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Morphological Phylogeny of the Bat Genus Platyrrhinus Saussure, 1860 (Chiroptera: Phyllostomidae) with the Description of Four New Species

November 2005
Fieldiana Zoology 105(Nov 2005):1-53

November 2005
105(Nov 2005):1-53

DOI:10.3158/0015-0754(2005)105[1:MPOTBG]2.0.CO;2

Authors:

Arcadia University

Platyrrhinus, comprising 10 species of broad-nosed bats, is one of the most diverse genera in the Neotropical family Phyllostomidae. To evaluate the content and limits of this genus, the phylogenetic relationships among these species and other closely related taxa were examined using a data set of 60 morphological characters including external, cranial, dental, and post-cranial characters. Four species (Carollia subrufa, Sturnira erythromos, Uroderma magnirostrum, and Vampyrodes caraccioli) of the family Phyllostomidae were used as outgroups to test the monophyly of Platyrrhinus and to resolve intrageneric relationships. Parsimony analysis resulted in two most parsimonious trees. The results support Platyrrhinus monophyly with a bootstrap value of 84% and a Bremer support of 3. Three synapomorphies are recognized for the genus. The results also support the recognition of P. umbratus as a junior synonym of P. dorsalis, the division of P. dorsalis into three distinct species, and the discovery of two new species previously considered part of P. helleri and P. vittatus. Taxonomic diagnoses of all the species of Platyrrhinus based on character optimizations as well as descriptions of four new species are presented. This revision brings the total number of species of Platyrrhinus to 14, making it the most speciose genus in the family Phyllostomidae.

. List of examined taxa.

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. Selected measurements a,b of members of the genus Platyrrhinus.

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. Continued.

…

. Selected measurements a,b of Platyrrhinus ismaeli and P. masu.

…

Posterior view of the temporal region, showing details of zygomatic arch (ZA) and fossa on the squamosal end of the zygomatic arch: (A) Platyrrhinus brachycephalus (MUSM 13793); (B) P. infuscus (MUSM 9897); (C) P. dorsalis (FMNH 128141). Scale bar 4 mm.

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Zoology

NEW SERIES, NO. 105

Morphological Phylogeny of the Bat Genus

Platyrrhinus Saussure, 1860 (Chiroptera:

Phyllostomidae) with the Description

of Four New Species

Paul M. Velazco

Division of Mammals

Department of Zoology

Field Museum of Natural History

1400 South Lake Shore Drive

Chicago, Illinois 60605-2496

U.S.A.

Department of Biological Sciences

University of Illinois at Chicago

Chicago, Illinois 60680-4348

U.S.A.

