et al., 1971a

Jul

Panama

Mar

Fleming et al., 1972

Apr

BIOLOGY OF THE PHYLLOSTOMATIDAE

353

Table 9. — Continued.

Trinidad

Aug

Goodwin and Greenhall, 1961

Sep

Colombia

Jan

Thomas, 1972

Peru

Aug

Tuttle, 1970

Chiroderma salvini

Chihuahua

Jul

Anderson, 1972

Sinaloa

Jan

Jones et al., 1972

Jalisco

Feb

Watkins et al., 1972

Colima

Sep

Villa-R., 1966

Honduras

Jul

Carter et al, 1966

Jul

LaVal, 1969

Aug

Colombia

Jan

Thomas, 1972

Mar*

Apr*

May

Jun

Jul

Oct

Nov

Dec

Chiroderma trinitatum

Panama

Feb

Fleming et al, 1972

May

Sep

Trinidad

Mar

Goodwin and Greenhall, 1961

Colombia

Jul

Thomas, 1972

Peru

Jul

Tuttle, 1970

Brazil

Jun

USNM

Jul

Ectophylla macconnelli

Colombia

Jan

Thomas, 1972

Peru

Aug

Tuttle, 1970

‘Pregnant and lactating.

Vampyressa brocki

The only published record of reproductive activity in V. brocki is that of
Baker et al. (1972), who reported one lactating and two pregnant females from
Colombia that were taken in June and July.

Vampyressa bidens

Davis (1975) reported two of 14 females pregnant in December in Peru. This
appears to be the only published record of reproductive activity for this species.

354

SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

Chiroderma doriae

Nothing is known about the reproductive pattern of C. doriae.

Chiroderma improvisum

No information is available on reproduction in this species.

Chiroderma villosum

Although the records listed in Table 9 are diverse, they are too insufficient to
have much predictive value. Davis et al. ( 1 964) suggested that this species breeds
throughout the year on the basis of their specimens from Chiapas. These data
also fit the pattern of bimodal polyestry fairly well, but unfortunately there are no
records from late in the rainy season.

Chiroderma salvini

This species is obviously polyestrous in Colombia, and when further data are
available, may prove to have a bimodal pattern similar to that found in other
species of Colombian phyllostomatids. See Table 9.

Chiroderma trinitatum

Analysis of the reproductive pattern of C. trinitatum must await further data
(see Table 9). Pregnancy records are all from early in the rainy season and late
the dry season.

Ectophylla alba

Gardner et al. (1970) reported a pregnant female in March and a lactating
female in April from Costa Rica. LaVal (1977) recorded pregnant females in
February and August in Costa Rica. He also found one lactating female in March,
and postlactating animals in September and November.

Ectophylla macconnelli

In addition to the records shown in Table 9, A. L. Gardner (personal com¬
munication) collected a lactating female in May and a pregnant female in July
from Peru.

Artibeus cinereus

The records for Colombia (Table 10) are in accord with the pattern of bi¬
modal polyestry as suggested for several other Colombian species. Larger samples
would help to define pregnancy and birth peaks.

Artibeus glaucus

I can find no published records of reproductive activity for A. glaucus, but
there is a USNM specimen from Venezuela recorded as lactating in August.

BIOLOGY OF THE PHYLLOSTOMATIDAE

355

Alfred L. Gardner (personal communication) has collected inactive females in
Peru in April and May.

Artibeus watsoni

In addition to the data in Table 10, Davis (1970) recorded pregnant females
from the months of February, March, April, July, August, and November from
throughout the range of A. watsoni (southern Mexico-Panama). Fleming (1973)
suggested that this species provides an example of bimodal polyestry in Panama.
The few data from Nicaragua also fit this pattern.

Artibeus phaeotis

Davis (1970) reported pregnant females in January, February, April, June,
July, and August, and inactive females from all other months except November
from throughout the range of A. phaeotis (Sinaloa to Panama). Fleming (1973)
reported A. phaeotis as seasonally polyestrous in Panama, and the data from
Mexico seem to support this. Heithaus et al. (1975) suggested bimodal polyestry
as the pattern in Costa Rica as well. See Table 10.

Artibeus toltecus

Davis (1969) recorded pregnant females in each month from January through
August in Mexico and Central America (Table 10). Davis et al. (1964) sug¬
gested an extended breeding season for A. toltecus and mentioned the possibility
of their having two births per year. The data support this assertion.

Artibeus aztecus

Davis (1969) mentioned three pregnant and two inactive females taken in
March and April in either southern Mexico, Guatemala, or Honduras. The data
in Table 10 from northern Mexico suggest that these bats are pregnant during
the summer months. Additional information from other times of the year would
be useful in clarifying the reproductive cycle.

Artibeus hirsutus

Anderson (1960) suggested that A. hirsutus lacks a restricted breeding season.
In support of this claim, Findley and Jones (1965) found in Sonora that two of
the lactating females had placental scars while a third had sperm in the uterus.
They also found three males with sperm and eight without in the same sample.
As they pointed out, spermatogenesis, copulation, lactation, and parturition all
were occurring at the same time. See Table 10.

Artibeus inopinatus

Reproductive information for A. inopinatus seems to be lacking, but in the
description of the species, Davis and Carter (1964) mentioned five young animals
taken in August in Honduras. Two of the young bats appeared to be about one

356

SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

Table 10. — Reproductive data for the genus Artibeus.

Place Date Pregnant Lactating Inactive Reference

Artibeus cinereus

Trinidad

Sep

Goodwin and Greenhall, 1961

Oct

Venezuela

Jul

USNM

Aug

Colombia

Jan

1* 2

Thomas, 1972

Feb

Mar

Apr

May

1 1

Jul

Aug

1 2

Sep

1 4

Oct

Nov

Dec

Aug

5 9

Arata and Vaughn, 1970

Peru

Jul

Tuttle, 1970

Brazil

Jun

USNM

Jul

Artibeus watsoni

Guatemala

Mar

Jones, 1966

Nicaragua

Feb

Jones et al., 1971a

Aug

Panama

Jan

Fleming et al, 1972

Feb

Apr

Jun

Aug

Dec

Artibeus phaeotis

Sinaloa

Jul

Jones et al., 1972

Oct

Jalisco

Jan

Watkins et al., 1972

Apr

Jun

Aug

Campeche

Jan

Jones et al., 1973

Feb

Mar

Quintana Roo

Aug

Apr

Birney et al., 1974

So. Mexico

Jan

Villa-R., 1966

Feb

BIOLOGY OF THE PHYLLOSTOMATIDAE

357

Table 10. — Continued.

So. Mexico

Sep

Villa-R., 1966

Oct

Guatemala

Mar

Jones, 1966

Apr

Murie, 1935

May*

Rick, 1968

Panama

Jan

Fleming et al., 1972

Feb

Mar

Apr

Jun

Aug

Artibeus toltecus

Tamaulipas

Jul

de la Torre, 1954

Sinaloa

Jan

Jones et al., 1972

May

Oct

Jalisco

Jan

Watkins et al., 1972

Feb

Mar

Apr

Jun

Jul

Aug

Sep

Puebla

Jan

LaVal, 1972

Chiapas

May

Davis et al., 1964

Jun

Aug

El Salvador

Jan

Burt and Stirton, 1961

Nicaragua

Apr

Jones et al., 1971 a

Jun

Artibeus aztecus

Tamaulipas

Jul

Alvarez, 1963

Aug

Durango

Jul

Baker and Greer, 1962

Sinaloa

Jul

Jones et al., 1972

Queretaro

Jan

Schmidly and Martin, 1973

Mexico

Sep

Villa-R., 1966

Artibeus hirsutus

Chihuahua

Jul

Anderson, 1972

Sonora

Apr

Cockrum and Bradshaw, 1963

May

Cockrum, 1955

May

Anderson, 1960

Sep

6 4

Findley and Jones, 1965

Sinaloa

Jun

1 1

Jones et al., 1972

Jul

Aug

Dec

Jalisco

Feb

Watkins et al., 1972

Jun

X 15

Aug

X 1

358 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

Table 10. — Continued.

Guerrero

May

Anderson, 1960

Artibeus jamaicensis

Tamaulipas

Mar

de la Torre, 1954

May

Alvarez, 1963

San L. Potosf

Jun

Cockrum, 1955

Sinaloa

Jan

Jones et al., 1972

Feb

Apr

May

Jun

Jul

Sep

Nov

Jun

Anderson, 1960

Nayarit

Apr

Villa-R., 1966

Jalisco

Jul

Anderson, 1960

Jan

Watkins et al., 1972

Feb

Mar

Apr

May

Jun

Jul

Oct

Guerrero

Feb

Villa-R., 1966

Morelos

Jul

Novick, 1960

Queretaro

Jan

Schmidly and Martin, 1973

Puebla

Jan

LaVal, 1972

Veracruz

Feb

Hall and Dalquest, 1963

Jul

Webb etal., 1967

Aug**

Barlow and Tamsitt, 1968

Yucatan Pen.

Apr

Bowles, 1973

May

Feb

Jones et al., 1973

Apr

May

Jul

Aug

Mar

Birney et al, 1974

Apr

Isla Cozumel

Aug

Jones and Lawlor, 1965

Oaxaca

Apr

USNM

Guatemala

Jan

Jones, 1966

Feb

Mar

Aug

May

Rick, 1968

El Salvador

Dec

Burt and Stirton, 1961

Costa Rica

Jan

Fleming et al., 1972

Feb

Mar

BIOLOGY OF THE PHYLLOSTOMATIDAE

359

Table 10. — Continued.

Costa Rica

Apr

Fleming et al., 1972

May

Jun

Jul

Sep

Oct

Nov

Dec

Aug*

Tamsitt and Valdivieso, 1961

Panama

Jan

Fleming et al., 1972

Feb

Mar

Apr*

May*

Jun

Jul

Aug

Sep

Oct

Nov

Dec

Jamaica

Dec

Goodwin, 1970

Feb

McNab, 1976

Providencia

Jan

Tamsitt and Mejia, 1962

Puerto Rico

Feb

Fenton, 1969

Feb

Tamsitt, 1970

Mar

Apr

Jun

Anthony, 1918

Aug

Tamsitt and Valdivieso, 1970

Virgin Is.

Apr**

Barlow and Tamsitt, 1968

Jul**

Trinidad

Feb

Goodwin and Greenhall, 1961

Mar

Apr

May

Jun

Jul

Sep

Jun

Jones, 1946

Colombia

Jun

Tamsitt and Valdivieso, 1963 6

Jul

Arata and Vaughn, 1970

Aug

Jan

Thomas, 1972

Feb

Mar*

Apr*

May

Jun

Jul*

Aug

360

SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

Table 10. — Continued.

Colombia

Sep

Thomas, 1972

Oct

Nov

Dec

Venezuela

Jul

Smith and Genoways, 1974

Peru

Jun

Tuttle, 1970

Jul

Aug

Artibeus lituratus

Tamaulipas

Mar

de la Torre, 1954

May

Alvarez, 1963

Durango

Jun

Jones, 1964 c

Sinaloa

Feb

Jones et al., 1972

Apr

Jun

Jul

Oct

Nov

Anderson, 1960

Jalisco

Mar

Watkins et al., 1972

Apr

Jun

Jul

Aug

Sep

Oct

Morelos

May

Cockrum, 1955

Queretaro

Jan

Spenrath and LaVal, 1970

Veracruz

Feb

Hall and Dalquest, 1963

Yucatan Pen.

Jan

Jones et al., 1973

Feb

Apr

Jul

Oaxaca

Apr

Villa-R., 1966

Guatemala

Feb

Jones, 1966

Mar

May

Rick, 1968

El Salvador

Jul

Starrett and de la Torre, 1964

Costa Rica

Jul

Jan

Fleming et al., 1972

Feb

Apr

May

Jul

Sep

Oct

Nov

Dec

Panama

Jan

Mar

Apr

May

BIOLOGY OF THE PHYLLOSTOMATIDAE

361

Table 10. — Continued.

Panama

Aug

Fleming et al., 1972

Sep

Oct

Mar

Bloedel, 1955

Apr

Hall and Jackson, 1953

May

Trinidad

Feb

Goodwin and Greenhall, 1961

Mar

Apr

May

Jun

Jul

Aug

Sep

Oct

Venezuela

Aug

USNM

Jul

Smith and,Genoways, 1974

French Guiana

Aug

Brosset and Dubost, 1967

Sep

Colombia

Jan

Tamsitt and Valdivieso, 1965 a

Feb

Mar*

Apr

May

Jun*

Jul

Aug

Sep*

Oct*

Nov*

Jan

Thomas, 1972

Feb

Mar

Apr*

May*

Jun

Jul*

Aug*

Sep

Oct

Nov

Dec

Jul

Arata and Vaughn, 1970

Aug

Peru

Jul

Tuttle, 1970

Brazil

Jun

USNM

Jul

Peracchi and Albuquerque, 1971

Aug

’Pregnant and lactating.
’’Twins.

362

SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

month old and the others were older, but still in subadult pelage. Baker and Jones
(1975) also recorded August-taken young from Nicaragua.

Artibeus concolor

The only record of reproduction in A. concolor is that of Thomas (1972), who
collected a pregnant female in February in Colombia.

Artibeus jamaicensis

Artibeus jamaicensis is one of the few species for which adequate information
on reproduction is available (Table 10). Goodwin (1970) reported that breed¬
ing is generally synchronized in Jamaica; Tamsitt and Mejia (1962) discussed a
restricted season on Providencia; and Felten (1956a) suggested that breeding
occurs in the dry season in El Salvador. On the other hand, Tamsitt (1966) and
Jones et al. (1973) argued for continuous or acyclic breeding behavior in Colom¬
bia and the Yucatan Peninsula, respectively. Fleming et al. (1972) and Fleming
(1973) have shown this species to be seasonally polyestrous in Panama and
Costa Rica. Fleming et al. (1972) also presented data on testis size correlated
with spermatogenic activity in males. Heithaus et al. (1975) supported the case
for bimodal polyestry in Costa Rica, pointing out that the two birth peaks occur
at times of peak flower and fruit availability.

Fleming (1971) has shown that A. jamaicensis has a unique seasonally poly¬
estrous cycle in Panama. A peak in parturition occurs in March and April, fol¬
lowed by postpartum estrous and a second peak in parturition in July and August.
“Blastocysts conceived after the second birth implant in the uterus but are dor¬
mant from September to mid-November, when normal development again re¬
sumes” (Fleming, 1971:402). Embryos then develop and young are bom during
the March-April birth peak.

Artibeus lituratus

In the northern part of its distribution, A. lituratus produces only one young
per year, but farther southward, the period of reproductive activity is extended
(Table 10). In Costa Rica and Panama, these bats probably are on a bimodal
cycle with a quiescent period after the second birth peak in the rainy season
(Heithaus et al., 1975). In Colombia, breeding proceeds throughout the year
(Tamsitt and Valdivieso, 1963a). Tamsitt (1966) noted that A. lituratus is an
acyclic or continuous breeder in Colombia. Tamsitt and Valdivieso (19656)
studied the reproductive cycle of males in Colombia and their data, based on
presence of sperm, length and tubule diameter of the testes, and diameter of the
epididymides, indicate that males are capable of reproductive activity at any time
of the year, and that the reproductive pattern is acyclic without any suggestion of
seasonal variation.

Thomas (1972) presented a much more extensive sample from Colombia and,
although he has confirmed year-round activity with the presence of pregnant,

BIOLOGY OF THE PHYLLOSTOMATIDAE

363

lactating, and inactive females in every month of the year, his data indicate bi-
modal activity peaks. Pregnancy peaks occur in December and May, with lacta¬
tion peaks lagging about a month behind, as would be expected.

Enchisthenes hartii

Gardner et al. (1970) suggested that E. hartii is reproductively active through¬
out the year in Costa Rica. The only inactive animals they found were subadults,
one in May and three in July. This species may be found to undergo a period of
reproductive inactivity when data become available from later in the year. See
Table 1 1.

Ardops nichollsi

I can find no records other than those of Jones and Schwartz (1967) who re¬
ported four pregnant females in March and one lactating and two pregnant fe¬
males in April from Dominica.

Phyllops falcatus

No information is available on reproduction in this species.

Phyllops haitiensis

Nothing is known about the reproductive pattern of P. haitiensis.

Ariteus flavescens

No data are available concerning the reproductive pattern of this species.

Stenoderma rufum

Tamsitt and Valdivieso (1966 b) described parturition in S. rufum. This
species seems to be polyestrous on Puerto Rico, but data from the period Septem¬
ber through December are needed in order to clarify their reproductive pattern.
See Table 11.

Pygoderma bilabiatum

Peracchi and Albuquerque (1971) reported a pregnant female collected in
August in Brazil.

Ametrida centurio

Nothing is known about the reproductive pattern of this species.

Sphaeronycteris toxophyllum

Nothing has been published about the reproductive pattern of S. toxophyllum.

364

SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

Table 1 1. — Reproductive data for the genera Enchisthenes, Stenoderma, and Centurio.

Place

Date

Pregnant Lactating Inactive

Reference

Enchisthenes hartii

Honduras

Aug*

LaVal, 1969

Costa Rica

Jan

Gardner et al., 1970

May

Jun

Jul

Aug

Armstrong, 1969

Aug

LaVal, 1977

Colombia

Apr

Thomas, 1972

May

Jul

1 3

Aug

Sep

Peru

Nov

Gardner, 1976

Stenoderma rufum

Puerto Rico

Feb

Tamsitt, 1970

Mar

May

Jul

Aug

Nov

Jul

1 9

Jones et al., 1971 b

Jul

Genoways and Baker, 1972

Aug

Tamsitt and Valdivieso, 19666

Centurio senex

Tamaulipas

Jun

Alvarez, 1963

Jalisco

Mar

Watkins et al., 1972

Aug

Apr

Jones, 19646

Veracruz

Apr

Villa-R., 1966

Yucatan Pen.

Jan

Jones et al., 1973

Feb

Jul

Oaxaca

Mar

Villa-R., 1966

Mar

USNM

Chiapas

Apr

Davis et al., 1964

Jul

Honduras

Aug

LaVal, 1969

Nicaragua

Feb

Jones et al., 1971 a

Mar

Costa Rica

Mar

Mares and Wilson, 1971

Trinidad

Jan

Goodwin and Greenhall, 1961

Oct

Pregnant and lactating.

BIOLOGY OF THE PHYLLOSTOMATIDAE

365

Centurio senex

Although there are a fair number of records for C. senex (Table 11), the data
from any given area are too few to decipher reproductive patterns. Pregnancies
from February and July on the Yucatan Peninsula suggest the possibility of
either polyestry or asynchrony.

Brachyphylla cavernarum

Anthony (1918) reported lactating females in July in Puerto Rico, and Nellis
(1971) found a lactating female in April on St. Croix. Walker (1975) mentioned
pregnant females in February on Puerto Rico, March on St. Croix, and a lactating
female in April on Puerto Rico.

Buden (1977) collected 12 females, all of which were pregnant, in March on
the Island of Caicos in the West Indies. All fetuses were 24 to 34 mm. in length,
suggesting a synchronized cycle. The females lactating in July (Anthony, 1918)
suggest the possibility of a second period of parturition as well.

Brachyphylla nana

Nothing is known about the reproductive pattern of this species.

Erophylla bombifrons

Although the data are sparse (Table 12), they suggest a restricted breeding
season. Females are pregnant from February to June and lactating in July. This
would result in the production of young early in the rainy season, a time when
resources should be most plentiful.

Erophylla sezekorni

Buden (1976) summarized data based on 91 pregnant or lactating females and
immatures. He suggested a gestation period during the first part of the year with
parturition in early summer. He found females carrying small fetuses in February
and larger fetuses in April and May. Lactating females were taken in June and
many juveniles in July. Immature animals approaching adult size were taken in
August. Thus, the pattern appears identical to that described above for E. bombi¬
frons. See Table 12.

Phyllonycteris poeyi

Miller (1904) reported that all of the females he examined from Cuba were
pregnant in June.

Phyllonycteris major

Nothing is known about reproductive patterns of P. major, a bat which is likely
extinct.

366

SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

■

Table 12. — Reproductive date for the genus Erophylla.

Place

Date

Pregnant Lactating Inactive

Reference

Erophylla bombifrons

Hispaniola

Feb

Buden, 1976

Puerto Rico

Jul

Apr

Barlow and Tamsitt, 1968

Jun

Valdivieso et al., 1968

Mar

Walker, 1975

Apr

May

Jul

Erophylla sezekorni

Cuba

Feb

Buden, 1976

Bahamas

Apr

May

Jun

Jul

Jun

8 2

Blake, 1885

♦Plus many immatures.

Phyllonycteris aphylla

The only record of reproductive activity in this species is that of Goodwin
(1970), who reported a pregnant female taken in January on Jamaica.

Desmodus rotundus

More is known about the reproduction of D. rotundus than about any other
phyllostomatid (Table 13). DeVerteiul and Urich (1936) apparently were the
first to suggest that D. rotundus breeds year-round, based on their work on
Trinidadian populations. Wimsatt and Trapido (1952) confirmed this in Panama
by presenting data on both males and females, and suggested a gestation period of
five to six months. Burt and Stirton (1961) reported continuous breeding in El
Salvador. Goodwin and Greenhall (1961) recorded the same thing for popula¬
tions on Trinidad and reported pregnant females, lactating females, and young
animals in every month, although the highest incidence of young was in April and
May and again in October and November. They also suggested that males may
roost separately from females when the young are born.

Crespo et al. (1961) gave a detailed account of reproduction in vampires
based on their work in Argentina during September and November. They found
that in males both testes are active and coincide in their activity rhythm. Sexually
active males with well-developed epididymides containing spermatozoa and in¬
active males with small epididymides and no spermatozoa were found in the
same population at the same time of year. Sexually active males were present in
September and November. In some instances, adult males have epididymides with
few spermatozoa mixed with resting cells, which could be interpreted as the
beginning of a new cycle of activity.

BIOLOGY OF THE PHYLLOSTOMATIDAE

367

For females, Crespo et al. (1961) found that both ovaries are functional, with
only a slight difference in degree of development of follicles. Ovaries are in a
periovarian capsule, and the fallopian tubes begin in the walls of the capsules.
There is always only one embryo, which occupies one uterine horn first but, with
development extends into both horns and the body of the uterus, obliterating the
partitioning of the uterus. At the end of a pregnancy, the ovary without the corpus
luteum is in early proestrous and will produce the next ovum. One postpartum
specimen had a corpus luteum in one ovary and a corpus albicans representing a
previous pregnancy in the other ovary.

In September and November, there are proestrous immature animals bearing
primary and secondary follicles. None of the animals examined had vaginal plugs
or sperm in the uteri.

Hall and Dalquest (1963) mentioned that these animals seem to have no
regular breeding season in Veracruz. They found a few young in various stages
of development, pregnant females, and inactive females in all of the colonies
examined. Dalquest (1955) had earlier pointed this out for San Luis Potosf
populations, and suggested that young are bom in all months of the year.

Villa-R. (1966) found pregnant females, lactating females, and newborn
young at all times of the year during 1 5 years work in Mexico.

Greenhall (1965) described mating behavior (including copulation), preg¬
nancy, and young animals in captivity. Schmidt and Manske (1973) found a
gestation period of seven months and lactation period of three to nine months for
captive animals. Linhart (1971) compiled a useful bibliography of vampire bats.

Diaemus youngii

The only recorded reproductive information for this species is that of Goodwin
and Greenhall (1961) for Trinidad. They found two lactating females in August
and in October they took one immature male, four pregnant females, one lactating
female, and two inactive females.

Diphylla ecaudata

Dalquest (1955) reported that D. ecaudata seems to have a well-defined breed¬
ing season and may have a single young per year in eastern Mexico. Felten
(1956a), however, felt that they breed in both dry and wet seasons in El Salvador
and postulated two litters per year. From the scatter in the records listed in Table
13, I am inclined to agree with Felten.

Summary

The three most obvious reproductive strategies found in the family Phyl-
lostomatidae are summarized in Fig. 1. The most critical environmental param¬
eter is the seasonality of the rainfall pattern. Although a great deal of geographic
variation exists, the pattern of a dry season during the months of January to April
or May is common in Middle America and in many areas in northern South
America. In tropical Mexico, the rains often begin as late as June, but as one

368

SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

Table 13. — Reproductive data for the genera Desmodus and Diphylla.

Place Date Pregnant Lactating Inactive Reference

Tamaulipas

Mar

May

Jun

Aug

Chihuahua

May

Durango

Jun

Sinaloa

Jan

Mar

May

Dec

Nayarit

Jan

Jalisco

Jan

Feb

Mar

Apr

May

Jun

Jul

Aug

Sep

Colima

Mar

May

Jul

Zacatecas

Oct

Michoacan

Jul

Guerrero

Jun

Aug

Sep

Nov

Queretaro

May

Jun

Dec

Puebla

Jan

Morelos

Jan

Mexico

Jan*

Veracruz

Feb

Jun

Jul

Yucatan Pen.

Jan

Feb

Mar

Apr

Jun

Jul

Aug

Apr

Desmodus rotundus

1
2

1 5

9
4

1
1
1
1
1
1

X
X
X
X
X
X
X

X
X

52 1

36 9

2 23

1
1

22
16
21
10
1
1

3
2
1
1

4
1
1
1
2

4 X

1 X
X

Alvarez, 1963

Anderson, 1972
Jones, 1964c
Jones et al., 1972

Cockrum, 1955
Watkins et al., 1972

Burns and Crespo, 1975

Cockrum, 1955
Hall and Villa-R„ 1949
Forment et al., 1971

Schmidly and Martin, 1973

LaVal, 1972
Burns, 1970

Hall and Dalquest, 1963
Lackey, 1970

Jones et al, 1973

Birney et al., 1974

BIOLOGY OF THE PHYLLOSTOMATIDAE

369

Table 13. —

■Continued.

Guatemala

Mar

Jones, 1966

El Salvador

Feb

Felten, 1956c

Mar

May

Jul

Aug

Oct

Nov

Costa Rica

Jan

Fleming et al„ 1972

Feb

Mar

Apr

May

Jul

Aug

Oct

Nov

Dec

Panama

Apr

May

Feb

Wimsatt and Trapido, 1952

Apr

May

Jul

Nov

Trinidad

Jan

DeVerteiul and Urich, 1936

Jun

Nov

Dec

Colombia

Nov

Tamsitt and Valdivieso, 1963 6

Jul

Arata and Vaughn, 1970

Apr

Thomas, 1972

May

Oct

Venezuela

Apr

Pirlot and Leon, 1965

Brazil

Jan

Peracchi and Albuquerque, 1971

Diphylla ecaudata

Tamaulipas

Nov

Alvarez, 1963

San L. Potosi

Mar

Dalquest, 1953

Jul

Yucatan

Nov

Hatt, 1938

May

Birney et aL, 1974

Mexico

Aug

Villa-R., 1966

Oct

Nov

El Salvador

Aug

Felten, 1956c

Honduras

Jul

Valdez and LaVal, 1971

Nicaragua

Apr

Jones et at ., 1971 a

370

SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

ASEASONAL POLYESTRY

LACTATION LACTATION

GESTATION PARTURITION GESTATION

BIMODAL POLYESTRY

GESTATION PARTURITION

MONESTRY

LACTATION

GESTATION PARTURITION

DESMODUS

PARTURITION

GLOSSOPHAGA
CAROLL1A
URODERMA
ART I BE US

LACTATION

MAC ROT US
LEPTONYCTERIS

GESTATION PARTURITION LACTATION

maximum

stress

MAXIMUM FRUIT

MAXIMUM INSECTS

dry

rains

begin

heavy

rains

JAN

FEB MAR

APR

MAY

JUN JUL AUG SEP

OCT NOV DEC

Fig. 1. — Summary of the three common reproductive patterns and of the environmental
events affecting them.

moves southward, the length of the dry season generally decreases, and in some
areas may be only a month or less in duration. Also, annual variations occur in
any given locality. Nevertheless, for purposes of this discussion, the pattern rep¬
resented in Fig. 1 may be taken as representative.

This environmental seasonality affects reproductive cycles of bats through the
food supply. The time of maximum abundance of a wide variety of both fruits and
insects is just after the beginning of the rainy season. Thereafter, a general de¬
cline is seen, culminating in a period of minimal abundance during the dry season.

The critical time for most bat populations seems to be the period of weaning
of the young (Wilson and Findley, 1970; Fleming et al., 1972). Thus, although
it may be possible for females to undergo gestation and lactation during the stress¬
ful time of year, the young are usually weaned during the most energetically
favorable periods.

In monestrous species of the Phyllostomatidae, there is a distinct period of
reproductive activity culminating with weaning of the young shortly after the be¬
ginning of the rainy season when food is plentiful. This pattern is seen in some
species at the northern limit of the range of the family, where the time of maxi¬
mum food availability is fairly short. The nectarivorous bats of the genus
Leptonycteris show this pattern in the southwestern United States, where they
migrate northward from Mexico and have their young in May or June. These
young are weaned in July or August, the peak of the rainy season and the period

BIOLOGY OF THE PHYLLOSTOMATIDAE

371

Table 14. — Reproductive patterns of the 20 species for which adequate data exist.

Macrotus waterhousii

delayed development and monestry

Glossophaga soricirui

continuous or bimodal polyestry

Leptonycteris sanborni

monestry or bimodal polyestry

Choeronycteris mexicana

monestry

Carollia castanea

bimodal polyestry

Carollia subrufa

continuous or bimodal polyestry

Carollia brevicauda

bimodal polyestry

Carollia perspicillata

bimodal polyestry

Sturnira 1 ilium

bimodal polyestry

Uroderma bilobatum

bimodal polyestry

Vampyrops helleri

bimodal polyestry

Vampyrodes caraccioli

bimodal polyestry

Vampyressa pusilla

bimodal polyestry

Vampyressci nymphaea

bimodal polyestry

Artibeus cinereus

bimodal polyestry

Artibeus watsoni

bimodal polyestry

Artibeus phaeotis

bimodal polyestry

Artibeus jamaicensis

bimodal polyestry and delayed development

Artibeus lituratus

geographically variable

Desmodus rotundus

continuous polyestry

of peak flower abundance. In October, individuals migrate back to Mexico for
the winter. A variation of this pattern is found in Macrotus californicus , where
the embryos undergo delayed development during the autumn and winter months
and begin developing at a more normal rate in spring. This results in parturition
and weaning periods similar to those of Leptonycteris. There is a possibility that
some individuals of Leptonycteris sanborni have a second period of reproductive
activity resulting in the production of offspring in Mexico in November. If so this
species would more properly belong in the next category, that of bimodal
polyestry.

The majority of species of phyllostomatids for which there is ample data show a
reproductive pattern involving an extended breeding season with two birth peaks
a year. In these species (for example, some members of the genera Glossophaga,
Carollia, Uroderma, and Artibeus), the young from the first birth peak are weaned
at the beginning of the rainy season, and those from the second pregnancy of the
year are weaned well into the rainy season. These two peaks are followed by an
inactive period, which results in no young being weaned during the stressful dry
season.

At the other extreme from monestry are those animals that are completely
polyestrous and produce young continuously and asynchronously throughout the
year. The evidence to date shows only the vampire bat Desmodus rotundus to
be in this category. These animals are adapted to a food supply (primarily blood
from domestic cattle) that is available throughout the year over much of their
range. However, because their gestation period is five or six months long, the
net result still is only two young per year.

372

SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

Table 14 summarizes the type of reproductive pattern for the 20 species for
which there is a reasonable amount of data. It should be noted that many of these
species will show geographic variation in the timing of reproductive events, and,
in some cases ( Artibeus lituratus, for example), the species may have completely
different patterns in different areas. This is hardly surprising in view of the wide
geographic and ecologic range of many of the species.

All of these patterns may be thought of as variations on a single theme —
maximizing the production of offspring with available environmental energy
resources. Further study will undoubtedly add a wealth of data on the fine tuning
of the various mechanisms involved in selecting for a particular reproductive
strategy for a given species.

Acknowledgments

I gratefully thank the following people: Michael A. Bogan, Estella C. Duell,
Robert D. Fisher, Theodore H. Fleming, Alfred L. Gardner, Barbara A. Harvey,
Daniel H. Janzen, Clyde Jones, Patricia Mehlhop, S. Jerrine Nichols, and William
A. Wimsatt.

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Rasweiler, J. J. 1972. Reproduction in the long-tongued bat, Glossophaga soricina.

I. Preimplantation development and history of the oviduct. J. Reprod. Fert.,
31:249-262.

Rick, A. M. 1968. Notes on bats from Tikal, Guatemala. J. Mamm., 49:516-520.

Rouk, C. S., and D. C. Carter. 1972. A new species of Vampyrops (Chiroptera:

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Schaldach, W. J., Jr. 1965. Notas breves sobre algunos mamiferos del sur de Mexico.
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Schmidly, D. J., and C. O. Martin. 1973. Notes on bats from the Mexican state of
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Schmidt, U., and U. Manske. 1973. Die Jugendentwicklung der Vampirfledermause
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Schwartz, A., and J. K. Jones, Jr. 1967. Bredin-Archbold-Smithsonian Biological
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Smith, J. D., and H. H. Genoways. 1974. Bats of Margarita Island, Venezuela, with
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Spenrath, C. A., and R. K. LaVal. 1970. Records of bats from Queretaro and San Luis
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Starrett, A., and L. de la Torre. 1964. Notes on a collection of bats from Central
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Taddei, V. A. 1976. The reproduction of some phyllostomidae (Chiroptera) from the
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Tamsitt, J. R. 1966. Altitudinal distribution, ecology, and general life history of bats in
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Wimsatt, W. A., and H. Trapido. 1952. Reproduction and the female reproductive cycle
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Anat., 91:415-445.

EMBRYOLOGY

William J. Bleier

Over the years, there have been numerous reports concerning reproduction
in the phyllostomatid bats, but a survey of the literature reveals that data on
the embryology of the Phyllostomatidae are limited to gross morphological
observations of reproductive tissues, embryos, and mammary glands of
individuals from natural populations. These reports have provided useful
information concerning times of pregnancy, lactation, and spermatogenesis.
Thus, a basic knowledge of reproductive cycles for a number of the phyl¬
lostomatid bats has been accumulated (for review, see Wilson, this volume).

However, there have been few microscopic studies of reproduction and
embryological development in the Phyllostomatidae. With respect to the
details of the embryology of these bats, only seven species representing
five genera have been studied microscopically. Some of these works are
based on tissues collected from natural populations; others, on tissues from
laboratory colonies.

This paper reviews the data now available on the embryology of the
Phyllostomatidae. In order to facilitate this presentation, developmental
events will serve as major subdivisions, and, within these subdivisions, the
data available on the various species will be presented. The subdivisions to
be considered are ovulation, fertilization, preimplantation embryonic
development, implantation, postimplantation embryonic development, and
placentation.

Ovulation

Macrotus californicus. — Studies of M. californicus indicate that ovulation
is from the right ovary only (even though both ovaries develop Graafian
follicles), and, typically, that only one ovum is released. It is not known if
ovulation is spontaneous in Macrotus (Bradshaw, 1961).

Glossophaga soricina. — Ovulation in G. soricina may occur from either
ovary, and there is a tendency for it to alternate between the two. Ovulation is
spontaneous and usually only one ovum is released per cycle. Menstruation
occurs in G. soricina and ovulation takes place at, or very close to, the time
of menstruation (Hamlett, 1935; Rasweiler, 1972).

Carollia perspicillata, C. brevicauda, and Desmodus rotundus. — Ovulation
in these three species is basically the same as in G. soricina. However, it is
not known if ovulation is spontaneous. Menstruation in Carollia and Desmodus
is similar to that of G. soricina.

Artibeus lituratus. — Ovulation in Artibeus lituratus may occur from either
ovary (Tamsitt and Valdivieso, 1963, 1965).

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Fertilization

Because there is no evidence for sperm storage in the female reproductive
tracts of phyllostomatid bats, it appears that fertilization occurs shortly after
copulation. Hence, the phenomenon of delayed fertilization that has been
observed in some of the Vespertilionidae (Wimsatt, 1942) has not been reported
in any of the Phyllostomatidae.

Preimplantation Embryonic Development

Macrotus californicus. — Studies on M. californicus have revealed the
sequence of events prior to implantation; however, the timing of these events
has not been determined (Bleier, 1975a). Development to a blastocyst
occurs in the oviduct and was predicted to require 10 to 20 days (Bleier, 1975/?).
Embryonic development follows the pattern typical for other therian mammals.
There is no information concerning the loss of the zona pellucida in Macrotus.

Glossophaga soricina. — In studies of a laboratory colony, Rasweiler
(1972) was able to time the sequence of events in embryonic growth of G.
soricina. The two-celled stage of development is attained by day 2 or 3 post¬
ovulation. The eight-celled stage is reached by days 5 to 7; the 32-celled stage,
by day 8; the blastocyst stage, by day 10. Compared to development in other
mammals, cleavage rate in Glossophaga is slow. The zona pellucida is usually
lost on day 1 2 or 1 3, and, prior to its loss, the embryo has been contained within the
ampulla of the oviduct. Upon loss of the zona pellucida, the embryo is located in
the intramural uterine cornu, which is the site of implantation. There is no evidence
of differentiation of germ layers during this preimplantation period.

Carollia perspicillata and C. brevicauda. — Cleavage in C. perspicillata
and C. brevicauda also proceeds slowly. De Bonilla and Rasweiler (1974)
reported that the first blastocyst was observed on day 10 postcoitum. Again,
development to the blastocyst stage and loss of the zona pellucida occurs in the
oviduct. Earliest loss of the zona pellucida was day 10.

Artibeus jamaicensis. — The only information available on early embryonic
development in A. jamaicensis was reported by Fleming (1971), who found
two reproductive cycles per year in Panamanian populations and noted that
the embryo reaches the blastocyst stage before entering the uterus. An unusual
feature is that during one of the cycles (August to March) there is a 2.5-month
period of delayed embryonic development. During this period of retarded
development, the only noticeable morphological change is an increase in the
size of the blastocyst.

Desmodus rotundus. — Slow cleavage also is characteristic of D. rotundus.
Quintero and Rasweiler (1974) observed a two-celled embryo as late as day
7 postcoitum in an individual from a laboratory colony. A blastocyst was not ob¬
served until day 15. Loss of the zona pellucida occurred in the oviduct, and the
earliest date of this loss was day 15. Wimsatt (1954) noted that endoderm dif¬
ferentiation in the blastocyst begins while the blastocyst is still in the oviduct.

BIOLOGY OF THE PHYLLOSTOMATIDAE

381

Implantation

Macrotus californicus. — Several reports are available concerning implantation
in M. californicus. Bradshaw (1962) noted that implantation occurs during
early gestation. Later studies by Bodley (1974) and Bleier (1975a, 19756)
have provided more details concerning the process in Macrotus. Central
implantation is initiated shortly after the arrival of the blastocyst into the uterus.
Early stages are characterized by a deterioration of the uterine epithelium
such that the invading trophoblast comes into contact with the basal lamina of
the uterine epithelium. Endoderm differentiation is initiated at this time.
By the end of October, implantation has progressed to the point that the entire
uterine epithelium that once surrounded the embryo has now' been obliterated.
The trophoblast is largely multilayered at this time, but unilaminar portions
may be observed in the abembryonic regions. Reichert’s membrane separates
the trophoblast from the remaining fetal tissue and becomes continuous
throughout the embryonic and abembryonic regions. The age of an embryo at
this stage is estimated to be 20 to 30 days (Bleier, 19756). By mid December,
syncytiotrophoblast has differentiated; there is considerable proliferation
of the syncytiotrophoblast by the end of January. At this time, an interstitial
membrane (presumptive intrasyncytial lamina) is conspicuous between the
maternal tissue and the syncytiotrophoblast. Reichert’s membrane, which
reaches its greatest thickness in late January, disappears by mid February.
Endoderm completely surrounds the yolk sac cavity at this stage. By mid
February all the layers that comprise the definitive placenta are present
(Bodley, 1974; Bleier, 19756).