Accepted July 22, 2005

Published November 15, 2005

Publication 1535

2005 Field Museum of Natural History

ISSN 0015-0754

PRINTED IN THE UNITED STATES OF AMERICA

iii

Table of Contents

BSTRACT

............................................... 1

NTRODUCTION

.......................................... 1

ACKGROUND

........................................... 2

ATERIALS AND

ETHODS

.......................... 4

Taxonomic Sample and Outgroup Deter-

mination ........................................ 4

Sources of Data .................................... 5

Deﬁnition of the Characters and Ordering

of Character States .......................... 5

Polarity ............................................... 5

Methods of Phylogenetic Analysis ............ 5

HARACTER

ESCRIPTIONS

........................... 6

Pelage and Integument ........................... 6

Skull .................................................. 10

Dentition ............................................ 12

Postcranial Skeleton .............................. 17

ESULTS

................................................. 18

ISCUSSION AND

ONCLUSIONS

.................... 18

Taxonomic Diagnoses of the Platyrrhinus

Species ......................................... 20

CKNOWLEDGMENTS

.................................. 42

ITERATURE

ITED

.................................... 42

PPENDIX

1: S

PECIMENS

XAMINED

.............. 45

PPENDIX

2: D

ATA

ATRIX

........................ 51

PPENDIX

3: O

PTIMIZATION OF

HARACTERS

... 51

List of Illustrations

1. Frontal view of Platyrrhinus sp. nov. ...... 2

2. Owen’s (1987; redrawn from Fig. 17)

hypothesis of relationships among Pla-

tyrrhinus species, Sturnira erythromos,

Uroderma magnirostrum, Chiroderma

villosum, and Vampyrodes caraccioli ..... 3

3. Schematic view of the heads of three

hypothetical bats ................................ 7

4. Ventral view of the chin of ﬁve hypo-

thetical bats ....................................... 8

5. Frontal view of the noseleaf of Sturni-

ra erythromos, Platyrrhinus infuscus,

and P. nigellus ................................... 9

6. Detail of the posterior margin of the

hard palate of Platyrrhinus brachyceph-

alus and P. helleri ‘‘Western’’ ............. 10

7. Detail of the postorbital processes of

Platyrrhinus infuscus and P. nigellus .... 11

8. Detail of the mastoid and paraoccipital

processes of Platyrrhinus helleri ‘‘West-

ern,’’ P. nigellus, P. vittatus ‘‘South-

ern,’’ and Sturnira erythromos .............. 11

9. Detail of the fossa on the squamosal

end of the zygomatic arch of Platyr-

rhinus brachycephalus, P. dorsalis, and

P. infuscus ....................................... 12

10. Detail of the posterior clinoid process-

es of Platyrrhinus infuscus and Uro-

derma magnirostrum .......................... 13

11. Occlusal view of the maxillary tooth-

row of Platyrrhinus infuscus and Stur-

nira erythromos ................................ 13

12. Lateral view of the maxillary toothrow

of Platyrrhinus infuscus ...................... 14

13. Occlusal view of M1-M2 of Pla-

tyrrhinus infuscus and P. dorsalis

‘‘Norte’’ ........................................... 15

14. Strict consensus tree of the morpholog-

ical analysis with the characters unor-

dered ............................................... 19

15. Distributional map of Platyrrhinus al-

bericoi sp. nov. ................................. 21

16. Labial views of the left p2 and p4 of

Platyrrhinus vittatus and P. albericoi

sp. nov. ........................................... 25

17. Distributional map of Platyrrhinus au-

rarius and P. brachycephalus .............. 26

18. Distributional map of Platyrrhinus cho-

coensis and P. dorsalis ....................... 27

19. Distributional map of Platyrrhinus hel-

leri ................................................. 28

20. Distributional map of Platyrrhinus in-

fuscus and P. lineatus ........................ 29

21. Frontal view of Platyrrhinus ismaeli

sp. nov. ........................................... 30

22. Distributional map of Platyrrhinus is-

maeli sp. nov. ................................... 31

23. Occlusal views of the left M1 and M2

of Platyrrhinus ismaeli sp. nov. and P.

dorsalis ........................................... 33

24. Frontal view of Platyrrhinus masu sp.

nov. ................................................ 34

25. Distributional map of Platyrrhinus

masu sp. nov. ................................... 35

26. Distributional map of Platyrrhinus ma-

tapalensis sp. nov. ............................. 38

27. Labial views of the left p2 and p4 of

Platyrrhinus matapalensis sp. nov., P.

helleri, and P. brachycephalus ............. 40

28. Distributional map of Platyrrhinus ni-

gellus, P. reciﬁnus and P. vittatus ........ 41

29. Strict consensus tree with nodes num-

bered as referenced in Appendix 3 ....... 52

List of Tables

1. Taxa examined .................................... 4

2. Measurements of the type series of Pla-

tyrrhinus albericoi sp. nov. ................... 21

3. Selected measurements of members of

the genus Platyrrhinus ......................... 22

4. Measurements of the type series of Pla-

tyrrhinus ismaeli sp. nov. ..................... 30

5. Measurements of the type series of Pla-

tyrrhinus masu sp. nov. ........................ 34

6. Selected measurements of Platyrrhinus

ismaeli sp. nov. and P. masu sp. nov. ..... 36

7. Measurements of the type series of Pla-

tyrrhinus matapalensis sp. nov. .............. 37

FIELDIANA: ZOOLOGY, N.S., NO. 105, NOVEMBER 15, 2005, PP. 1–54 1

Morphological Phylogeny of the Bat Genus

Platyrrhinus Saussure, 1860 (Chiroptera:

Phyllostomidae) with the Description of

Four New Species

Paul M. Velazco

Abstract

Platyrrhinus, comprising 10 species of broad-nosed bats, is one of the most diverse genera

in the Neotropical family Phyllostomidae. To evaluate the content and limits of this genus, the

phylogenetic relationships among these species and other closely related taxa were examined

using a data set of 60 morphological characters including external, cranial, dental, and post-

cranial characters. Four species (Carollia subrufa, Sturnira erythromos, Uroderma magniros-

trum, and Vampyrodes caraccioli) of the family Phyllostomidae were used as outgroups to test

the monophyly of Platyrrhinus and to resolve intrageneric relationships.

Parsimony analysis resulted in two most parsimonious trees. The results support Platyrrhinus

monophyly with a bootstrap value of 84% and a Bremer support of 3. Three synapomorphies

are recognized for the genus. The results also support the recognition of P. umbratus as a junior

synonym of P. dorsalis, the division of P. dorsalis into three distinct species, and the discovery

of two new species previously considered part of P. helleri and P. vittatus. Taxonomic diag-

noses of all the species of Platyrrhinus based on character optimizations as well as descriptions

of four new species are presented. This revision brings the total number of species of Platyr-

rhinus to 14, making it the most speciose genus in the family Phyllostomidae.

Introduction

The genus Platyrrhinus Saussure, 1860, be-

longs to the family Phyllostomidae (Mammalia:

Chiroptera). The distribution of this family is re-

stricted to the Neotropics. Morphologically, it is

characterized by the presence of a noseleaf, de-

veloped in most of its members, as well as the

presence of two synapomorphic characters that

differentiate it from the other Microchiropterans:

oviductal folds restricted to the extramural ovi-

duct (Hood & Smith, 1982) and friction lock in

the digits of feet (Simmons & Quinn, 1994; Sim-

mons, 1998). Within the Phyllostomidae, Platyr-

rhinus is placed in the subfamily Stenodermati-

nae, which is diagnosed by a combination of the

presence of short and ﬂattened muzzle, short and

wide nose leaf, narrow uropatagium, lack of tail,

and frequent presence of facial and/or dorsal

stripes (Linares, 1986; Albuja, 1999). This sub-

family also has two synapomorphic characters: a

single common uterine lumen, without remnants

of cornual lumina, and the oviducts enter the fun-

dic border of the uterine body near the midsagittal

line (Hood & Smith, 1982).

The species of Platyrrhinus (Fig. 1) are thought

to be monophyletic (Owen, 1987; Lim, 1993;

Wetterer et al., 2000). Platyrrhinus is diagnosed

from other Stenodermatinae by a combination of

three characters: two accessory cusps on the pos-

terior face of P4, presence of three upper molars,

and presence of a fringe of hair on the edge of

the uropatagium. Although other genera also have

these characters, no other genus possesses all

three (Lim, 1993; Albuja, 1999).

Ten nominal species of Platyrrhinus are com-

monly recognized (Owen, 1987; Koopman,

1993; Nowak, 1999). The distributional range of

2 FIELDIANA: ZOOLOGY

. 1. Frontal view of Platyrrhinus sp. nov. (FMNH 172108); adult male photographed at Pillahuata in the

Cultural zone of the Manu Biosphere Reserve, Cuzco, Peru, by P. M. Velazco.