Glossophaga soricina. — Implantation in G. soricina is initially central and
secondarily interstitial (Rasweiler, 1974). Rasweiler (1974) divided this
process of implantation into eight stages. Stage I (12 to 14 days postcoitum)
blastocysts resemble ampullary blastocysts; however, there is some hypertrophy
of the trophoblast in Stage I embryos. The uterine epithelium is intact but at
times flattened. The blastocyst is oriented such that the inner cell mass is
toward the cephalic side of the blastocystic cavity. The first appearance of
endoderm differentiation is at this stage. Stage II blastocysts (days 13 to 15)
are characterized by a bilaminar and multilaminar trophoblast in the embryonic
polar region, whereas the trophoblast of the abembryonic region remains
unilaminar. Necrosis of the maternal epithelium has begun in the bilaminar and
multilaminar regions and the trophoblast has penetrated the basal lamina of the
uterine epithelium. Stage III blastocysts (days 14 to 16) resemble Stage II blas¬
tocysts, but the uterine epithelium has deteriorated further. In some areas, the
trophoblast has penetrated to the maternal basal lamina. Endoderm is clearly
recognized in all specimens from Stage III. Stage IV specimens (days 15 to 17) are
characterized by complete obliteration of the uterine luminal epithelium with
encroachment of the trophoblast to the uterine glands. A decidual reaction
first appears at this stage. During Stage IV, the endoderm and inner cell mass

382 SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

fill almost the entire space of the blastocystic cavity. Solid multilayered masses
of endoderm occur on the ventral side of the inner cell mass, and, by days
16 to 17, pockets have begun to develop in the endoderm. Endoderm appears
on the lateral and dorsal surfaces of the inner cell mass. Stage V (days 16 to 21)
is recognized by the presence of syncytiotrophoblast in the region of the
embryonic pole. Cytotrophoblast at this stage is present outside of the
syncytiotrophoblast, in addition to its position inside the syncytiotrophoblast,
and in some regions has penetrated the glandular epithelium. The fluid-filled
pockets in the endoderm are more pronounced, and in one specimen had
coalesced to form a unilocular condition. By Stage VI (days 20 to 22) and
Stage VII (days 23 to 25), the syncytiotrophoblast has proliferated further
and has begun to penetrate the decidua basalis. There is an increase in vascular
lacunae and a decrease in maternal endothelium in Stage VII individuals.
A lamina that is probably an extension of the abembryonic portion of Reichert’s
membrane is interposed between the inner cell mass and the endoderm dorsal
and lateral to the inner cell mass. Coalescence of the pockets in the endoderm
has continued so that most embryos are unilocular. In Stage VIII (days 26 to
30), the cytotrophoblast has penetrated deep into the syncytiotrophoblast.
During this stage, the intrasyncytial lamina is observed and significant quantities
of maternal blood in the labyrinth first appear. Amniogenesis by cavitation
has begun at this stage. By day 32, differentiation of ectoderm has been
initiated, and thinning of the roof of the amnion has begun. The endoderm and
Reichert’s membrane, in the region of the embryonic pole, have disappeared.
The fate of Reichert’s membrane is currently unknown.

Carollia perspicillata and C. brevicauda. — Little is known about implantation
in Carollia. De Bonilla and Rasweiler (1974) found that the site of implantation
in C. perspicillata and C. brevicauda is similar to that reported for G. soricina;
that is, implantation occurs in the segment between the end of the oviduct
and the main cavity of the uterus.

Artibeus jamaicensis. — The only report on A. jamaicensis is that of Fleming
(1971). Implantation is similar to that observed for Glossophaga soricina and
Desmodus rotundus, including “(i) precocious development of the blastocyst,
which by the time it reaches the uterus, has differentiated into a trophoblast
thickened at the embryonic pole and an embryonic cell mass . . . and (ii)
implantation that is interstitial and cytolytic.”

Desmodus rotundus. — The only observations of implantation in D.
rotundus were reported by Wimsatt (1954): implantation is “cytolytic and
completely interstitial,” occurring antimesometrially in the middle of the uterine
cornu and on the same side as is the ovary from which ovulation occurred.
During early implantation, the embryo is secured to the uterus only in the
region of the embryonic cell mass, thereby exposing the abembryonic surface
to the uterine cavity. The trophoblast near the embryonic pole is multilaminar,
whereas the trophoblast associated with the free surface (abembryonic) is
unilaminar. Beneath the inner cell mass, the endoderm has hypertrophied;

BIOLOGY OF THE PHYLLOSTOMATIDAE

383

in other regions it remains flattened. Wimsatt (1954) also observed precocious
formation of mesoderm, but Rasweiler (1974) speculated that this may
actually be endoderm.

In a second, older specimen, Wimsatt (1954) noted that implantation was
complete. By this stage, the embryo is completely embedded in the endometrium,
and the trophoblast is multilayered in the embryonic region but still largely
unilaminar in the abembryonic region. In both specimens, there is a marked
decidual reaction but it is most pronounced in the older specimen. Amniogenesis
is accomplished by cavitation.

Postimplantation Embryonic Development

Macrotus californicus. — Embryonic growth in M. californicus to the end
of implantation is slow. Fertilization in Macrotus most often occurs during
October, and, by the end of implantation (mid February), amniogenesis by
cavitation has begun. Therefore, the embryo requires approximately four
months to reach the embryonic-disc stage (Bleier, 1975 a). Growth accelerates
during March, and embryos at the limb-bud stage (crown-rump length
approximately 4.5 millimeters) of development are observed. Embryonic
growth continues at a rapid rate, and most parturitions occur during June.
Growth and differentiation of the embryonic tissues and organs, following the
period of slow development, are similar to the pattern that has been described
for other therian mammals.

Glossophaga soricina. — Hamlett (1935) described the embryonic
growth in G. soricina following implantation. His description included a
discussion of the primitive streak and mesoderm formation. Primary mesoderm
is formed early; however, Rasweiler (1974) provided evidence that this
“primary mesoderm” is most likely endoderm. Formation of secondary
mesoderm (that derived from the primitive streak) and subsequent primitive
streak activity are similar to that of any typical mammal. By the six-somite
stage the coelom is present (but absent at the medullary-fold stage) and the
mesoderm has split into splanchnic and somatic layers. The yolk sac remains
large, but the yolk stalk disappears before the 2.5-millimeter stage. There is no
evidence of the allantois in the six-somite specimen (length is one millimeter
from head fold to end of primitive streak), but by the time the embryo reaches
2.5 in length, the allantois has attained its maximum relative size.

Placentation

Macrotus californicus. — Bradshaw (1961) noted that the definitive
placenta in M. californicus is hemochorial. Recently, Bodley (1974) used
electron microscopic techniques that revealed the definitive placenta to be
hemodichorial. Development of the placenta is such that it is large enough to
be readily visible with the naked eye by late March. At this time, reduction of
the cytotrophoblast to a single cell layer begins and syncytial blocks

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(derivatives of the syncytiotrophoblast) replace the maternal endothelium
(Bodley, 1974; Bleier, 1975 b). Changes from March to June involve maturation
of the placenta, but there is no change in the number of cell layers. The layers
of the hemodichorial placenta in Macrotus include syncytial blocks, intrasyncytial
lamina, syncytiotrophoblast, cytotrophoblast, fetal basal lamina, and fetal capillary
endothelium (Bodley, 1974).

Glossophaga soricina. — Hamlett (1935) and Rasweiler (1974) classified
the placenta in G. soricina as discoidal and hemochorial, and Rasweiler (1974)
indicated that formation was rapid. There is an interstitial lamina present,
but its origin is uncertain — Rasweiler (1974) suggested that it is derived from
the trophoblast. The trophoblast differentiates into cytotrophoblast and
syncytiotrophoblast; however, the cytotrophoblast disappears by midgestation.
In addition, the walls and endothelium of the maternal blood vessels are eroded
(Hamlett, 1935) so that there are three cell layers that separate the fetal and
maternal blood streams. These layers are fetal endothelium, loose mesenchyme,
and syncytiotrophoblast.

Carollia perspicillata. — Little is known concerning the placenta in C.
perspicillata. Wimsatt (1958) noted that the placenta is discoidal and
endotheliochorial. Also, he implied that the cytotrophoblast does not persist
to the end of gestation. There is a conspicuous interstitial membrane between
the syncytiotrophoblast and maternal endothelium. However, this observation
was made by using light microscopy. Recent studies indicate that other
phyllostomatid bats have a hemodichorial type placenta and that the “maternal
endothelium” is actually syncytiotrophoblast (Bjorkman and Wimsatt, 1968;
Rasweiler, 1974; Bodley, 1974; Bleier, 1975b). Therefore, it would not be
surprising if it were determined that the “maternal endothelium” in the placenta
were syncytiotrophoblast. If this were true, and if the cytotrophoblast is
lost, then the placenta of Carollia would be a hemochorial type. Further
investigations are needed to confirm the type of placental barrier characteristic
of Carollia.

Artibeus jamaicensis. — Wislocki and Fawcett (1941) stated that the placenta
is discoidal and hemochorial.

Desmodus rotundus. — Initial reports indicated that the placenta in D. rotundus
is discoidal and endotheliochorial (Wimsatt, 1954, 1958). However, by using
electron microscopic methods, Bjorkman and Wimsatt (1968) concluded
that the definitive placenta is hemodichorial, but in earlier stages before the
loss of the maternal endothelium it is endotheliochorial. Thus, the definitive
placenta consists of the following layers: intrasyncytial lamina, syncytiotropho¬
blast, cytotrophoblast, a thick basement membrane, mesenchyme, and fetal
endothelium.

Summary and Conclusions

From the data summarized in this paper, several trends can be seen in the
embryology of the Phyllostomatidae. In general, ovulation may occur from

BIOLOGY OF THE PHYLLOSTOMATIDAE

385

either ovary, except in Macrotus californicus, and fertilization follows
immediately after ovulation and copulation. Embryonic development to the
blastocyst stage appears to be similar to that reported for other therian
mammals; however, the process seems to be considerably slower in the
phyllostomatid bats studied thus far. Implantation is interstitial except in
M. californicus. The placenta is discoidal, and it is likely that the placental
barrier is either hemodichorial or hemochorial.

There are several features of phyllostomid embryology that should stimulate
further investigations of the species reported in this paper. In addition, studies
of other species should be encouraged for they might reveal embryological
strategies other than the ones presently known. Some of the areas deserving the
application of sophisticated research techniques include ovulation from only the
right ovary in M. californicus, delayed embryonic development in M. californicus
and Artibeus jamaicensis, the length of gestation in Desmodus , and menstruation
and interstitial implantation in Glossophaga, Carollia , and Desmodus.

Literature Cited

Bjorkman, N. H., and W. A. Wimsatt. 1968. The allantoic placenta of the vampire
bat ( Desmodus rotund us murinus): a reinterpretation of its structure based on
electron microscopic observations. Anat. Rec., 162:83-98.

Bleier, W. J. 1975a. Early embryology and implantation in the California leaf-nosed
bat, Macrotus californicus. Anat. Rec., 182:237-254.

- . 1975ft. Fine structure of implantation and the corpus luteum in the California

leaf-nosed bat, Macrotus californicus. Unpublished Ph.D. dissertation,
Texas Tech Univ., 75 pp.

Bodley, H. D. 1974. Ultrastructural development of the chorioallantoic placental
barrier in the bat Macrotus waterhousii. Anat. Rec., 180:351-368.

Bradshaw, G. VR. 1961. A life history study of the California leaf-nosed bat, Macrotus
californicus. Unpublished Ph.D. dissertation, Univ. Arizona, 89 pp.

- . 1962. Reproductive cycle of the California leaf-nosed bat, Macrotus californicus.

Science, 136:645-646.

De Bonilla, H„ and J. J. Rasweiler, IV. 1974. Breeding activity, preimplantation
development, and oviduct histology of the short-tailed fruit bat, Carollia ,
in captivity. Anat. Rec., 179:385-404.

Fleming, T. H. 1971. Artibeus jamaicensis: delayed embryonic development in
a Neotropical bat. Science, 171:402-404.

Hamlett, G. W. D. 1935. Notes on the embryology of a phyllostomatid bat. Amer.
J. Anat., 56:327-349.

Quintero, F., and J. J. Rasweiler IV. 1974. Ovulation and early embryonic
development in the captive vampire bat, Desmodus rotundus. J. Reprod.
Fert., 41:265-273.

Rasweiler, J. J., IV. 1972. Reproduction in the long-tongued bat, Glossophaga
soricina. I. Preimplantation development and histology of the oviduct. J. Reprod.
Fert., 31:249-262.

- . 1974. Reproduction in the long-tongued bat, Glossophaga soricina. II. Im¬
plantation and early embryonic development. Amer. J. Anat., 139:1-36.

Tamsitt, J. R„ and D. Valdivieso. 1963. Reproductive cycle of the big fruit-eating
bat, Artibeus lituratus Olfers. Nature, 198:104.

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SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

- . 1965. Reproduction of the female big fruit-eating bat, Artibeus lituratus

palmarum, in Colombia. Caribbean J. Sci., 5:157-166.

Wimsatt, W. A. 1942. Survival of spermatozoa in the female reproductive tract
of the bat. Anat. Rec., 83:299-307.

- . 1954. The fetal membranes and placentation of the tropical American vampire

bat Desmodus rotundus murinus with notes on the histochemistry of the placenta.
Acta Anat., 21:285-341.

- . 1958. The allantoic placental barrier in chiroptera: a new concept of its

organization and histochemistry. Acta Anat., 32:141-186.

Wislocki, G. B., and D. W. Fawcett. 1941. The placentation of the Jamaican bat
(Artibeus jamaicensis parvipes). Anat. Rec., 81:307-317.

ONTOGENY AND MATERNAL CARE

D. G. Kleiman and T. M. Davis

Although many aspects of phyllostomatid biology have received increasing
attention in recent years, there is still a dearth of information on the growth and
behavioral ontogeny of this diverse family of bats. This is in contrast with
studies of the Vespertilionidae, where both field and laboratory investigations
of development have been common, although by no means numerous (Jones,
1967; Pearson et ai, 1952; Kleiman, 1969; Orr, 1970; Gould, 1971). The
lack of interest in chiropteran ontogeny is discouraging because the special
adaptations for flight, including echolocation, and diverse feeding strategies
should provide fertile ground for developmental studies, as Gould (1970)
has pointed out.

In this chapter we will attempt to review some aspects of ontogeny in the
phyllostomatid bats, concentrating on growth and development in Carollia
perspicillata, which we have studied in captivity. Field and laboratory observations
of other species will be included where they are available. The vampire bat,
Desmodus rotundus, is the only other phyllostomatid for which detailed in¬
formation is available (Schmidt and Manske, 1973).

The colony of Carollia perspicillata was originally captured in Trinidad in
April 1972 and maintained at Johns Hopkins University for six months by
E. Gould. During this period, several births occurred. Sixteen Carollia were
brought to the National Zoological Park, Washington, D.C., in October 1972.
At this time, one female had a small infant; a second female gave birth three
days after the arrival of the colony. Both young were reared. Table 1 presents
the history of the colony between January 1973 and January 1974. Three
Glossophaga soricina (two males, one female) were acquired with the Carollia,
of which one adult male died and one male was born. Nine Anoura geoffroyi
(four males, five females) also were received, but all but a pair died within
the first three days. No breeding of Anoura occurred.

The colony was housed in a climate-controlled room measuring approximately
3 by 3 by 2.5 meters. Temperatures averaged 29 °C (range 27 to 31 °C); relative
humidity, 70 per cent (range 50 to 80 per cent). A light cycle of 12 hours of
light to 12 hours of dark was used. Two wire mesh cages with wooden frames
and burlap covers were provided for roosts in an elevated position. Several
branches were placed between the roosts and from the roosts to the floor.

Bats were fed a peach-nectar mixture developed by Rasweiler and De Bonilla
(1972) for nectarivorous phyllostomatids, although there is evidence that
Carollia also feeds on insects (Pine, 1972; Ayala and D’ Alessandro, 1973).
Water was available ad libitum, as were ripe, peeled bananas that were
suspended from branches. Dishes with the nectar diet were placed in brackets

387

388

SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

Table 1. — History o/Carollia perspicillata colony from January 1973 to January 1974.

Males

Females

Total

Number of original adults

Number of births

Number of deaths:

Adults

Juveniles

*One mother and young died accidentally.

attached to the outside of the roosts so that bats could feed while in flight or
while hanging on the roost.

Bats were caught with butterfly nets; adults initially were examined
bimonthly beginning in January 1973, but weekly examinations were instituted
in April 1973. Young were weighed and measured every two to four days.
Individuals were identified by a number punch marked on the wing membrane
(Bonaccorso and Smythe, 1972; Kleiman and Davis, 1974). Behavioral
observations and retrieval tests were conducted at irregular intervals.

Reproductive Cycle

After the two births in October 1972, there were three birth peaks in
Carollia: February 1973, June and July 1973, and November and December
1973 (Table 2). Known interbirth intervals ranged from 115 to 173 days.
During the first peak of parturition, females were highly synchronized — nine
of 1 1 females gave birth within a 17-day period. The births were more scattered

Table 2. — Dates of birth and interbirth intervals for 12 Carollia perspicillata females,

between January 1973 and January 1974.

No. of
females

Birth no. 1

Birth no. 2

Birth no. 3

Interbirth

interval

(days)

12 Feb. 73

1 July 73

25 Oct. 73

138,

116

26 Jan. 73

29 May 73

18 Nov. 73

123,

173

28 Feb. 73*

23 June 73

6 Dec. 73*

115,

165

20 Feb. 73*

21 June 73*

121

14 Feb. 73*

5 Mar. 73

21 July 73*

10 Dec. 73

138,

142

16 Feb. 73*

3 Aug. 73*

168

12 Feb. 73

24 June 73

8 Nov. 73*

132,

137

18 Feb. 73*

1 July 73

15 Nov. 73*

133,

137

22 Feb. 73*

12 July 73*

7 Dec. 73*

140,

148

22 Feb. 73*

20 June 73*

10 Dec. 73

118,

173

14 Jan. 74*

"■Indicates

accurate birth date. Other

dates are estimated

and parturition might

have occurred a

maxi-

mum of three days earlier.

BIOLOGY OF THE PHYLLOSTOMATIDAE

389

Pre-partum Post-partum

Weeks

Fig. 1. — Average weights and ranges of weights in pre and postpartum Carollia
perspicillata, based on 25 births of 1 1 females.

during the succeeding two parturition periods. The shortest interbirth intervals
ranged from 115 to 123 days (N = 5). Rasweiler and De Bonilla (1972)
found an implanting blastocyst in a female killed 21 days postpartum,
suggesting that estrus may occur shortly after parturition. If an immediate
postpartum heat occasionally occurs, the gestation period for Carollia perspicillata
may be approximately 115 to 120 days. The single Glossophaga soricina female
gave birth in March 1973 and did not become pregnant again for a full year.

A total of 30 Carollia young were born (see Table 1), of which 24 survived
through weaning. No females aborted nor were any infants rejected after birth.
The majority of juvenile deaths occurred at weaning, and at least four of these
might have resulted from feeding on spoiled food or a disfunction in the
humidity control, which caused a rapid drop in humidity in the flight room.
Adult losses were limited to a single female and her young, which died
accidentally.

390

SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

Females gained approximately one-third of their initial weight during
pregnancy (see Fig. 1). Average weight during the last week of pregnancy was
22.9 grams as compared with 17.3 grams during the first week postpartum.
During the final weeks of pregnancy, females were reluctant to fly and
maneuvered less efficiently when they flew. Fetuses were palpable from
about five to six weeks before birth, and were in a transverse position.

The nipples of pregnant females were not obvious prior to birth, but within
two days of parturition the surrounding fur had been shed and the mammary
region had become pink in color. Thick milk could be expressed from the
nipples up until approximately 33 days postpartum (range 21 to 49 days).
Thereafter, the milk began to thin, but fluid could be expressed until
approximately 56 days after birth (range 42 to 72 days). The area around
the nipples began to assume a darker pigmentation and the fur began to re¬
appear from 48 days postpartum (range 37 to 66 days); however, the mammary
region did not assume prepartum condition until 72 days postpartum (range
64 to 87 days). From these observations, it would appear that heavy lactation
continues for slightly over one month after birth, but females continue to
produce milk until approximately 1.5 to 2 months postpartum.

Data available for length of lactation in other phyllostomatids indicate a
lactation period of one to two months (Jenness and Studier, 1976). In the
vampire bat, Desmodus, nursing may continue for nine months although
weaning is initiated at three (Schmidt and Manske, 1973). In Macrotus and
Leptonycteris , lactation continues for one month and four to eight weeks,
respectively (see Jenness and Studier, 1976). A single Glossophaga soricina
female in our colony continued lactating for approximately two months.

Maternal Care

No births were observed in Carollia although females were seen eating
placentas and licking newborn young. The umbilical cord was rarely severed
at the base, but usually dried up and fell off within a day following birth.

Parturition has been described for Stenoderma rufum (Tamsitt and
Valdivieso, 1966b), Artibeus lituratus, Glossophaga soricina, Vampyrops
helleri (Tamsitt and Valdivieso, 1965), and Choeronycteris mexicana
(Barbour and Davis, 1969). In all species, parturition occurred in the normal
head-down position; this seems to be typical of phyllostomatids but rare in
vespertilionids (Wimsatt, 1960), except for Nyctalus noctula (Kleiman, 1969).

In the species observed by Tamsitt and Valdivieso (1965, 1966b), a head
presentation was found. Placentophagia has not been reported for the above-
mentioned species, nor for Desmodus (Schmidt and Manske, 1973).

During the first few days, young Carollia were carried parallel to the mother’s
body and held under the wing. Thereafter, the typical carrying position, both
at rest and in flight, was cross-wise on the mother’s ventral surface, just posterior
to the throat. Carollia infants (up to 14 days) were rarely observed hanging
alone. Young attached themselves primarily with the mouth and hind feet;

BIOLOGY OF THE PHYLLOSTOMATIDAE

391

the wings were tightly closed and partially covered the infant’s body. Claws on
the thumbs were not used for clinging because the distal portion of the forearm
was pressed tightly against and covered the infant’s head and ears. Young removed
from the mother’s nipple occasionally remained in this carrying posture for
several seconds, even when placed on their back. The cross-wise carrying
posture was also seen in our individual of Glossophaga soricina, Desmodus
(see fig. 2 in Schmidt and Manske, 1973), and might be present in Choeronycteris
(see fig. 8 in Barbour and Davis, 1969). It appears to be an adaptation for carrying
young while the female is flying. For the first 10 days, captive young of Artibeus
were reported (Novick, 1 960) to hang head down under the mother’s wing with the
hindfeet around the mother’s thigh.

Carollia mothers preferred to hang freely from a horizontal ceiling when
carrying attached young. Thus, it was impossible for infants to be attached
simultaneously to the nipple and support themselves by the hind feet until
they were about half the size of the mother. Young were capable of hanging
from the ceiling by the age of 18 days, but still remained attached to a nipple.
Similar observations were made of a young Glossophaga. In Desmodus, young
do not support themselves until at least two weeks of age (Schmidt and Manske,
1973).

From our observations, it appeared that resting Carollia females supported
the bulk of their infant’s weight for at least 14 days. An added advantage to
the cross-wise carrying position assumed by the young, other than providing
balance, was that they did not need to readjust their position when a female
flew. Young were last observed attached to the mother approximately 23.5
days postpartum (range 19 to 31; N = 15), when they were approximately
57 per cent of the mother’s weight.

Because we were unable to observe the bats without disturbing them,
especially at night, we do not know whether females foraging in the wild carry
their young or, if they do, for how long. One 1 1 -day-old young was seen
hanging alone next to its mother approximately 45 minutes after the lights
went out, but the infant attached to the nipple and moved back into a cross-wise
carrying position immediately after we entered the room. The mother flew
as soon as the young attached. This suggests that mothers may detach from the
young at night, but we had no evidence that young were ever left in a creche.
Mothers with attached young were more reluctant to fly when disturbed than
were unencumbered bats but did so, nevertheless, and seemed able to
maneuver efficiently.

Observations of development in a single young Glossophaga soricina were
similar to those for Carollia. The young was last seen attached to the mother
when it was 20 days old.

Both from our Carollia observations and some field reports, it appears
as though some species of phyllostomatid bats commonly carry their young
and, unlike vespertilionids (see Fenton, 1969; Davis, 1970), do not leave them
in creches.

392

SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

Felten (in Pine, 1972) apparently netted a Carollia perspicillata with a
half grown young, and Tamsitt and Valdivieso (1963 a) caught lactating
Artibeus lituratus and Glossophaga sorcina carrying young in the vicinity of
fruit trees where presumably they were foraging. One A. lituratus female
carried a young 53.8 per cent of her weight (Tamsitt and Valdivieso, 1965).
Mumford and Zimmerman (1964) reported netting lactating Choeronycteris
mexicana with attached young at a distance of approximately 200 yards from
the main daytime roost. Bradshaw (1961) captured a female Macrotus
californicus in a roost carrying a young weighing 57 per cent of her weight;
Cockrum (in Davis, 1970) observed female Leptonycteris sanborni moving
young within a cave as well as carrying advanced young to a previously
abandoned roost. Schmidt and Manske (1973) indicated that Desmodus
females can carry young up to eight weeks old. A. M. Greenhall (personal
communication) has observed Desmodus females with attached young of
unknown age feeding on cattle; however, these bats were similar in size to
young that he had observed crawling around in roosts without the mothers.
These young were not newborn and might have been approaching weaning
age.

Observations discussed above suggest that phyllostomatids may carry
attached young of an advanced age. Whether females forage with the young or
simply move them from roost to roost remains to be determined. Certainly,
except for Macrotus waterhousii (Goodwin, 1970), Leptonycteris sanborni
(Hoffmeister, 1959), and Phyllostomus hastatus (J. Bradbury, personal
communication), one does not find reports of creches of infants in phyllostomatids,
although lactating females may roost colonially and segregate themselves
from males. Bradbury (personal communication) suggested that female
Carollia, for example, may move their babies from the day roost to a night
roost prior to foraging, which may partly account for the well-developed
tendency to carry young in captivity.

Retrieval of young Carollia was observed under several experimental
conditions. Mothers and young were released into a small holding cage after
being weighed and measured; typically, they reestablished contact within
30 minutes to an hour (that is, before being released into the flight room).
On several occasions, young were deliberately separated and hung on the
outside of the roost, after which time the other bats were released into the
flight room. Several different bats would fly past hanging infants, pausing
briefly to hover, as though to inspect the young. Usually, a juvenile was inspected
several times (both by its mother and other bats) before the mother would
alight above her offspring and crawl down to it.

Juveniles that were too young to fly were never observed attempting to regain
contact with their mother by climbing higher on the roost. Normally, they
hung motionless until the mother made tactile contact with them. Audible
vocalizations (ultrasonic calls were given by the mother and young, Gould,
1975) were not heard nor did the infant reveal much sign of disturbance.
Licking of the young by the mother usually accompanied retrieval, especially

BIOLOGY OF THE PHYLLOSTOMATIDAE

393

before the mother flew again. The latency to retrieve was highly variable
in the females, ranging from two to 30 minutes. The age of the young did not
seem to affect this latency because infants between one and three days old
were retrieved within two to 22 minutes.

Mothers clearly recognized their own offspring; we never caught a female
with an alien young attached to her. Moreover, mothers and young retained
an association (roosted near each other) long after weaning. One Carollia
mother and daughter were regularly caught together until the daughter was
five months old, about a week prior to the next birth.

Development of Young

Carollia are born in an advanced state, with the eyes open (Fig. 2).
Neonates are fully furred on the dorsum, and the more sparsely furred venter
and muzzle become covered within two to three days after birth. The dark
brown juvenile pelage is complete by day 7 to 10.

Of the neonatal phyllostomatids observed, Macrotus, Leptonycteris
(Gould, 1975), Carollia, Glossophaga (this study and Klfma and Gaisler,
1968), Choeronycteris (Mumford and Zimmerman, 1964), and Artibeus
(Tamsitt and Valdivieso, 1966a) are born well furred. Desmodus (Schmidt
and Manske, 1973; Gould, 1975), Phyllostomus discolor (Klima and Gaisler,
1968), and P. hastatus (Gould, 1975) are sparsely furred at birth.

Eyes are open at birth in Carollia (this study), Artibeus (Tamsitt and
Valdivieso, 1966a), Desmodus (Schmidt and Manske, 1973; Gould, 1975),
Macrotus (Gould, 1975), and Phyllostomus hastatus (Gould, 1975). Only
Leptonycteris and Phyllostomus discolor have been reported (Tamsitt and
Valdivieso, 1963 a) to have the eyes closed at birth.

Carollia neonates were active from birth and when handled would
squirm, try to crawl away, and often vocalize. This contrasted with their
behavior in the flight room during retrieval tests when they hung motionless
on the bat roost. The increased activity might have been caused by the
temperature of the room in which weights and measurements were taken,
which was cooler than was the flight room. Gould (1975) stated that the young
of Desmodus, Phyllostomus hastatus, and Leptonycteris sanborni are active
during reunions with the mother, whereas those of Macrotus californicus
are passive.

C. perspicillata young are born with a complete set of 22 deciduous teeth,
with the formula di 2-2/2 -2, dc 1 -1/1-1, dpm 3-3/2-2 = 22. A comparison
of preserved skulls from the U.S. National Museum with living neonates suggests
that only 16 of the 22 deciduous teeth are functional. The four lower incisors,
barely penetrating the gingivum, disappear several days after birth, and the first
upper deciduous premolars are not even visible in live specimens. Lower deciduous
premolars are simple, highly reduced spicules, undifferentiated in width from root
to crown. The second and third upper premolars, although more prominent than
the lower ones, are tiny pegs that taper to a fine point at the crown. The second milk

394

SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

Fig. 2. — Neonate of Carollia perspicillata on the day of birth. Note that the
eyes are open, and the animal’s dorsum is fully furred. The venter typically has only
sparse fur.

BIOLOGY OF THE PHYLLOSTOMATIDAE

395

Fig. 3. — The loss of deciduous teeth in juvenile Carollia perspicillata. Observations
within a given time period may include the same individual. (Symbols are closed circles,
upper outer incisors; open circles, upper inner incisors; closed triangles, lower canines;
open triangles, upper canines; closed squares, lower premolars; and open squares,
upper premolars).

premolar is weakly recurved. Lower canines are slender, mildly recurved spicules
that gradually taper to a point. The upper canines and upper outer incisors are the
largest, most strongly recurved stylettes; also, they are retained longest. Upper
inner incisors are bifid at the distal extremity.

The comparative rate of loss of deciduous teeth is represented in Fig. 3.
Lower deciduous premolars are lost during the first two weeks postpartum.
Lower canines, upper premolars, and upper inner incisors are shed next. The
upper canines and upper outer incisors are retained until one month postpartum,
the last milk tooth being lost at 34 days postpartum. The permanent dentition
of the upper jaw emerged first. By day 22 postpartum, one-third of the
permanent teeth had emerged; by day 26, two-thirds; and by day 31, all were
present.

Deciduous upper and lower canines and upper outer incisors are the teeth
primarily used to attach to a nipple. Two observed perforations in a female’s
nipple were a clear result of the upper canines, the distances between the
perforations and the canines both measuring 2.6 millimeters. Carollia
resembles Tonatia, Mimon, Chrotopterus, Choeronycteris, and Phyllostomus
in that the upper outer incisors are more prominent than the upper inner ones.

396

SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

In Macrotus, Glossophaga, and Leptonycteris, both outer and upper inner
incisors are functional (Phillips, 1971).

In general, the deciduous dentition of most phyllostomatids is reduced
and less complex than that of vespertilionids (Phillips, 1971; Miller, 1907).
This seems to correlate with the tendency to carry attached young rather than
deposit them in creches, thus suggesting that increased complexity in the
deciduous dentition of vespertilionids may function to grasp the returning
mother (or any female in species that nurse promiscuously) rather than to main¬
tain a hold on the nipple when already attached.

The development of flight in C. perspicillata was investigated by periodically
dropping infants and juveniles. Prior to day 14, all young drop straight to the
ground, with the wings extended. As infants approached 14 days of age, they
occasionally flapped their wings once or twice as they fell. Between days 14 and
16, young bats began flapping the wings when dropped, but could not maintain
altitude or turn. They also were unable to land and often collided with obstacles or
eventually dropped to the floor. By day 1 8, they could maintain (and gain) altitude,
take off from a roosting position, turn, and avoid obstacles. However, their landing
ability was poor, and they often landed with the wings extended. Between days 20
and 23, the ability to land upside-down with the wings folded perfected, and,
after day 24, flight development essentially was complete. Juveniles, however,
could be distinguished from adults by their flight patterns for several weeks more
because they flew more slowly and erratically. Juveniles were first captured inde¬
pendent of the mother on an average of 27.6 days (range 23 to 31, N= 16) after
birth.

There is little information available on flight development in other young
phyllostomatids. In Desmodus, young achieve flight capability at eight to ten
weeks of age (Schmidt and Manske, 1973); Novick (1960) reported that a
young Artibeus began to fly at approximately 28 days of age. A single juvenile
Glossophaga soricina was first found separate from its mother and flying at
age 25 to 28 days.

Neonates of Carollia perspicillata average 5.0 grams at birth (range 4.1 to
5.9; N= 13), which is 28.4 per cent of the postpartum weight of females.
Initial growth in weight is rapid (Fig. 4), but juveniles do not achieve adult
weight until 10 to 13 weeks of age. Forearm length at birth is 24.4 millimeters
(range 22.4 to 27.5 mm; N= 10), and forearm growth essentially is complete
at six weeks (Fig. 4). At approximately 24 days of age, when the young first
begin to fly, forearm length is 93.4 per cent and weight 63.0 per cent of that
for adults (N= 10).

Neonatal and postpartum weights and measurements are not available for
most phyllostomatid bats. Table 3 presents some accurate and estimated
neonatal to mother weight and measurement ratios for both phyllostomatid
and vespertilionid bats, based on known and derived data. Weights and
measurements were taken from full-term fetuses and nonlactating females.
Young- to-mother weight ratios are poor for comparative purposes because
weights tend to fluctuate seasonally, captive and field weights frequently

BIOLOGY OF THE PHYLLOSTOMATIDAE

397

Days Weeks

Age

Fig. 4. — Increase in average weight (bottom) and forearm length (top) for Carollia
perspicillaux. A, average day when young were last observed attached to the mother;
B, average day when the mother’s milk began to thin; C, average day when milk no longer
could be expressed from the mother's nipples. These averages are based on measurements
from 17 individuals (8 females, 9 males) of known age. The open squares indicate the
mean weight and forearm length (and range) for 12 adult males for comparison.

differ, and species may have one to three young per litter. However, most
phyllostomatids exhibit ratios greater than 0.25 (for single births). Orr (1970)
noted that the ratio in vespertilionids depends on species size, larger species
tending to have a smaller ratio. Neonatal-to-mother forearm ratios are a
better comparative measure. Table 3 indicates that phyllostomatid bats may
be born in a more advanced stage than vespertilionids because seven of eight
species of phyllostomatids have a ratio usually exceeding 0.41 whereas this
ratio is exceeded in only three of 1 3 vespertilionids.

Discussion and Conclusions

The paucity of information on phyllostomatid development not withstanding,
available data suggest that ontogeny and maternal care in phyllostomatids
differs in several characteristics from those in vespertilionids.

Table 3. _ Average neonate to adult weight and forearm length ratios in selected phyllostomatid and vespertilionid bats. Weights are given

398

SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

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BIOLOGY OF THE PHYLLOSTOMATIDAE

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1. Phyllostomatids generally are born in a more precocial condition,
(furred, eyes open, mobile, and size large relative to that of the mother) than
vespertilionids. As Gould (1975) pointed out, there is no clear dividing line
between altriciality and precociality, but within the two families, the degree
of overlap in such characteristics as mobility, eye opening, and pelage
development is small.

2. In phyllostomatids, deciduous teeth are reduced in size, relatively simple
in form, and functional teeth are fewer in number. The deciduous dentition
might be related to permanent dentition and different feeding strategies, but
it might also correlate with maternal care patterns, as discussed in point 3
below.

3. Phyllostomatid young usually are not deposited in large creches by
foraging mothers. Instead, they remain attached to the mother in the roost
during the day and might be carried during foraging. Bradbury (personal
communication) suggested that young might be carried to a nocturnal roost
before the female begins to forage. The cross-wise position assumed by attached
young could be an adaptation of phyllostomatids to frequent carrying by the
mother.

The occurrence of these three characteristics in many species of phyllostomatid
bats is intriguing, especially when considering how such adaptations evolved.
Carrying young during foraging or transferring young to individual nocturnal
roosts before foraging could serve as an antipredator strategy for bats living under
conditions where other bats have evolved as predators. However, transferral
to a nocturnal roost might be an adaptation that could evolve only under stable
tropical conditions where temperature fluctuations are not great. By contrast,
creches of vespertilionids might function, in part, to retain heat in the altricial
young. Clearly, behavioral studies in the field are needed to determine how onto-
gency and maternal care in the Phyllostomatidae relate to feeding strategies, social
organization, roosting behavior, and possible antipredator mechanisms.

Acknowledgments

We are grateful to E. Gould for providing us with the captive colony of
Carollia perspicillata, Anoura geoffroyi, and Glossophaga soricina. The
study specimens examined from the United States National Museum of
Natural History, Smithsonian Institution, included five Carollia perspicillata
and two C. brevicaudatum (USNM 104551; 104552; 179612; 284511-
284513; 65475). We thank R. H. Pine for his assistance. The literature review
for this article was completed in early 1975.

Literature Cited

Ayala, S. C., and A. D’Alessandro. 1973. Insect feeding of some Colombian fruit¬
eating bats. J. Mamm., 54:266-267.

Barbour, R. W., and W. H. Davis. 1969. Bats of America. Univ. Press Kentucky,
Lexington, 286 pp.

BIOLOGY OF THE PHYLLOSTOMATIDAE

401

Bogan, M. A. 1972. Observations on parturition and development in the hoary bat,
Lasiurus cinereus. J. Mamm., 53:61 1-613.

Bonaccorso, F. J., and N. Smythe. 1972. Punch-marking bats: an alternative to
banding. J. Mamm., 53:389-390.

Bradshaw, G. V. R. 1961. A life history study of the California leaf-nosed bat,
Mcicrotus californicus. Unpublished Ph.D. dissertation, Univ. Arizona,
Tucson, 89 pp.

Burns, R. J. 1970. Twin vampires born in captivity. J. Mamm., 51:391-392.

Crespo, R. F., R. J. Burns, and S. B. Linhart. 1970. Loadlifting capacity of the vampire
bat. J. Mamm., 51:627-628.

Davis, R. 1969. Growth and development of young pallid bats, Antrozous pallidus.
J. Mamm., 50:729-736.

- . 1970. Carrying of young by flying female North American bats. Amer.

Midland Nat., 83:186-196.

Davis, W. H., R. W. Barbour, and M. D. Hassell. 1968. Colonial behavior of Eptesiats
fuscus. J. Mamm., 49:44-50.

Fenton, M. B. 1969. The carrying of young by females of three species of bats. Canadian
J. Zool., 47:158-159.

Goodwin, G. G., and A. M. Greenhall. 1961. A review of the bats of Trinidad and
Tabago. Bull. Amer. Mus. Nat. Hist., 122:195-301.

Goodwin, R. E. 1970. The ecology of Jamaican bats. J. Mamm., 51:571-579.

Gould, E. 1970. Echolocation and communication in bats. Pp. 144-161, in
About bats (B. H. Slaughter and D. W. Walton, eds.), Southern Methodist
Univ. Press, Dallas, vii + 1-339.

- . 1971. Studies of maternal-infant communication and development of

vocalizations in the bats Myotis and Eptesicus. Comm. Behav. Biol., 5:263-313.

- . 1975. Neonatal vocalizations in bats of eight genera. J. Mamm., 56:15-29.

Hoffmeister, D. F. 1959. Distributional records of certain mammals from

Southern Arizona. Southwestern Nat., 4: 14-19.

Jenness, R„ and E. H. Studier. 1976. Lactation and milk. Pp. 201-218, in
Biology of bats of the New World family Phyllostomatidae. Part 1 (R. J. Baker,
J. K. Jones, Jr., and D. C. Carter, eds.), Spec. Publ. Mus., Texas Tech
Univ., 10:1-218.

Jennings, W. L. 1958. The ecological distribution of bats in Florida. Unpublished
Ph.D. dissertation, Univ. Florida, Gainesville, 126 pp.

Jones, C. 1967. Growth, development, and wing-loading in the evening bat
Nycticeius humeralis. J. Mamm., 48:1-19.

Kleiman, D. G. 1969. Maternal care, growth rate, and development in the noctule
( Nyctalus noctula), pipistrelle ( Pipistrellus pipistrellus), and serotine
{Eptesicus serotinus) bats. J. Zool., 157:187-211.

Kleiman, D. G., and T. M. Davis. 1974. Punch-mark renewal in bats of the genus
Carollia. Bat Research News, 15:29-30.

Klima, M., and J. Gaisler. 1968. Study on growth of juvenile pelage in bats, III. Phyl¬
lostomatidae. Zoolog. Listy, 17:1-18.

Kunz, T. H. 1973. Population studies of the cave bat ( Myotis velifer ): reproduction,
growth, and development. Univ. Kansas Occas. Papers Mus. Nat. Hist.,
15:1-43.

Lane, H. K. 1946. Notes on Pipistrellus subflavus subflavus (F. Cuvier) during the
season of parturition. Proc. Pennsylvania Acad. Sci., 20:57-61.

Medway, L. 1972. Reproductive cycles of the flat-headed bats, Tylonycteris pachypus
and T. robustula (Chiroptera: Vespertilioninae) in a humid equatorial environ¬
ment. Zool. J. Linn. Soc., 51:33-61.

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SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

Miller, G. S., Jr. 1907. The families and genera of bats. Bull. U. S. Nat. Mus.,
57:1-282.