Platyrrhinus includes North, Central, and South

America, from southern Mexico to Bolivia, Par-

aguay, Uruguay, and southeastern Brazil (San-

born, 1955; Cabrera, 1957; Jones & Carter, 1976;

Hall, 1981; Koopman, 1982, 1993). Some spe-

cies of this genus are widely distributed—for ex-

ample, P. helleri is distributed from southern

Mexico to Peru, Bolivia, and Brazil as well as

Trinidad and Tobago (Koopman, 1982; Ferrell &

Wilson, 1991).

The genus Platyrrhinus presents difﬁculties

both in terms of its diagnosis and in how many

species it comprises. The present work seeks to

elucidate the phylogenetic relationships among

the members of Platyrrhinus. To do so, cladistic

analyses of discrete morphological characters

were conducted. External, cranial, dental, and

postcranial skeletal characters were examined.

The results from these analyses offer a new inter-

pretation of morphological diversity within the

genus and a reassessment of the hypotheses of

relationships presented by Owen (1987), Lim

(1993), Baker et al. (2000, 2003), and Wetterer et

al. (2000).

Background

All the species of Platyrrhinus Saussure, 1860,

described before 1990 were recognized under the

genus Vampyrops Peters, 1865. Gardner and Fer-

rell (1990) reviewed the nomenclatural history of

these bats and determined that Platyrrhinus and

Vampyrops were synonymous and that Platyr-

rhinus had priority over Vampyrops.

Koopman (1993) recognized 10 nominal spe-

cies of Platyrrhinus:P. aurarius (Handley & Fer-

ris, 1972), endemic to the Guianan Shield (south-

ern Venezuela, Guyana, and Suriname); P. bra-

chycephalus (Rouk & Carter, 1972), distributed in

the Amazon and the coasts of northern South

America and adjacent islands (northern Brazil,

Colombia to Guyana, Ecuador, Peru, andBolivia);

P. chocoensis Alberico and Velasco, 1991, dis-

tributed in the Choco´ region (western Colombia,

in lowlands between the western slope of the An-

des Cordillera and the Paciﬁc coast); P. dorsalis

(Thomas, 1900), distributed on both slopes of the

Andes Cordillera and Central America (Panama

to Peru and Bolivia); P. helleri (Peters, 1866), dis-

3VELAZCO: PLATYRRHINUS PHYLOGENY

. 2. Owen’s (1987; redrawn from Fig. 17) hy-

pothesis of relationships among Platyrrhinus species,

Sturnira erythromos, Uroderma magnirostrum, Chirod-

erma villosum, and Vampyrodes caraccioli, based on a

consensus of continuous and discrete characters.

tributed from Central America (Oaxaca and Ve-

racruz [Mexico]), south to Peru, Bolivia, the Am-

azon of Brazil, and Trinidad and Tobago; P. in-

fuscus (Peters, 1880), distributed on the eastern

slope of the north part of the Andes Cordillera

and in the Amazon (Colombia to Peru, Bolivia,

and northwestern Brazil); P. lineatus (E

´. Geof-

froy, 1810), distributed in the Pampean subregion,

Amazon, and the eastern slope of the north part

of the Andes Cordillera (Colombia to Peru, Bo-

livia, Uruguay, northern Argentina, and southern

and eastern Brazil); P. reciﬁnus (Thomas, 1901),

distributed in the highlands and coast of eastern

Brazil and the Amazon (eastern Brazil); P. um-

bratus (Lyon, 1902), distributed in the eastern

slope of the northern Andes Cordillera, north

coast of South America and adjacent islands, and

the Paciﬁc coast of Ecuador and Colombia (Pan-

ama, Colombia, and northern Venezuela); and P.

vittatus (Peters, 1860), distributed on the eastern

slope of the north part of the Andean Cordillera,

north coast and islands of South America, and

Central America (Costa Rica to Venezuela and

from Peru to Bolivia) (Koopman 1982, 1993; Al-

berico 1990; Lim & Engstrom, 2000).