Mumford, R. E., and D. A. Zimmerman. 1964. Notes on Choeronycteris mexicana. J.
Mamm., 43:101-102.

Novick, A. 1960. Successful breeding in captive Artibeus. J. Mamm., 41:508-509.

O'Farrell, M. J., and E. H. Studier. 1973. Reproduction, growth, and development
in Myotis thyscinodes and M. lucifugus (Chiroptera: Vespertilionidae). Ecology,
54:18-30.

Orr, R. T. 1954. Natural history of the pallid bat, Antrozous pall id us ( LeConte). Proc.
California Acad. Sci., 28:165-246.

- . 1970. Development: prenatal and postnatal. Pp. 217-231, in Biology of

Bats (W. A. Wimsatt, ed.). Academic Press, New York, 1:406 pp.

Pearson, O. P., M. A. Koford, and A. K. Pearson. 1952. Reproduction of the
lump-nosed bat ( Corynorhinus rafinesquii) in California. J. Mamm., 33:273-320.

Phillips, C. J. 1971. The dentition of the glossophagine bats: development, morphological
characteristics, variation, pathology and evolution. Misc. Publ. Mus. Nat.
Hist., Univ. Kansas, 54:1-138.

Pine, R. H. 1972. The bats of the genus Carollia. Tech. Monogr., Texas Agric.

Exp. Sta., Texas A&M Univ., 8:1-125.

Rasweiler, J. J., and H. De Bonilla. 1972. Maintaining nectarivorous phyllostomatid
bats in the laboratory. Lab. Anim. Sci., 22:658-663.

Schmidt, U., and U. Manske. 1973. Die Jugendentwicklung der Vampirfledermause
(Desmodus rotundus). Z. Saugetierk., 38:14-33.

Tamsitt, J. R., and D. Valdivieso. 1963a. Records and observations on Colombian
bats. J. Mamm., 44:168-180.

- . 1963 6. Reproductive cycle of the big fruit-eating bat, Artibeus lituratus,

Olfers. Nature, 198:194.

- . 1965. Reproduction of the female big fruit-eating bat, Artibeus lituratus

pal mar um, in Colombia. Caribbean J. Sci., 5:157-166.

- . 1966a. Taxonomic comments on Anoura caudifer, Artibeus lituratus, and

Molossus molossus. J. Mamm., 47:230-238.

- . 19666. Parturition in the red fig-eating bat, Stenoderma rufum. J. Mamm.,

47:352-353.

Wimsatt, W. 1960. An analysis of parturition in Chiroptera, including new observations
on Myotis l. lucifugus. J. Mamm., 41:183-200.

GENERAL PHYSIOLOGY

John M. Burns

At first exposure to this volume, as well as its previous companions, one is
amazed at the amount of information that has accumulated concerning the biology
of New World leaf-nosed bats. Upon closer inspection, however, it is apparent
that the vast majority of this information deals with taxonomy, distribution,
natural history, and various aspects of morphology. Physiological study of these
biologically important mammals has been a neglected area, at least as judged by
the published literature.

Two physiological systems that have been examined to a substantial degree and
warrant separate consideration are sensory physiology (primarily echolocation)
and thermoregulation. Gould (1977) and McManus (1977) have provided excel¬
lent reviews of these respective topics and I shall not atterqpt to duplicate here
the information presented in these two papers.

Endocrine studies, in particular, are lacking for phyllostomatids. The reason
for this probably can be attributed to the fact that bats are small and therefore
have small blood volumes. Until the last decade, measurements of hormone
concentrations were dependent mostly on bioassays that required blood to be
pooled from several bats. Determination of hormone concentration is no longer
a major problem because such techniques as radioimmunoassay (RIA) and
fluorescent immunoenzyme assay require only 50 to 100 microliters of plasma.
Echolocation and thermoregulation studies, on the other hand, have allowed
investigators to work with entire animals without the need for expensive equip¬
ment.

Many interesting questions can be raised as to the role of chiropteran endocrine
systems in such physiological endeavors as water balance, bone and calcium
metabolism, and digestion. For the moment, we can only surmise that such
endocrine regulation is similar to that known for other mammals.

Reproductive Physiology

In view of the great deal of emphasis placed on reproductive physiology of
animals over the past several decades, one would be inclined to suppose that there
is a wealth of such information for phyllostomatid bats. However, the vast
majority of literature on reproduction in leaf-nosed bats deals with studies of
comparative anatomy, morphology, natural history, and fecundity rather than
with the physiological processes of reproduction. As Wilson (this volume)
pointed out, reproductive strategies of phyllostomatids are varied. These include
such schemes as monestry, polyestry, and a system that Bradshaw (1962) termed
delayed development for Macrotus californicus\ a similar system was reported
(Fleming, 1971) for Artibeus jamaicensis. A unique gestation pattern was re¬
ported in Macrotus californicus for thyroid hormone (Bums et al., 1972),

403

404

SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

estrogens (Burns and Wallace, 1975), and for progesterone (Burns and Easley,
1977). In each of these reports, biphasic patterns were described in which one
peak coincided with the fertilization and implantation period of October and
November, followed by a second peak in May and June that corresponded to
fetal maturation and parturition. The hormonal data (summarized in Table 1),
as well as the histological studies of Bleier (1975a, 1975 b), suggest that the
reproductive scheme in M. californicus is quite different from delayed im¬
plantation.

Krutzsch et al. (1976) reported changes in plasma testosterone and testicular
ascorbic acid in reproductively active male M. californicus ; testosterone and
testicular ascorbic acid reached a peak concentration of 2.7 ng/ml and 38 ug/ml,
respectively, in late summer, and spermatazoa were present in the epididymides
from August to early December. The testes began to atrophy by late September,
and the levels of testosterone and testicular ascorbic acid declined by December
but were detectable the year around (minimum concentrations observed for
testosterone were 0.25 ng/ml; ascorbic acid, 1 ug/ml.).

The seminal vesicles and prostate glands were at maximum size in September
(15 mm diameter, 19 mg weight) and slowly digressed beginning in late autumn.

In my studies at Texas Tech University, I also found that M. californicus is
an adaptable animal for laboratory study. After individuals are fed by hand for
2 to 3 days, they are tamed quite rapidly. When bats were housed in large cages,
which allow for adequate freedom of flight, attempts to establish breeding colonies
proved successful (unpublished).

Thyroid

Reports on thyroid physiology are scarce and usually play a minor role in larger
studies related to thermoregulation or reproduction. Sadler and Tyler (1960a)
examined thyroid function in a nonhibernating bat, Macrotus californicus , by
means of 131 1 uptake. Animals were tested over a temperature range of 24° to
37 °C, and it was found that chronic exposure to these temperatures did not
influence thyroid activity. This is quite different from responses of hibernating
species of vespertilionids, which show drastic changes in the rate of thyroid
uptake of radioactive iodine when subjected to a similar temperature regime as
described above for M. californicus (Sadler and Tyler, 1960 b).

Burns et al. (1972) reported a drastic decrease in plasma thyroxine for M.
californicus during the second trimester of pregnancy (see Table 1). It was found
later, however, that triiodothyronine (T3) levels were elevated throughout the
gestation period to the extent that total thyroid hormone concentration in the
blood during pregnancy remained essentially unchanged (unpublished).

Adrenal Glands

Studies that attempt to describe the role of either adrenocortical or medullary
hormones in regulating a host of physiological processes in phyllostomatid bats
are unknown. Such a work would represent a “first” for comparative physiology.

BIOLOGY OF THE PHYLLOSTOMATIDAE

405

Table 1 . — Changes in plasma concentrations of various hormones during pregnancy /nMacrotus

californicus.

Hormone

Aug.

Sept.

Oct.

Nov.

Dec.

Jan.

Feb.

Mar.

Apr.

May

June

Thyroxine1

3.5

2.0

1.0

2.0

5.0

Progesterone2

1 1

Estrone3

Estradiol- 17 6

■Thyroxine concentrations were measured by a column chromatography-colorimeter technique and are
expressed as microgram per cent. The June sample is from lactating females. From Burns et at. (1972).

2Progesterone concentrations were determined by radioimmunoassay and are expressed as nanograms
per milliliter. The June sample represents preparturition samples. From Burns and Easley (1977).

3Estrogens determined by radioimmunoassay and expressed as picograms per milliliter. The values for
August mean that estrogen levels were not detectable with this assay. From Burns and Wallace (1975).

There are a few reports that describe the basic morphology of chiropteran adrenal
glands (for example, see Christian, 1963) but no attempt has been made to
elucidate the role of the adrenal glands in a physiological sense.

Parathyroid Glands

I was unable to find a published report of any investigation that dealt with the
function or particular characteristics of the parathyroid glands in any member
of the Phyllostomatidae.

Renal Physiology

The great success of Chiroptera in general, and phyllostomatids in particular,
suggests that some species have evolved elaborate and highly efficient renal
mechanisms for conserving water. Most studies of renal function pertain to
evaporative water loss, however, and are orientated more toward thermoregulation
than anything else (see McManus, 1977). There is a paucity of information
concerning renal physiology and evidently an absolute absence of data dealing
with the endocrine regulation of renal function in leaf-nosed bats.

McFarland and Wimsatt (1965, 1969) reported on the unusual ability of the
kidneys in the vampire bat Desmodus rotundus to concentrate urine. At first,
one might question the physiological demand that would result in development of
a versatile renal system in an animal with a diet that is approximately 98 per cent
water. McFarland and Wimsatt (1969) proposed that the majority of the water
content ingested with a blood meal must be eliminated rapidly for purposes of
flight. This would result in a meal residue composed almost entirely of protein,
which represents a nitrogen load that must be excreted with a minimum of urinary
water loss. McFarland and Wimsatt (1969) also reported that the vampire bat
concomittantly forms urine at a high rate (4 ml/kg/minute) and a low osmolality
(475 mOs) during feeding. Five to six hours after feeding, the rate of urine pro¬
duction falls to approximately 0.2ml/kg/minute, with a surprising high urine
concentration (4656 mOs). Wimsatt and Guerriere (1962) also reported on the
relationship of volume of blood consumed by D. rotundus to amount of urine

406

SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

excreted. For example, if the blood meal is 35 milliliters, the urine volume excreted
shortly after feeding is approximately 26 milliliters. Also of interest is the
observation (Wimsatt and Guerriere, 1962) that isolated D. rotundus have a
somewhat higher average daily consumption of blood than do bats held captive in
groups (21.2 as compared to 15.5 milliliters). The physiological significance of
these observations is not known.

Whereas Desmodus rotundus demonstrates a remarkable ability to concentrate
urine, the nectarivorous Leptonycteris sanborni has little physiological capability
in this regard. Carpenter (1969) showed that even when individuals of L. sanborni
collected from desert habitats were placed on a high protein diet, the maximum
urine concentration was only 342 mOs. This value is even less concentrated than
that reported by Schmidt-Nielsen and O’Dell (1961) for semiaquatic mammals
such as beaver, Castor canadensis. Normally, L. sanborni feeds on nectar from a
variety of desert plants that are high in water and carbohydrates. Howell (1974)
showed that this species obtains proteins and amino acids by consuming pollen
of the saguaro cactus as a dietary supplement. The pollen’s nitrogenous degradation
products are concentrated in the urine and then actively ingested by the bat. This
behavior results in a positive nitrogen balance, a condition otherwise impossible
on a pollen-free diet.

Respiratory Physiology

Inasmuch as bats lack the more efficient flow-through air sac arrangement
characteristic of birds, they must devote a substantially greater portion of their
body to respiratory surface tissue. For example, the common crow, Corvus
brachyrhynchos, has a respiratory surface area in its lungs of approximately

O. 6 square centimeter per gram of body weight (McCauley, 1971), whereas small
bats, such as those in the vespertilionid genus Myotis, must devote 100 square
centimeters per gram of body weight so as to meet the metabolic demand of
flight. It does not appear, however, that this poses an anatomical disadvantage
for bats because flight is an efficient method of travel for chiropterans. For
example, Thomas (1975) calculated that Phyllostomus hastatus requires
only one-sixth the energy needed by a terrestrial mammal of the same size to
cover a given distance. He also calculated the metabolic rate, in watts, for flying

P. hastatus (0.93 kg) as 130.4 w/kg-1. Thomas also stated that such metabolic
rates are essentially the same as the predicted values for flying birds of similar
body size, but that they are two and a half to three times greater than the highest
metabolic rates of which exercising terrestrial mammals of similar size appear
capable.

Thomas and Suthers (1972) provided some interesting data concerning the
differences in respiration at rest and during flight for Phyllostomus hastatus,
which are summarized in Table 2.

They also reported that the heart rate of preflight P. hastatus was 8.7 beats
per second as compared to 13 beats per second (780 beats per minute) in the
first few seconds of flight. Lastly, Thomas and Suthers recorded the hematocrit

BIOLOGY OF THE PHYLLOSTOMATIDAE

407

Table 2. — Comparison of difference in respiration for resting and flying Phyllostomus hastatus
(from Thomas and Slithers, 1972). Weight is given in grams and metabolic rate in terms of
milliliters of oxygen per gram of body weight per hour.

Weight

Metabolic rate
ml02 (gh)-1

Ventillation rate
(breaths/second)

Before flight

101

6.78 ± 0.85

2.8

6.12 ± 1.15

During flight

101

27.53 ± 0.79

10.6

24.68 ± 1.87

of P. hastatus as 60 per cent. This is considerably greater than the percentage
of red blood cells found in a given volume of blood from any avian species listed
by Sturkie (1965); the higher erythrocyte number probably reflects one of the
general physiological adaptations for flight in bats.

Electrophoretic properties of some phyllostomatid hemoglobins have been
described. Valdivieso et al. (1969) found a single, common hemoglobin band for
Monophyllus redmani, Artibeus jamaicensis , Stenoderma rufum, and Erophylla
bombifrons. A similar, more comprehensive electrophoretic survey was reported
by Mitchell (1966). Additional hematological data for leaf-nosed bats were
reported by Valdivieso and Tamsitt (1971), who concluded that hematocrit
values for frugivorous species are lower than those found in insectivorous bats.

Concluding Remarks

This contribution to the biology of New World leaf-nosed bats is an indication
of what little is known concerning their physiology rather than a survey and
review of a substantial body of knowledge. It also represents perhaps a subtle plea
to comparative physiologists to turn their attention to phyllostomatids. Techniques
now are available for measuring biological molecules in blood samples of small
volume. Hopefully, future investigators will take advantage of this technology.

Literature Cited

Bradshaw, G. V. R. 1962. Reproductive cycle of the California leaf-nosed bat, Macrotus
cal ifornicus. Science, 136:645.

Bleier, W. J. \915a. Early embryology and implantation in the California leaf-nosed
bat, Macrotus californicus. Anat. Rec., 182:237-254.

- . 19756. Fine structure of implantation and the corpus luteum in the California

leaf-nosed bat, Macrotus californicus. Unpublished Ph.D. dissertation, Texas
Tech Univ., Lubbock, Texas, vii+75 pp.

Burns, J. M„ and R. G. Easley. 1977. Hormonal control of delayed development in
the California leaf-nosed bat, Macrotus californicus. III. Changes in plasma
progesterone during pregnancy. Gen. Comp. Endocrinol., 32:163-166.

Burns, J. M., and W. C. Wallace. 1975. Hormonal control of delayed development
in Macrotus waterhousii. II. Radioimmunoassay of plasma estrone and estradiol
17B during pregnancy. Gen. Comp. Endocrinol., 25:529-533.

408

SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

Burns, J. M., R. J. Baker, and W. J. Bleier. 1972. Hormonal control of delayed
development in Macrotus waterhousii. I. Changes in plasma thyroxine during
pregnancy and lactation. Gen. Comp. Endocrinol., 18:54-58.

Carpenter, R. E. 1969. Structure and function of the kidney and the water balance of
desert bats. Physiol. Zool., 42:288-302.

Christian, J. J. 1963. Endocrine adaptive mechanisms and the physiologic regulation of
population growth. Pp. 189-353, in Physiological mammalogy, (R. G. Van
Gelder and W. Mayer, eds.). Academic Press, New York, l:xii+ 1-381.

Fleming, T. H. 1971. Artibeus jamaicensis : delayed embryonic development in a
neotropical bat. Science, 171:402-404.

Gould, E. 1977. Echolocation and communication. Pp. 247-279, in Biology of bats
of the New World family Phyllostomatidae. Part II (R. J. Baker, J. K. Jones, Jr.,
and D. C. Carter, eds.), Spec. Publ. Mus., Texas Tech Univ., 13:1-364.

Howell, D. J. 1974. Bats and pollen: physiological aspects of the syndrome of chi-
ropterophily. Comp. Biochem. Physiol., 48A:263-276.

Krutzsch, P. H., R. W. Watson, and C. P. Lox. 1976. Reproductive biology of the male
leaf-nosed bat, Macrotus waterhousii, in the southwestern United States. Anat.
Rec., 184:611-636.

McCauley, W. J. 1971. Vertebrate physiology. W. B. Saunders Co., Philadelphia,
xiv + 422 pp.

McFarland, W. N., and W. A. Wimsatt. 1965. Urine flow and composition in the
vampire bat. Amer. Zool., 5:662.

- . 1969. Renal function and its relation to the ecology of the vampire bat, Desmodus

rotundas. Comp. Biochem. Physiol., 28:985-1006.

McManus, J. J. 1977. Thermoregulation. Pp. 281-292, in Biology of bats of the New
World family Phyllostomatidae. Part II (R. J. Baker, J. K. Jones, Jr., and D.
C. Carter, eds.), Spec. Publ. Mus., Texas Tech Univ., 13:1-364.

Mitchell, H. A. 1966. Multiple haemoglobins in bats. Nature, 210:1067-1068.

Sadler, W. W., and W. S. Tyler. 1960a. Thyroidal activity in hibernating Chiroptera.
I. Uptake of 131I. Acta. Endocrinol., 34:586-596.

- . 19606. Thyroidal activity in hibernating Chiroptera. II. Synthesis of radio-

iodinated amino acids. Acta Endocrinol., 34:597-604.

Schmidt-Nielsen, B., and R. O’Dell. 1961. Structure and concentrating mechanism of
the mammalian kidney. Amer. J. Physiol., Zool., 200:1 1 19-1 124.

Sturkie, P. D. 1965. Avian physiology. Cornell University Press, Ithaca, New York,
2nd ed., xxvii + 766 pp.

Thomas, S. P. 1975. Metabolism during flight in two species of bats, Phyllostomus
liastatus and Pteropus gouldii. J. Exp. Biol., 63:273-293.

Thomas, S. P., and R. A. Suthers. 1972. The physiology and energetics of bat flight. J.
Exper. Biol., 57:317-335.

Valdivieso, D., and J. R. Tamsitt. 1971. Hematological data from tropical American
bats. Canadian J. Zool., 49:31-36.

Valdivieso, D., J. R. Tamsitt, and E. Conde-del Pino. 1969. Electrophoretic properties
of neotropical bat hemoglobin. Comp. Biochem. Physiol., 30:1 17-122.

Wimsatt, W. A., and A. Guerriere. 1962. Observations on the feeding capacities and
excretory functions of captive vampire bats. J. Mamm., 43:17-27.

POPULATION AND COMMUNITY ECOLOGY

Stephen R. Humphrey and Frank J. Bonaccorso

Bats are the numerically dominant group of mammals in the Neotropics.
They comprise 52 per cent of the mammalian species in Costa Rica (Robinson,
1971) and 46 per cent of those in Panama (Handley, 1966). The family
Phyllostomatidae accounts for 55 per cent of all Costa Rican bat species and
59 per cent of the species in Panama. In terms of number of individuals, the
density of some phyllostomatid species far exceeds that of any other kind of
mammal in Central America (F. J. Bonaccorso and D. Morrison, unpublished).
Additionally, phyllostomatids exhibit great diversity in the types of food used,
with specializations for eating fruit, nectar and pollen, insects, small land
vertebrates, and blood of birds and mammals. The importance of this family
in diversity and relative density suggests an equivalent functional importance
in tropical ecosystems.

A recurrent theme in tropical ecology and in this volume is the seasonal
variation of tropical climate. The dominant feature of tropical climate is an
annual cycle of wet and dry seasons (see Rumney, 1968). It is not uncommon to
find tropical dry or wet forests (forest types refer to the classification of Holdridge,
1967) that receive 200 to 400 millimeters of rain per month in the wet season
and no measurable rain in some dry season months. Tropical wet and rain
forests have less distinct dry seasons but predictably have reduced rainfall in
certain months. The influence of wet-dry seasonality on the foraging and
reproduction of tropical bats was discussed by Baker and Baker (1936),
Mutere (1968, 1970), Liat (1970), Mares and Wilson (1971), Fleming et al.
(1972), and Heithaus et al. (1975).

Foraging Strategy

Optimal foraging strategy requires that animals maximize food intake
(benefits) while minimizing expenditure of time and energy (costs) of acquiring
food. The distribution of food resources in time and space, the type of food
eaten, and competition for food all weigh heavily in shaping foraging strategy
(Schoener, 1969). Additionally, transitional stages in the evolution of species
or individual life histories may coincide with less than optimal time-energy
budgets when animals use excessive energy to exploit new resources. For
example, some phyllostomatids that change their diets seasonally may incur
such increased foraging costs.

In this section, we discuss factors influencing the foraging strategies of
phyllostomatids. We suggest that Neotropical bats feeding on vertebrates and
blood can rely on stable and abundant food resources throughout the year.
On the other hand, fruits, flowers, and insects are extremely seasonal in abundance.

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Some phyllostomatids specializing on these food types may encounter local
shortages at predictable times of the year. In order to survive such food shortages,
foraging strategies of tropical bats include migration, dietary changes, dis¬
continuation of reproduction, and successfully competing with other species
for limited food resources. In addition, phyllostomatids do have seasonal
fat cycles (McNab, 1976) and undergo at least diel torpor (McNab, 1969);
these strategies also might help in the accommodation of food shortages.

Fruit

Fruit availability in tropical forests varies in complex ways. Some tree
species produce fruit synchronously each year at a characteristic season. Some
fruit rhythmically but not every year. Others fruit with no discernable pattern
from once every few years to several times a year (Richards, 1973; Foster,
1973; Frankie et al., 1975).

Thorough studies of fruiting patterns in tropical dry, moist, and wet forest
plant communities have been conducted in Panama by Foster (1973) and in
Costa Rica by Frankie et al. (1975). These studies show that edible fruit is
available throughout the year, regardless of life zone, but that sharp seasonal
fluctuations occur in the number of species fruiting and the total fruit biomass.
In dry forest, a single peak in the number of species with mature fruits occurs
during the wet season. Both moist and wet forests have two peaks in the
number of species fruiting, one each in the dry and wet seasons.

Heithaus et al. (1975) studied the foraging patterns and resource use of
six fruit-eating phyllostomatids near Canas, Costa Rica. The tropical lowland
dry forest of Canas has a wet season from May to early November and a dry
season from mid-November through April. Virtually no rain falls in the dry
season, and the forests are semideciduous, with about half the tree species losing
their leaves (Daubenmire, 1972). In each month, between five and 10 species
of plants produce fruit eaten by bats. A single strong peak in the number of
plant species with “bat fruits” occurs from May through August. During the
early dry season, when the fewest kinds of fruits are ripe, a peak in the number
of species of blooming “bat flowers” occurs. At that time, all “fruit bats” at
Canas switch in part to a pollen and nectar diet. Three species ( Carollia
perspicillata, Sturnira lilium, and Artibeus jamaicensis ) that eat fruit,
nectar, and pollen reproduce twice each year — once coinciding with the dry
season and once with the fruit abundance in the wet season. S. lilium undergoes
a marked change from nectarivory in the dry season to frugivory in the wet
season. Thus, female S. lilium on a diet that is either primarily nectar and
pollen or primarily fruit are able to nurse young.

Our own unpublished data from a moist forest site, Barro Colorado Island,
Panama, reveal that from six to 19 species of bat fruits are available each month.
Again, two peaks in fruit abundance occur, one in the wet season and one in
the dry season (Fig. 1), and Artibeus jamaicensis correspondingly reproduces
twice each year. At this site, few bat flowers are available, and A. jamaicensis

BIOLOGY OF THE PHYLLOSTOMATIDAE

411

m -q
C m
Co *3

i_ m
> Z

n m

z o

CO -n
CO _

Fig. 1. — Seasonal reproduction of female Artibeus jamaicensis and of trees supplying
this species with food, on Barro Colorado Island, Panama.

relies on a diet of fruit from canopy trees throughout the year. The period of an
adult female mammal’s year that is most expensive energetically — lactation
(Miguela, 1969; Studier et al., 1973) — is even more costly in the first
reproductive peak of A. jamaicensis , which occurs during the late dry season.
Then most bats are simultaneously lactating and pregnant with embryos to be
born in the wet season. Selective pressure for this postpartum estrus probably
arose from the combination of the four-month gestation of A. jamaicensis
and the occurrence of the second fruiting peak four months into the wet season.
This reproductive adaptation places the end of the second lactation period
during the year’s second fruiting peak. Therefore, coincidence of the first
lactation with the year’s larger and longer fruiting peak is a doubly vital
phase of seasonal timing. The wet season fruiting peak is followed by two months
of fruit scarcity; it is accompanied by another postpartum estrus of A. jamaicensis,
but development of the embryo is delayed until the end of the wet season (Fleming,
1971).

The fruiting patterns of individual plant species often are less important to
bats than are the fruiting patterns of inclusive genera. Usually, all members of
a genus will have similar fruits, either edible or not. For example, the 1 8 species
of Miconia eaten by birds in Trinidad’s Arima Valley fruit for periods of one
to four months, with fruiting intervals spaced through the year so that from one
to seven species always are bearing fruit simultaneously (Snow, 1965). On
Barro Colorado Island, Panama, and at Cahas, Costa Rica, several species

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of the shrub genus Piper (Fig. 2) are sympatric. Each fruits cyclicly, with
species cycles offset so that fruit of the genus is available all year (Heithaus
et al ., 1975; our data). Pipers are the most important food species for bats of
the genus Carollia in Central America (Howell and Burch, 1974; Heithaus
et al., 1975; our data). The plant genus Ficus (figs) has many species that
consecutively serve as dietary staples for Artibeus and other stenodermines.
The same is true for Cecropia trees and Phyllostomus discolor, in our experience.
The year-round availability typical of such dietary staples may result from
long coevolution in response to mutualistic seed dispersal (Snow, 1965).

Some important bat fruits are available only for several months and do not
have congeners fruiting at other times of the year. For example, Spondias
mombin is ripe only from September to December on Barro Colorado Island
(Smythe, 1970). (Croat, 1974, reported that this species begins to fruit in
July, but this is true only along watercourses and drainage ditches.) Its only
congener, 5. radlkoferi, also fruits within this period. Both species of Spondias
are important food items for bats when few other fruits are available during the
heaviest rains of the wet season.

The low densities of tree species in heterogeneous Neotropical forests may
force large foraging distances upon herbivorous bats. Large-sized specialists on
fruit should have greater foraging distances than smaller generalists. For
example, Fleming et al. (1972) gave mean recapture distances of 347 meters
for Artibeus jamaicensis and 167 meters for Carollia perspicillata. Heithaus
et al. (1975) reported that small species feed on resources of high abundance,
whereas large species use resources that are more patchy in time and space.

Nectar and Pollen

Community patterns in the timing of flowering for Neotropical plants,
like fruiting patterns, are quite complex and vary by life zone. In dry forests,
most species flower during the dry season (Allen, 1956, summarized by Janzen,
1967; Fournier and Salas, 1966; Daubenmire, 1972; Frankie et al., 1975).
In the dry forest in Costa Rica (Heithaus et al., 1975), the number of species
of bat flowers in bloom varied from a low of two in July and August (wet season)
to a maximum of seven in January and March (dry season). During the dry
season, while bat flowers were abundant, seven species of phyllostomatids
were regularly covered with pollen from flower visits. At this time, the flowering
periods of plants were displaced — adaptations effectively avoiding competition
for the services of bat pollinators. However, as the wet season began and flowers
decreased, only Glossophaga soricina continued to visit flowers regularly for
food. Phyllostomus discolor apparently responded to the scarcity of flowers

Fig. 2. — Flowers and fruits used as food by phyllostomatids: A, Cecropia eximia
(Moraceae) fruit; B, Piper aeqaale (Piperaeae) fruit; C, Astrocaryum standleyaniun
(Palmaceae) fruit; D, Ochroma lagopus (Bombacaceae) flowers closed during daytime;
E, Pseudobombax septenatum (Bombacaceae) flower; F, Markea sp. (Solanaceae) flower.

BIOLOGY OF THE PHYLLOSTOMATIDAE

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by migrating. The other five species switched to fruit diets. This shows how
different species react with different strategies to a scarcity in food resources
that are shared during times of abundance.

In the moist forest on Barro Colorado Island, only five kinds of flowers appear
to be fed on by bats (see Figs. 2d and 2e\ our data). These flowers are available only
from late December to late March, during the dry season. Thus, for nine months
each year, little nectar or pollen is available to bats. Here, nectar and pollen are
important food sources only for G. soricina and P. discolor. The latter does not
migrate as it does in the dry forest studied by Heithaus et al. (1975); instead, it
switches to a diet of fruit and insects. G. soricina , which typically is more dependent
on floral resources, is rare on the island but apparently also switches to fruit.

We know of no dry-season flower feeders that switch wholly to insects in
the wet season. Instead, they switch to fruit or fruit and insects together (for
example, G. soricina , Fleming et al., 1972, and P. discolor at our Panamanian
site). It might be most realistic to view such species as herbivores with omnivorous
tendencies, in which case it is proper to wonder if a plant-adapted gastrointestinal
tract (Rouk and Glass, 1970) could function effectively on a wholly insectivorous
diet.

Taxa, such as Leptonycteris and Choeronycteris, that are exceptional in not
switching from plant food, migrate to stay permanently in “dry season” environ¬
ments by moving to subtropical and warm temperate thorny vegetation zones
where suitable flowers occur in summer. In view of the many potential competitors
among insectivorous and frugivorous bats, the selective pressure for migration
is true nectar-pollen specialists should not be underrated.

Insects

Wet-dry seasonality strongly affects the distribution and abundance of
Neotropical insects. The dry season presents many insects with food shortages
and water balance problems. Most tropical insects survive the dry season as
adults (Janzen and Schoener, 1968) rather than in diapause (as in winter
survival of temperate taxa). However, the precise impact of tropical seasons
on the food of insectivorous bats is difficult to assess, because few studies deal
with the particular insects of interest. These are nocturnal species either in
flight, for bats that catch flying prey, or active on leaves, tree trunks, and the
ground, for bats that feed by gleaning.

In a study of mosquito seasonality based on adults flying into a livestock-baited
trap, Bates (1945) showed that nocturnal species peak in abundance immediately
after the onset of the wet season. Some species exhibited a secondary peak
near the end of the wet season, and all species were least common in the dry
season. In addition to this annual periodicity, one species underwent population
irruptions, with a hundredfold difference in minimum and maximum numbers
over a two-year period.

Light-trap samples in moist forest in Panama (Smythe, 1974) document
remarkable seasonal changes, with up to eight times as much insect biomass

BIOLOGY OF THE PHYLLOSTOMATIDAE

415

Fig. 3. — Seasonality of tropical insect biomass (after Smythe, 1974). This pattern
occurs in Central America where distinct dry seasons occur. The timing varies
geographically by one to three months.

in the wet season as in the dry season. Large taxa ( > 5 millimeters long) were
responsible for this change, with Isoptera, Diptera, and Lepidoptera having
particularly dramatic population increases early in the wet season. By contrast,
small taxa (<5 millimeters long) were of constant abundance throughout the
year. Combined data (Fig. 3) show biomass increasing shortly after the wet
season begins, peaking about a month later when reproduction, growth, and
metamorphosis is complete, and remaining high for the next three or four
months. Biomass declines late in the wet season at the time when the heaviest
rains occur, and it remains low through the dry season.

Insectivorous phyllostomatids may exhibit at least three responses to the
seasonality of their food. One would be to bear young at the beginning of the
wet season; the limited data available (see Wilson, this volume) suggest that this
often may be the case. Another would be to switch to other types of food.
A partial shift occurs in Micronycteris hirsuta, which gleans insects as its primary
food but supplements this diet with fruit during the dry season (Wilson, 1971).
A third response would be to change foraging habitat. In the dry forest near
Canas, Costa Rica, Janzen (1973) noted that night-time numbers of beetle
and true bug species decreased much less during the dry season in riparian
forest than in nearby pasture land and upland deciduous forest. Thus, riparian
forest may serve as a dry season refuge for food of insectivorous bats, assuming
that the preferred insect taxa behave similarly.

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Few precise data exist on the food habits of insect-eating phyllostomatids.
Wilson reported that large roaches, Orthoptera, and scarabaeid beetles are
the most important items in the diet of Micronycteris hirsuta in Panama. These
insects spend much of their time walking and feeding on vegetation or detritus.
Wilson concluded that M. hirsuta captures most of its prey by gleaning. This
pattern appears to apply to other members of the genus as well (Gardner, 1977).
Macrotus californicus in the southwestern United States also feeds heavily on
large insects, including larval Lepidoptera that probably are gleaned from
foliage (Ross, 1967). A gleaning mode of foraging was confirmed for Macrotus
waterhousii by watching one (S. R. Humphrey, aided by an ultrasonic sensor
and a streetlight) for which the feeding flight was confined to the interior of
an almost spherical tree crown. F. J. Bonaccarso observed a captive pair of
Tonatia bidens take large cicadas, katydids, grasshoppers, and beetles by
picking them off the walls of their cage.

Phyllostomatids known to be mainly insectivorous have adaptations
characteristic of bats that glean prey from vegetation or the ground. Such
adaptations include large eyes, large ears, a long robust rostrum, long vibrissae,
and a low wing aspect ratio that promotes vertical flight and hovering. Many
insectivorous phyllostomatids with these features also have long nose leafs.
As suggested by Wilson (1971), many of the same food-gathering skills
probably are involved in securing fruit and resting insects. By contrast, none
of the Neotropical insectivores of other families (Emballonuridae, Mormoopidae,
Furipteridae, Thyropteridae, Vespertilionidae, and Molossidae, possibly
excepting Natalidae) appear to have this gleaning morphology, although
several Nearctic vespertilionids do ( Antrozous , Euderma, Plecotus, Idionycteris,
and several species of Myotis). We suspect an evolutionary character displacement
at the family level, in which phyllostomatids were decisively preeminent as
insect gleaners. A gleaning morphology well could have provided suitable
preadaptations for specializing on vertebrate prey, as in Chrotopterus auritus
and Vampyrum spectrum.

Vertebrates

The literature on foods of vertebrate-eating phyllostomatids is not detailed,
but at least lizards, birds, mice, and bats are taken (Goodwin and Greenhall,
1961; Gardner, 1977). We offer information on seasonal abundance of birds
by way of example. Peaks of bird breeding should coincide with high population
levels. Tropical birds may breed continuously, regularly in concert with wet-dry
seasons, or irregularly. In continuous breeders, individuals or pairs breed
according to their own activity cycles, with all reproductive stages present in
the population at any time. Most regular breeders in regions with a weak dry
season breed in the drier months. In regions with a pronounced dry season,
most breed in the wet season, but few specialists breed in the dry season. Regions
with two annual wet-dry cycles have some species breeding once a year and
others breeding twice (Immelmann, 1971). Superimposed on this complex

BIOLOGY OF THE PHYLLOSTOMATIDAE

417

pattern is the arrival of numerous migrants during austral and boreal winters.
Thus, at least in a general sense, it appears that night-roosting birds should
be in ample supply at all times of the Neotropical year.

Blood

Aside from man and domestic animals, food for blood-eating phyllostomatids
should be a seasonally stable resource, as the birds and mammals parasitized
are large and have long life spans. For the same reason, these hosts are likely
to occur at low densities and therefore to be difficult for vampires to locate.
Conversely, humans and domesticants are high-density hosts that are predictably
accessible in time and space. One account (Benzoni, in Turner, 1975) suggests
that humans were a major host for vampires along the east coast of Costa Rica
in the sixteenth century. Now, greater use of houses that limit access makes
human bites unusual, but vampires commonly and regularly feed on domestic
birds and mammals. In accord with these observations, our impression from
mist-netting is that sanguivorous bats are rare except where domestic animals
are abundant; see Fig. 6 for some illustrative data.

Roosting Strategy

Like other bats, phyllostomatids spend the daylight hours at rest. A good
roost should provide some protection from adverse weather, predators, and
nonresident parasites such as diurnal mosquitoes and biting flies. Beyond this,
a roost should afford microclimatic conditions that are not stressful and that
favor effective use of available energy. At the very least (though hardly a
problem), a roost should prevent prolonged exposure to direct sunlight,
because phyllostomatids die at body temperatures of 37 to 42 °C (McManus,
1977). More importantly, microclimate should be optimal for growth during
periods of gestation and lactation.

Roost Type

Phyllostomatids use an amazing variety of natural and man-made structures.
These include caves, culverts, buildings, bridges, cisterns, steam banks, cliff
crevices, tree foliage, tree hollows (even hollow tree trunks lying on the forest
floor), rabbit burrows, and old termite nests. The tent-making bat ( Uroderma
bilobatum ) makes shelters by clipping palm fronds so that the frond tips fold down.
Most species seem unrestricted to particular sorts of roosts. For example, Des-
modus rotundus occurs in tree hollows, caves, and culverts. Commonly several
species will share a roost, in bodily contact with each other (Goodwin and Green-
hall, 1961).

Roost Microclimate

The microclimates of phyllostomatid roosts are known from a single study.
McNab (1969) recorded air temperature and relative humidity in roosts of

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12 species at the time of capture. Temperature ranged from 13 to 29 °C and
humidity from 70 to 98 per cent, so microclimate is characteristically mild and
moist. Studies of diel, seasonal, and regional variation of these microclimates
have not been reported.

A few studies characterize tropical forest microclimate and indicate conditions
that might be encountered by a foliage-roosting bat. Dry season data at Barro
Colorado Island, Panama, show a weekly temperature range of 25.8 to 27.4 °C
at the ground, 24.9 to 33.0° in the subcanopy, and 27.0 to 37.5° in the canopy
(Allee, 1926). Relative humidity and light intensity were likewise stratified,
and conditions were more extreme in sunflecks than in the shade. Similar daytime
temperature profiles occur in other tropical forests (Hales, 1949; Baynton
et al., 1965). Allen et al. (1972) showed that such stratification is stable all
day, breaking down in the evening, and that it is caused by the ameliorating
effect of vegetation on air turbulence rather than any constancy of incident
conditions. Thus, near-lethal temperatures occur in the canopy, but a bat can
easily avoid them by seeking shaded sites in lower foliage.

Studies of phyllostomatid response to cooling, such as would be encountered
at high latitudes or altitudes, are inconsistent, apparently because of differing
experimental procedures. When bats were exposed to rapidly dropping
temperatures for two hours (McNab, 1969), four hours (Carpenter and Graham,
1967), or exposed to cold for several days with food provided ad libitum
(Arata and Jones, 1967; Arata, 1972), they responded endothermically,
surviving by increasing metabolic rate. Exceptions were small stenodermines
and the three vampire genera, which died quickly as temperature dropped.
Animals with food available fed many times a day when cold. Studier and
Wilson (1970) used fed animals but did not provide food during their experiments,
lowering temperatures stepwise from 34 to 2.5 °C over periods of seven to 10
hours, and allowing body temperatures to stabilize at each step. Most individuals
were wholly ectothermic or else partially so, maintaining body temperatures
5 to 15°C above ambient temperatures while both ambient and body temperatures
decreased. Below 8°C most bats went into torpor and died after failing to arouse.
One lactating female Carollia perspicillata remained endothermic at ambient
temperatures as low as 5.7 °C.

Obviously a bat in a roost with a temperature that is too low can leave for
an alternate site, but if it remains it cannot feed and would be exposed to roost
temperatures for approximately eight to 12 hours. Realistic thermoregulation
studies should employ microtemperatures that are stable or that increase
during the day. The limited data on thermal response lead us to hypothesize
that at low roost temperatures (1) reproducing female phyllostomatids
thermoregulate, incurring the consequent metabolic costs, and (2) nonreproducing
females thermoregulate weakly or not at all. In the latter case, presumably the
practice would not be fatal at roost microclimates encountered at low altitude
in the Neotropics. At higher latitudes or altitudes, ectothermy could be fatal
and perhaps phyllostomatids in such circumstances attempt to thermoregulate.

BIOLOGY OF THE PHYLLOSTOMATIDAE

419

Roosts as a Limiting Factor

Most phyllostomatids roost alone or in small colonies and are not strongly
specialized to be highly colonial in order to exploit particular roost types (Dwyer,
1971), as is common in families characteristic of temperate zones (Humphrey,
1975). Known highly gregarious exceptions are Phyllonycteris, Erophylla,
Desmodus, Brachyphylla, and Phyllostomus (Dalquest and Walton, 1970).
Satisfactory roosts are available in abundance in the Neotropics. For these
reasons, and because of the probable importance of food as a limiting factor
(McNab, 1971), it would be expected that roosts seldom limit phyllostomatid
abundance and community structure.

Distributional limits of herbivorous and nonmigratory carnivorous
phyllostomatids should be determined by food. However, roosts may be limiting
factors at the distributional limits of many carnivorous phyllostomatids,
including sanguivores and migratory vertebrate-eaters and insectivores. Dwyer
(1971) predicted that such bats — that is, tropical species adapted to tropical
roost microclimates — will be limited at higher latitudes and altitudes by
absence of suitable food and by the increased cost of thermoregulation in caves
with cool microclimates. Including bats of all feeding types, Dwyer judged
that food would be a more critical factor than roosts. In fact, McNab (1973)
calculated that the cost of thermoregulation in cool roosts prevents Desmodus
rotundus (which, as in any animal, can consume only so much food nightly)
from occupying higher latitudes, even though its preferred food is abundant.
That no insectivorous phyllostomatids are known to migrate to temperate
zones may reflect both their thermoregulatory disadvantage in cool roosts and
a probable competitive advantage on the part of vespertilionid insect-gleaners
that hibernate in winter. The migratory nectar-pollen feeders, Leptonycteris
and Choeronycteris, move only as far north as the hottest and driest areas of
Texas and Arizona, although one preferred food (Agave) occurs much farther
northward. That these bats appear to be highly colonial at the northern limits
of their range (Easterla, 1972) may reflect clustering thermoregulatory
behavior.

Demography

Every animal can be said to have a demographic strategy — a combination
of performances that adds individuals to the population in concert with factors
that subtract individuals, with a pattern of magnitude, balance, and timing that
differs for each species. A demographic strategy is a set of responses to an
environment; to some degree a species may vary its strategy among environments
(for example, in response to different seasonal regimens of climate and food in
tropical lowland dry as opposed to wet forest). On the other hand, the evolved
nature of some demographic phenomena (for example, biotic potential, the
dispersal effect of pioneering, and the ability to avoid predation) results in
reasonably fixed numerical expressions. The totality of demographic events
produces a growth rate that must be positive or zero over any substantial time

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interval if a population is to survive. Demographic lability enables a species
to survive in a variety of environments, but demographic limitations make
success in other environments unlikely. Values of demographic parameters
show specific things an animal does to succeed and simultaneously reveal
performances that must be modified to enable use of another location.

Ways that demographic factors interrelate and operate are presented in a
flow diagram (Fig. 4). If population size and the natural rate of increase (r)
are known, growth trends can be predicted and used to evaluate the progress
of the population. Three basic demographic parameters integrate to determine
population growth rate and size: natality rate, survival rate, and dispersal rate.
Factors affecting the values vary quantitatively as a function of population age
structure because the importance of each intrinsic factor (for example, emigration)
changes with age. The extrinsic factors and potential regulatory pathways are
speculative, as these seldom or never have been demonstrated to operate
among phyllostomatids or other bats. However, studies directed toward these
factors and pathways should reveal the implications of demographic adaptations
of phyllostomatids. Although no demographic strategy is documented thoroughly
for these bats, piecemeal data on intrinsic factors ( sensu Fig. 4) are reported
in recent literature.

Number of Births per Year

Most phyllostomatids studied to date are polyestrous (Fleming et al.,
1972; Wilson, this volume) with the maximum possible number of estrous
cycles being two (possibly three may occur in tropical vespertilionids, as shown
by Wilson and Findley, 1971). Two peaks in parturition clearly are indicated
by the bimodal pattern of pregnancy of most species, and a maximum number of
two is dictated by the long gestation period of Desmodus rotundus. Of much
more interest than the maximum number of births possible annually however,
is the average number actually occurring. To our knowledge such data are
unavailable. In cases of bimodal polyestry, individual females could produce
offspring at none, one, or both peaks during a year. Frequent observation of
females simultaneously lactating and pregnant shows the latter case to be
common. Macrotus californicus gives birth only once per year (Bradshaw,
1962). Members of the genera Leptonycteris and Choeronycteris that annually
migrate from tropical to warm temperate regions are parturient during the
temperate zone summer; available data do not preclude the possibility of
a second birth during the tropical dry season. Data on Leptonycteris sanborni
(Cockrum and Ordway, 1959; Howell, 1972) suggest two peaks in parturition
for each female or a single peak that occurs either in the temperate summer or
the tropical dry season.

Number of Offspring

All phyllostomatids presently are thought to have a single young at a time.
Carter (1970) termed the family “characteristically monotocous,” and

BIOLOGY OF THE PHYLLOSTOMATIDAE

421

Fig. 4. — Operational pathways of demographic factors.

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Fleming et al. (1972) stated that in seasonally polyestrous phyllostomatids
“one young is produced in each pregnancy.” Records of twinning prove to be
the exception, not the rule. Barlow and Tamsitt (1968) reported three sets of
unborn twins in 195 pregnancies in A. jamaicensis, one of 615 in Glossophaga
soricina, and one of 10 in Erophylla bombifrons. Burns (1970) noted one case in
Desmodus rotundus. In Artibeus lituratus, the single young results from
reducing the number of ova shed; in a sample of 49 females, the average number
of corpora lutea or mature follicles was 1.37, but only one ovum was released
at estrus (Tamsitt and Valdivieso, 1965).

Production of a single young in phyllostomatids is consistent with the
pattern generally expected (Spencer and Steinhoff, 1968; MacArthur, 1972)
in that animals in tropical latitudes have more numerous but smaller litters than
those in temperate latitudes. By contrast, Nearctic vespertilionids and molossids
are monestrous, with mean number of young ranging from 1 to 3.4 (Humphrey,
1975). The possibility of larger litters in phyllostomatids should not be ignored
entirely, however, as some tropical bats of other families regularly produce
twins. Examples are the tropical vespertilionids Rhogeesa parvula (Cockrum,
1955) and R. tumida (Goodwin and Greenhall, 1961).

Birth Rate

One important intrinsic factor can be measured by answering the question,
“for each species, what percentage of females gives birth during each birth
pulse?” The pregnancy rate may approximate the birth rate in species with
synchronous parturition if late abortions and stillbirths are few, as is true of
some temperate-zone vespertilionids. Single, fortuitous captures of pregnant
phyllostomatids (Mares and Wilson, 1971; Fleming et al., 1972) demonstrate
asynchrony or partial synchrony of breeding. Because births are not simultaneous
within seasonal birthpulses, these pregnancy data do not indicate which
individuals are breeding and which are not. However, in our experience,
properly timed samples accounting for pregnancies and early lactations can
generate estimates of the proportion reproducing. To our knowledge, no
phyllostomatid birth rate data have been published (the values for Desmodus
rotundus in Turner, 1975, do not permit calculation of annual or seasonal
rates).

Age at Sexual Maturity

In Macrotus californicus, females breed during their first autumn and give
birth at the age of one year (Bradshaw, 1962). Males do not breed until their
second autumn, which is not disadvantageous so long as enough males live
that long, and it may be an advantage in ensuring that only successful male
genotypes are perpetuated.

To our knowledge, this important factor has not been documented for any
other phyllostomatid, notwithstanding the unsubstantiated suggestion that
females of Artibeus jamaicensis become pregnant in the dry season following

BIOLOGY OF THE PHYLLOSTOMATIDAE

423

their birth (Fleming et al., 1972). Data on age at sexual maturity come only
from recapture of marked females of known age taken from one to several
breeding seasons after their birth. For example, one Desmodus rotundus
marked as an infant was pregnant when recaptured 18 months later (Turner,
1975). Careful study may reveal that this parameter varies in response to
unusual hardship dictated by climatic or habitat variation (see Christian, 1971).

Survival Rate

The only record of phyllostomatid longevity exceeding two or three years
is a recapture of an A. jamaicensis seven years after banding (Wilson and
Tyson, 1970). Longevity information is useful to indicate the maximum age
attainable by a species but is of little importance as a numerical expression of
demographic strategy. The parameter of interest is the mean life expectancy
(mean life span), the value designated e0 in a life table. Mean life expectancy
reflects the actual performance of a cohort of animals in nature, and it
integrates properly with the other rate values. For example, it allows easy
calculation of the number of offspring produced during the average lifetime of
a female.

The only proven technique for documenting bat survival is frequent re¬
capture of marked individuals of known age and sex. Failure to determine
age at the time of marking (for example, banding cohorts of bats during
temperate winters) has resulted in voluminous but not especially useful bat
survival data in the literature. Such data by definition produce a constant
rate of survival throughout life (or nearly so, depending on details of sampling
protocol), when actually mammals characteristically have lower survival in
immature and elderly stages than during adulthood (Caughley, 1966). The
most satisfactory period to mark cohorts of immature bats is just prior to
weaning. Obtaining accurate data then depends on recapturing all of the living
cohort members at least once annually until the last individual has died.
Mortality between birth and weaning should be documented to prevent
overestimation of survival. Because marking extremely young bats would
cause many deaths, the best available technique is to determine the number of
young born in a roost, remove all carcasses from the roost area, and count the
number of young dying before they begin to fly. Examples of such data are
preweaning survival of a molossid (Herreid, 1967), Tadarida brasiliensis,
and postweaning survival of a vespertilionid (Humphrey and Cope, 1976),
Myotis lucifugus. These studies must be done at roosts and require many years
for long-lived species.

Aging bats according to tooth cementum annuli is an exciting prospect,
as it would provide an “instantaneous” method of constructing survival curves.
A disadvantage is that the animals must be killed to acquire the data. This
technique has been applied to Desmodus rotundus, yielding mean age values
of 3.0 years for females and 1.5 years for males in Mexico (Linhart, 1973)
and 4.13 years for females and 3.01 years for males in Argentina (Lord et al.,

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1976). The latter author suggested that apparent differences in sex-specific
survival are artifacts of the social structure of the sampled populations.
Unfortunately, in neither study were annuli counts checked against known-age
control animals to confirm that a line represents one year’s growth. Why growth
should be periodic when food supply is constant is unclear.

Dispersal Rate

Pioneering is a vital phenomenon for finding available habitats and com¬
pensating for local extinctions. For demographic purposes, dispersal rate is the
net loss or gain of animals by one-way movement in proportion to the population
in a given area. No measurements of phyllostomatid dispersal rates occur in
the literature. Studies of dispersal rate of mobile animals must include large
geographical areas, and mammalian dispersal is seldom quantified. For examples
and discussion of procedural difficulties, see Barkalow et al. (1970) and
Humphrey and Cope (1976). Site attachment index values and associated
movement data on two species of temperate vespertilionids (Humphrey and
Cope, 1970, 1976; Humphrey, 1975) indicate little or no dispersal of recruited
females in undisturbed populations; whether such a pattern applies to phyl-
lostomatids in the tropics in unknown.

Migration has no effect on the dispersal rate if a migrating individual indeed
returns. If the migrator stays away, then it becomes a dispersor, and if it dies
while migrating the effect is on the survival rate. These distinctions help prevent
confusion about the demographic implications of migration. Migration has
not been demonstrated clearly for any phyllostomatid, but many sorts of
collateral evidence suggest that Leptonycteris and Choeronycteris are migratory
in the northern part of their range (Hayward and Cockrum, 1971). Further,
our unpublished data suggest that some species in Belize and Panama are
migratory or at least nomadic (see beyond).

Community Diversity

Field biologists recognize great differences in the various bat communities
that they sample. Although patterns of diversity occur in and among these
taxonomic communities, so many characteristics of species and habitat factors
are involved that these patterns are difficult to perceive and express.

Species Number

The simplest measure of diversity is the number of species present (in the
literature termed variously faunal size, species density, species diversity, and
species richness). Often this is the only useful measure of diversity available
from specimens taken for taxonomic purposes. Bat communities (and numbers
of phyllostomatid species) in the Americas are largest in tropical lowland rain
forest. Moving away from that life zone in moisture, altitude, or latitude, the
number of species diminishes (Fig. 5). Beyond this common observation,
analysis of species number reveals little about the nature of bat communities.

BIOLOGY OF THE PHYLLOSTOMATIDAE

425

COOL TEMPERATE

Desert

Desert Scrub

Steppe

Moist Forest

Wet Forest

Rain Forest

0.89(6,2)

0.05(71)

1.32(75)

99/0

83/0

100/0

Desert

2.2808,1)

99/0

Rain Forest

Desert

Rain Forest

Desert

2.04(12,2)

93/6

1.8502,3)

86/0

Rain Forest

Desert

Rain Forest

WARM TEMPERATE

Desert

Deserf Scrub
1.33(?1)
76/0

Thorn Steppe

165(10,1)

91/0

Dry Forest

Moist Forest
128(8,2)
95/0

Wet

Forest

Rain Forest

SUBTROPICAL

Desert

Desert Scrub

Thorn

Woodland

Dry Forest

Moist Forest

Wet

Forest

Rain Forest

SUBALPINE

MONTANE

LOWLAND

TROPICAL

Desert

—

0 80(3,1)
0/100

2 26(16,1)
2/84

2 60(1^1)
5/89

Rain Forest

LOWER

MONTANE

PRE-

MONTANE

LOWLAND

Desert

Desert Scrub

Thorn

Woodland

Very Dry
Forest

Dry Forest

169(18,5)

1/95

Moist Forest

2.13(18,8)

6/85

Wet Forest

214(20,6)

5/94

Rain Forest

2.36(29,1)

1/99

Fig. 5. — Characteristics of bat community structure according to life zone (after
Holdridge, 1967). Numbers are : top line, average diversity value, (average number of
species, number of samples); bottom line, proportion of the diversity contributed by
vespertilionids/phyllostomatids. Because no data are available from boreal or subpolar
latitudes or alpine altitudes, corresponding life zones are omitted. All samples were
mist-netted in tropical wet seasons, temperate summers, or year-round. Although the
best available, these samples are not ideal for diversity analysis. Samples vary in habitat
(for example, mature forest, riparian forest, slash-and-burn agriculture) and adequacy of
netting vertical strata and full nights. All tropical samples inadequately represent high-flying
molossids and taxa more difficult to net than phyllostomatids (for example, emballonurids,
mormoopids, and vespertilionids). Summaries and references to sample data are available from
the senior author on request.

Species Diversity

More can be learned by finding a concise way to compare communities
and the abundance of species within and among communities. Such comparison
is afforded by a species diversity index. Details of rationale and application
of this analytical tool to bat communities are presented by Humphrey (1975).
Briefly, the standard index is that of Shannon and Weaver (1949), H' =
— 2 PilogePi , where p{ is the number of individuals in the i,h species divided by
sample size. The contribution of species n to its community’s diversity is
H'n= ~ Pnl°SePn-

Parallel to the pattern of species number, species diversity (Fig. 5) is
highest in tropical lowland rain forest and decreases along gradients of moisture,
altitude, and latitude. The most diverse single sample (//' = 2.65) was taken in
garden and forest habitats at San Pablo, Peru (Tuttle, 1970); no doubt this is
an overestimate, as data from two habitats are pooled. Average diversity of

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warm temperate montane dry forest also is overestimated, because all three
sites were chosen for exceptional diversity in topography and vegetation.

Although fairly diverse communities continue into middle latitudes and
zones of intermediate moisture, a pronounced shift in the importance of
phyllostomatids occurs between subtropical and warm temperate zones. In
warm temperate zones, phyllostomatids are replaced by vespertilionids.
Presence of two species of nectar-feeding phyllostomatids in warm temperate
montane thorrujsteppe ( Choeronycteris mexicana and Leptonycteris sanborni
in Arizona samples) results from migration to take advantage of seasonally
available Agave and cactus flowers. Bat communities are least diverse in zones
of extreme dryness or high altitude or latitude. These correlations with the
Holdridgean axes of precipitation, humidity, and temperature suggest that
phyllostomatids, as a family, are best adapted to regions where 1) annual
precipitation exceeds 1000 mm, or 2) the ratio of potential evapotranspiration
to precipitation is less than two, and 3) a mean annual biotemperature about
17°C is available for at least one season of the year (as by migration). The
ultimate factors responsible for this pattern will become clear as the functions
of the morphological, behavioral, demographic, and physiological adaptations
of these bats are better understood. We infer that the pattern represents
phyllostomatid response either directly to climate or to biological factors such
as vegetation or food.

The general lack of anomalies in diversity trends of phyllostomatid-
dominated faunas is striking. One exception is in tropical lower montane dry
forest, where small sample size (10) of the single sample may account for
low diversity. By contrast, no clear life-zone pattern appears in diversity of
warm and cool temperate bat faunas. As shown by Humphrey (1975), the
presence of suitable roosts enables strongly roost-adapted vespertilionids and
molossids to become exceptionally abundant there. A super-abundant species
affects the diversity value because H'n 1< H'n 2, lowering H' . Thus for roost-adapted
taxa, perhaps including the tropical mormoopids, we expect such factors as
karst topography and forest management practices to be of primary importance.

Some indication of the importance of certain species to their bat communities
is given in Figs. 6 and 7. Consistently important species in lowland forest are
the feeding generalists Carollia perspicillata and small species of Artibeus,
which eat a wide variety of fruits. When fruit is scarce, C. perspicillata also
will consume nectar, pollen, and insects. Other generalists such as species of
the genus Sturnira, however, are consistently minor community members.
Specialists on large fruit, Artibeus jamaicensis and species of Vampyrops, do
best in wet climates and decrease in importance in drier forests. High H' „
of A. jamaicensis in dry forest is an artifact in that all samples there were in
fruit plantations or riparian gallery forest that included many fig trees. High
importance of both A. jamaicensis and small species of Artibeus in forest of
intermediate moisture accords with our unpublished data that these bats eat
different species of fruit, partitioning food on the basis of particle size.
Glossophaga soricina, a species that specializes on nectar and pollen but

BIOLOGY OF THE PHYLLOSTOMATIDAE

427

TROPICAL LOWLAND FOREST

Fig. 6. — Contribution to bat community diversity by species of phyllostomatids along
a moisture gradient in tropical lowland life zones. Data are from samples used in Fig. 5.

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TROPICAL WET FOREST

Fig. 7. — Contribution to bat community diversity by species of phyllostomatids
along an altitudinal gradient in tropical wet forest life zones. In effect, this graph adds a
third dimension, elevation, to Fig. 6, with the origin at the point “tropical lowland wet
forest.”

switches to fruit in the wet season when flowers are scarce, is of complementary
importance to those bats that eat large fruits, increasing its contribution in
dry communities. Trachops cirrhosus, a specialist on insects and small vertebrates,
and Phyllostomus hastatus, a large bat that eats fruit, insects, and some
vertebrates, are predictably unimportant; we do not understand the higher
contribution of the latter in wet forest. Desmodus rotundus, a specialist on

BIOLOGY OF THE PHYLLOSTOMATIDAE

429

mammal blood, is a minor constituent of wet zones but becomes increasingly
important in drier forest, probably a function of increasing livestock density.

Altitudinal data (Fig. 7) reinforce the general conclusions derived from
lowland data. Carollia perspicillata is an important community member at
all elevations sampled. G. soricina, already seen to be a minor member in wet
lowland forest, is equally unimportant in higher forest. The large fruit
specialists A. jamaicensis and Vampyrops, which respond similarly to moisture
change in lowland forest, show opposite trends at high altitudes in wet forest —
prominence of Vampyrops and disappearance of A. jamaicensis. The frugivorous
generalists of the genus Sturnira and small species of Artibeus show a pattern
similar to that of Vampyrops , except that small Artibeus decrease in importance
at the highest elevation. Perhaps this pattern represents a competition-based
displacement or poor response by small kinds of Artibeus to the greater daily
variation of climate in high altitude forests. The prominence of Sturnira may
indicate some special adaptations to highlands in view of its unimportance
in all lowland forests sampled.

Careful work that samples vertical strata and accounts for differences of
habitat and season will reveal much more about tropical bat communities
and phyllostomatids. For example, ground-level nets at a site at Belem,
Brazil, yielded 18 species with a diversity of 1.93; simultaneous netting with
subcanopy and canopy nets placed above the ground nets added seven species
of phyllostomatids and increased diversity to 2.40 (Handley, 1967). In all-night
netting, LaVal (1970, personal communication) alternately sampled riparian
forest and banana groves at Finca La Pacifica, near Canas, Costa Rica. A
decrease in diversity in the plantation (1.63 as opposed to 1.84 in the riparian
forest) was accompanied by marked shifts in species composition; most striking
was the omnivorous Phyllostomus discolor, rare in the forest but by far the
most abundant species in banana groves, where presumably it ate banana nectar
and pollen. In an area of slash-and-bum agriculture near Frijoles, Canal Zone,
D. E. Wilson (personal communication) sampled during the wet season (shortly
after the main pulse of births for the year) and in the following dry season.
Wet season diversity was 1.93 with 14 species, but dry season values dropped
to 1.74 and seven, respectively; all species lost in the dry season were
phyllostomatids. Such data suggest exciting patterns of phyllostomatid
specialization in foraging strata and habitat and the possibility of seasonal
migration or nomadism among habitats.

Ecosystem Functions

The dynamics of energy flow are receiving increased attention by ecologists.
Producers and decomposers are recognized as the important organisms in
contributing to net productivity. Consumers account for little of the energy
flowing through their ecosystems (Fleharty and Choate, 1973; Fittkau and
Klinge, 1973). However, they do play key roles in directing ecosystem
dynamics. Long-term growth, succession, and stability of plant associations
depend partly on ecosystem functions performed by herbivores. Examples are

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seed dispersal by birds, primates, rodents, and bats; pollination by bees, moths,
and bats; and successional retardation by voles, prairie dogs, and some ungulates.

Phyllostomatid bats fall into several consumer trophic levels. These range
from the second level for fruit and nectar-pollen consumers, the third for
insectivores and sanguivores (most of the time), and the fourth or fifth for
carnivorous taxa such as Vampyrum spectrum and Trachops cirrhosus,
which often eat other carnivores. The interactions of the phyllostomatids with
other organisms and the ecosystem functions performed by phyllostomatids
will be discussed in the following sections.

Seed Dispersal

Phyllostomatid bats, birds, primates, and rodents are the most important
agents of seed dispersal in the Neotropical region (van der Pijl, 1972). Few
data exist for shrubs, vines, or epiphytes, but phyllostomatids act as dispersal
agents for up to 24 per cent of the forest tree species at some sites. At Finca la
Selva near Puerto Viejo in Costa Rica, Gary Hartshorn made the following
unpublished observations in a tropical wet forest (4000 millimeters of
rain per year; no dry season): in an area containing 20 species of fruit-eating
bats, 24 per cent of the 273 tree species counted bore bat-dispersed seeds.
F. J. Bonaccorso (unpublished data) recorded similar information for a tropical
moist forest (rainfall, 2750 millimeters per year; four to five month dry season)
on Barro Colorado Island, Panama: 7.7 per cent of the approximately 350
tree species identified carried seeds dispersed by bats; 16 species of fruit-eating
bats were present. In both of these works, a fruit-eating bat was defined as one
with 20 per cent or more of its diet consisting of fruit; trees, as being greater than
10 centimeters dbh or 5 meters tall. These observations suggest that
phyllostomatids become increasingly important as dispersal agents in wetter
forests, as evidenced by the percentage of trees dispersed by bats and increased
number of frugivorous bat species. Where long dry seasons occur, persistent
winds disperse the seeds of many trees. At sites without a strong dry season
and associated winds, animals (and water) play a major role in dispersing plant
species. Some dry forest sites may support higher than expected numbers of
frugivorous species because of the abundant quantities of bat flowers during the
dry season. At that time, these usually frugivorous bats switch to diets of nectar
and pollen (Fleming et al. , 1972; Heithaus et al., 1975).

The bat-fruit syndrome. — The relationship between bats and fruiting plants
is mutualistic. The plants expend energy on production of edible, nutritious
fruits as well as on olfactory and visual stimuli that attract bats. In eating fruits,
bats usually transport seeds away from the parental crown and discard them
at potential germination sites.

Fruit-eating bats and bat fruits have undergone considerable coevolution,
and the resulting set of adaptations are characterized as the “bat-fruit syndrome”
(Table 1). Exceptions to this syndrome occur, but when several or all characters
of the syndrome occur in a fruit, it is likely to be dispersed by bats.

BIOLOGY OF THE PHYLLOSTOMATIDAE

431

Table 1. — The bat-fruit syndrome (after Pijl, 1972).

Fruit characteristics Bat characteristics

1 . Strong, musty odor

2. Dull color, often green or brown, fruits
visually inconspicuous

3. Exposed position outside dense foliage on
periphery of branches or on pendulous branches

4. Attachment to tree through maturity

5. Hard skin or pulp to deter other frugivores
(many bat fruits are soft externally)

6. Requires animal agent to disperse seed

Good sense of smell
Large eyes for orientation,
probably color-blind
Approach fruit from air

Harvest fruit from tree, not ground
Strong dentition for tearing fruit

Carry fruits from fruiting
trees to night roosts

Fruiting plants are under selective pressure to attract seed dispersal agents
and repel or temporally avoid seed predators, such as squirrels and peccaries
that frequently eat fruit and seeds. Also, some animals may eat fruits yet
neither destroy the seeds nor disperse them, but simply discard seeds below
the parental crown. Ceboid monkeys often discard large seeds in this manner.
Mature fruits on the tree are available to bats, birds, and arboreal animals,
but not to terrestrial rodents, deer, and peccaries. Pendulant fruits of tree
species such as Cecropia (Fig. 2), exclude arboreal rodents, but not birds or
bats that pluck fruits in flight, nor monkeys that hang from prehensile tails
and arms, nor procyonids or primates that pull branches with their feet (see
Kaufmann, 1962). The strong odor of fruits attracts olfactory-orienting mammals,
whereas dull coloration camouflages fruits from visually orienting birds.
Fruits having hard edible parts or edible parts covered by a tough husk exclude
most birds but do not hinder fruit-eating bats with their strong teeth.

Seed survival and mortality. — Once a plant releases its fruit to a dispersal
agent, the chances of seed mortality are high. In response, many plants produce
vast numbers of seeds, a few of which survive losses to seed predators, parasites,
mechanical damage, and inhospitable germination sites. For seeds to be

dispersed successfully by bats, the following conditions are requisites: 1)

fruit harvest must occur at maturity; 2) seed displacement must be beyond the
crown of the parent plant (in some species); 3) seed deposition must be at a
site suitable for germination; and 4) seeds must not be severely damaged.

In our experience, frugivorous phyllostomatids select ripe fruit. Several
factors promote this pattern. Unripe fruits have little odor and would not
attract bats. Also unripe fruits are hard and difficult to chew, and some are
distasteful or toxic when immature. For example, some immature fruits of

Passiflora contain deadly cyanide compounds but when ripe are eaten by

mammals (Saenz and Nassar, 1972). Unripe figs contain latex, which is
gummy, and have a bad taste (to humans, at least). The great difference in
sugar content of green and ripe fruit (Snow, 1971) suggests that there is little

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selective advantage in exploiting unripe fruit. These mechanisms effectively
protect seeds until they are mature and viable.

Janzen (1971 cr, 1971 b) and Wilson and Janzen (1972) demonstrated that
seed predators cause heavy mortality of seeds falling under the parental
crown because the foraging or egg-laying strategy is to locate areas of high
seed density. Seed survival frequently is related to the seed’s ability to
escape predators in space. Figs that are not taken away by bats or other
dispersors, for example, are susceptible to heavy seed predation by lygaeid
bugs (Slater, 1972).

As a rule, the Phyllostomatidae and the paleotropical Pteropodidae carry
fruits from resource trees to night feeding roosts (Greenhall, 1956, 1965;
Nellis, 1971; Osmaston, 1965; Jones, 1972), which may change every few
days (D. Morrison, personal communication). Because hundreds, or even
thousands, of bats may come to large fruiting trees in a single night, the use
of feeding roosts may alleviate crowding and aggression at resource trees.
Additionally, this behavior may reduce the attractiveness of bat feeding
aggregations to predators such as owls and opossums.

Each plant species has particular requirements with respect to soil, nutrients,
drainage, and lighting conditions conducive to subsequent growth and
development. Once taken to a feeding roost, seeds are either discarded as the
fruit is eaten (somatochory) or ingested with fruit pulp (endochory).
Somatochores (Fig. 2C) have seeds too large to swallow, and their dispersal
by phyllostomatids is limited by the nature and location of night feeding roosts.
Of these, caves and buildings are particularly bad places for seeds to germinate.

Endochores (Fig. 2 A, B) have small, numerous seeds scattered through
the edible pulp. Endochores are eliminated with the feces and have the
potential to land any place a bat moves during a night. Alimentary passage
time is usually less than three hours for seeds (Arata, 1972; S. Farkas,
personal communication) and typically may be half an hour (Klite, 1965). Quick
seed passage time keeps bats at low flight weights and also ensures that
many seeds will be eliminated before the bat returns to the day roost, which
usually is a poor germination site.

Seed damage may result from mastication or digestion. Bats rarely damage
small seeds, and excreted seeds have a high germination rate (S. Gaulin,
personal communication). We know of only one species of large-seeded fruit
species, Anacardium excelsum, regularly damaged by bats. A. excelsum
seeds commonly are eaten by Carollia perspicillata, which acts as a seed
predator.

Pollination and the Bat-Flower Syndrome

Based on floral form, Vogal (1969) estimated that bats play some part
in pollination of at least 500 Neotropical plant species of 96 genera. It appears
that phyllostomatids increase in importance as pollinating agents from mesic
to xeric habitats. This pattern is the opposite of that of phyllostomatids acting

BIOLOGY OF THE PHYLLOSTOMATIDAE

433

Table 2. — The hat-flower syndrome ( after Faegri and Pijl, 1971).

Flower characteristics Bat characteristics

1. Nocturnal anthesis

2. Strong, musty odor

3. Dull color, often whitish, creamy, or purple

4. Exposed position outside dense foliage on
periphery of branches or on pendulous branches

5. Large flowers

6. Copious nectar and pollen production

7. Flower tube-like with anthers protruding,
or brush-shaped flower

Nocturnal foraging
Good sense of smell
Large eyes for orientation,
probably color-blind
Approach flower from air

Large body size compared to
other pollinators

High metabolic rate and large body
size

Elongate snout and protrusible tongue
for probing deep into flowers

as seed dispersal agents. Glossophaga soricina in Panama uses flower resources
for only five months of the year (Fleming et al., 1972), whereas in the dry
forest of Costa Rica, G. soricina uses flowers all year long (Heithaus et al .,
1975).

Flowers pollinated by bats are distinguished by drab colors, musty odors,
tube or brush shapes, position free of foliage, large size, nocturnal anthesis,
and copious nectar and pollen production. These floral characters (Fig. 2)
and corresponding adaptations found in nectarivorous bats are summarized
in Table 2. Caution must be taken because bats are highly opportunistic in
foraging habits and may sometimes take advantage of flowers not precisely
fitting the “bat-flower syndrome.” Furthermore, bats may ingest nectar or
pollen (or both) of a particular plant species and yet not provide pollination
services. Floral parasites are common in nature (for example, flower-piercing
hummingbirds and bees). Ratcliffe (1931) reported that flying foxes of
Australia eat entire flowers, but we have found no reports of phyllostomatids
regularly eating flowers. Baker et al. (1971) suggested that Leptonycteris
sanborni occasionally eats anthers.

The association between these bats and flowering plants is mutualistic.
Plants divert energy into production of odors and floral parts that attract
bats as well as nectar and pollen that feed bats. In moving from flower to flower
for food, bats transport some pollen, which results in fertilization.

It has long been obvious that flower bats obtain carbohydrate in the form of
sugars from floral nectaries. Recently it has been demonstrated that pollen is
an important source of protein to these bats (Howell, 1974). The cellular
contents of pollen grains begin to extrude through the micropores when pollen
begins to germinate in the gut. Protein then is extracted by hydrochloric acid
produced in the stomach, and protein is leached further by urea from urine
ingested by the bat (at least in Leptonycteris sanborni). Howell also found that
the protein content of pollen eaten by L. sanborni was 44 per cent for saguaro
and 23 per cent for paniculate agave, much higher than in pollen of closely

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related plants for which pollen is not eaten or dispersed by bats. Inasmuch
as herbivores must have a rich source of plant protein in order to maintain a
high rate of metabolism, the concentration of protein in pollen may be an
important avenue of coevolution.

If flower bats eat pollen, then how do they function as pollinators? While
probing the flower corolla for nectar, bats become dusted with pollen from
noseleaf to uropatagium (Baker, 1970). Phyllostomatids eat pollen only as they
groom their flight membranes and fur after a foraging bout (Howell and Hodgkin,
1976; Heithaus et al., 1975). Such behavior would provide for floral pollination
during visits to successive flowers while foraging and still permit the bat later
to eat the excess pollen covering its body.

Some bats adapted to the exploitation of flowers are known to feed on insects
and fruits. Whether insects are taken in the process of nectar-feeding or hunted
separately is unknown.

Impact of Predation by Bats

Little is known of the precise diets of insect or vertebrate-eating phyllostomatids,
so the impact of their predation on prey populations is undocumented. The
prominence of biotic limiting factors in the Neotropics suggests that investigation
of this impact is worthwhile.

Competition

Competition for sunlight or food is thought to be the dominant limiting
factor for Neotropical organisms (see Janzen, 1967, and MacArthur, 1969).
The likelihood of interspecific competition in tropical bats has been discussed
(Tamsitt, 1967; McNab, 1971; Dwyer, 1971; Fleming et al., 1972; Howell
and Burch, 1974; Heithaus et al., 1975), though documentation of such
competition awaits further study. Intraspecific competition for both food and
roost space are most likely to occur in the most colonial phyllostomatids,
mentioned above.

Enough is known about the three main categories of phyllostomatid food —
fruit, nectar and pollen, and insects — to discuss them briefly. Potential
competitors in all three categories include insects, birds, arboreal mammals,
and other bats, plus insectivorous spiders. Observations cited above of times
when food may be in short supply suggest when competition could be acute.

Fruit. — Because fruits are available on a “first come, first served” basis,
fruit searching success may be an important component of potential competition.
Birds feed heavily on ripe fruits of species eaten by bats. Two of the most
important bat fruits in Trinidad — Cecropia and Piper (Fig. 2) — are eaten in
significant quantities by tanagers and honeycreepers (Snow and Snow, 1971). For
example, Eisenmann (1961) recorded 24 species of birds feeding on Cecropia fruit
in Panama. Potential mammalian competitors include monkeys, marsupials,
rodents, and procyonids. Monkeys may be especially important in eating large
quantities of unripe fruit (Daubenmire, 1972).

BIOLOGY OF THE PHYLLOSTOMATIDAE

435

Nectar and pollen. — Diurnal birds and insects seldom compete with bats
for nectar and pollen, because many flowers are adapted for pollination
at one time of day and might not be open or produce nectar at other times (for
example, Bauhinia pauletia, Heithaus et al., 1974; Janzen, 1968). Thus, most
flowers used by Neotropical hummingbirds (Wolf, 1970; Snow and Snow,
1972) are not visited by bats. However, Baker et al. (1971) have shown that
Ceiba acuminata may be pollinated by both bats and hummingbirds, because
these flowers open at night but continue to secrete nectar the next day and are
visited by both kinds of animals. Balsa ( Ochroma ) flowers open at dusk
(Fig. 2) and are visited by phyllostomatid bats and sphingid moths. Some
flowers used by bats are destroyed when monkeys or insects eat them. Alvarez
and Gonzalez Q. (1970) concluded that little or no competition for flowers
occurs among six genera of glossophagines in Mexico. Heithaus et al. (1975)
also found nectarivorous bats to feed as generalists with high dietary overlap.

Insects. — Seasonally rapid recruitment rates for insects (as when a hatch
is under way) may enable bats to partition temporally a common resource
during the night. Additional partitioning is possible because insect taxa differ
in periodicity of night-time activity, at least in temperate zones (Williams,
1935, 1939; Lewis and Taylor, 1965). Potential night-time competitors
include spiders, tree frogs, caprimulgiform birds, owls, night monkeys,
marsupials, rodents, and procyonids.

Roost space. — We know of no published evidence of phyllostomatids
competing for roost space. However, such competition frequently may be
provided by the more colonial taxa. On Barro Colorado Island, Panama, a
group of Phyllostomus hastatus displaced a hollow tree colony of Carollia
perspicillata (S. Graetz, personal communication). F. J. Bonaccorso observed,
on the same island, a displacement of C. perspicillata and Saccopteryx bilineata
from their hollow tree roost by a colony of Desmodus rotundas.

Predation on Bats

Little is known about causes of phyllostomatid mortality or the food habits
of their potential predators. Reviewers of temperate zone data judge that
predation on bats is opportunistic but seldom regular (Allen, 1939; Gillette
and Kimbrough, 1970). In the New World tropics, predation on bats may
well be more important, in view of the general prominence of biotic interactions
and the dominant numbers of bats in mammal faunas. In Haiti, 27 of 147 prey
items of a Hispanolean barn owl ( Tyto glaucops) were phyllostomatid bats
(Wetmore and Swales, 1931). In Panama, three species of owls have killed
bats in our nets. Arboreal opossums, procyonids, and snakes may wait for
bats visiting resource trees. Opossums ( Didelphis virginiana and Philander
opossum) are known to eat bats (Campbell, 1925; Rice, 1957; our observations).
The bat falcon, Falco rufigularis, may specialize on bat prey. The largest
phyllostomatid bats ( V ampyrum spectrum, Chrotopterus auritus, and
Phyllostomus hastatus) are suspected or known to eat smaller bats (Goodwin
and Greenhall, 1961 ; Valdivieso, 1964; Greenhall, 1968).

436

SPECIAL PUBLICATIONS MUSEUM TEXAS TECH UNIVERSITY

An important clue to the role of predation on phyllostomatids may be
the inverse relationship between moonlight and flight activity of Desmodus
rotundus (Crespo et al. , 1972; Turner, 1975) and Artibeus jamaicensis
(Morrison, 1975). Our qualitative observations at several Central American
sites are that phyllostomatid foraging is characteristically maximal when
no moonlight is incident and minimal under full moonlight. To explain
avoidance of moonlight as a response to heightened predator success would
be uncomplicated for bats that feed on plants; the behavior of predatory bats
must additionally account for the possibility of similar responses by their own
prey species.

Acknowledgments

We appreciate the helpful criticisms of R. B. Foster, D. J. Howell, E. Leigh,
J. J. McManus, P. A. Opler, and C. M. Simon, and the art work of N. Halliday
and S. J. Scudder. Contributions of unpublished data by G. W. Frankie, E. R.
Heithaus, D. J. Howell, B. K. McNab, D. Morrison, and N. Smythe have
enabled us to explain much more tropical ecology than is possible from the
presently available literature. Our own data were acquired under the following
grant support : National Institutes of Health Biomedical Sciences grant no.
RR 7021-07 from the University of Florida Division of Sponsored Research
to S. R. Humphrey; National Science Foundation grant no. GB-36068 to
J. H. Kaufmann; and Smithsonian Tropical Research Institute grants to F. J.
Bonaccorso.

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phidae) in North and Middle America, 81 pp., 14 figs . $2.00

Genoways, H. H. 1973. Systematics and evolutionary relationships of spiny pocket
mice, genus Liomys, 368 pp., 66 figs . $7.00

Northington, D. K. 1974. Systematic studies of the genus Pyrrhopappus (Com-
positae, Cichorieae), 38 pp., 14 figs . $1.00

King, M. E., and I. R. Traylor, Jr., eds. 1974. Art and environment in native
America. 169 pp . $5.00

Pence, D. B. 1975. Keys, species and host list, and bibliography for nasal mites of
North American birds (Acarina: Rhinonyssinae, Turbinoptinae, Speleognathi-
nae, and Cytoditidae), 148 pp., 728 figs . $4.00

Bowles, J. B. 1975. Distribution and biogeography of mammals of Iowa, 184 pp.,
62 figs . $5.00

Baker, R. J., J. K. Jones, Jr., and D. C. Carter, eds. 1976. Biology of bats of the
New World family Phyllostomatidae. Part I., 218 pp . $6.00

Foster, D. E. 1976. Revision of North American Trichodes (Herbst) (Coleoptera:
Cleridae), 86 pp., 17 figs . $2.00

Mitchell, R. W., W. H. Russell, and W. R. Elliott. 1977. Mexican eyeless chara-
cin fishes, genus Astyanax: environment, distribution, and evolution, 89 pp.,
21 figs . $5.00

Baker, R. J., J. K. Jones, Jr., and D. C. Carter, eds. 1977. Biology of bats of the
New World family Phyllostomatidae. Part II., 364 pp . $16.00

Francke, O. F. 1978. Systematic revision of diplocentrid scorpions (Diplocentridae)
from circum-Caribbean lands, 93 pp., 146 figs . $7.00

Carter, D. C., and P. G. Dolan. 1978. Catalogue of type specimens of Neotropical
bats in selected European museums, 136 pp . $8.00

Baker, R. J., J. K. Jones, Jr., and D. C. Carter, eds. 1979. Biology of bats of the
New World family Phyllostomatidae. Part III., 441 pp . $20.00

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