Sanborn (1955), Gardner and Carter (1972b),

and Carter and Rouk (1973) considered Platyr-

rhinus umbratus to be conspeciﬁc with P. dorsa-

lis, synonymizing the former with the latter. How-

ever, Handley (1976) reported that P. umbratus

differed from P. dorsalis, which was followed by

Koopman (1993) and Owen (1987). Velazco and

Solari (2003) considered P. aquilus, P. oratus,

and P. umbratus as junior synonyms of P. dor-

salis, and they stated that P. dorsalis as it is now

recognized is a complex of three taxa. One of

these corresponds to P. dorsalis (Thomas, 1900),

with aquilus, oratus, and umbratus as junior syn-

onyms; the other two taxa are undescribed spe-

cies, denoted P. dorsalis ‘‘Norte’’ and P. dorsalis

‘‘Centro-Sur.’’ Platyrrhinus dorsalis ‘‘Norte’’ is

distributed on both slopes of the Andes in Ecua-

dor and Peru, and P. dorsalis ‘‘Centro-Sur’’ is dis-

tributed on the eastern slope of the Andes in Peru

(Velazco & Solari, 2003).

Tuttle (1970) regarded Platyrrhinus infuscus as

a junior synonym of P. vittatus, but Gardner and

Carter (1972b) and Koopman (1978) considered

P. infuscus distinct from P. vittatus. Gardner and

Carter (1972a) described P. nigellus as a species

present in Colombia and probably Ecuador and

Peru, but Koopman (1978, 1982), Jones and Cart-

er (1979), and Willig and Hollander (1987) treated

nigellus as a subspecies of P. lineatus. Owen

(1987) and Velazco and Solari (2003) recognized

P. nigellus as a species distinct from P. lineatus.

The ﬁrst explicit hypothesis of phylogenetic re-

lationships within Platyrrhinus was presented by

Owen (1987; Fig. 2) in the context of his analysis

of the subfamily Stenodermatinae. That analysis

has been criticized both generally (Lim, 1993) as

well as for speciﬁc genera included in his analysis

(Pacheco & Patterson, 1991). In that analysis,

Owen used 22 discrete and 33 continuous exter-

nal, cranial, and dental characters.

Owen’s preferred phylogenetic hypothesis

(Owen, 1987; Fig. 2) placed the monotypic genus

Vampyrodes as a sister genus of Platyrrhinus. He

considered Platyrrhinus to be a monophyletic

group and indicated two clades that were not re-

solved within Platyrrhinus, one containing P. ni-

gellus

P. helleri

P dorsalis and the other P.

vittatus

P. aurarius

P. lineatus

P. umbra-

tus. The relationships of the remaining species in

the tree (P. brachycephalus, P. infuscus, and P.

reciﬁnus) also were unresolved. Lim (1993) sub-

sequently argued that Owen (1987) did not judi-

ciously select the best characters for his cladistic

analysis and used distance methods not suitable

for the reconstruction of evolutionary history.

Lim (1993) used 20 external, cranial, dental,

and internal morphological characters to generate

a hypothesis of relationships at the generic level

for the Stenodermatinae. In this analysis, he ob-

4 FIELDIANA: ZOOLOGY

ABLE

1. List of examined taxa.

Ingroup

Playtrrhinus aurarius

Platyrrhinus brachycephalus

Playtrrhinus chocoensis

Platyrrhinus dorsalis

Platyrrhinus dorsalis ‘‘Norte’’

Platyrrhinus dorsalis ‘‘Centro-Sur’’

Platyrrhinus helleri ‘‘Eastern’’

Platyrrhinus helleri ‘‘Western’’

Platyrrhinus infuscus

Platyrrhinus lineatus

Platyrrhinus nigellus

Platyrrhinus reciﬁnus

Platyrrhinus vittatus ‘‘Northern’’

Platyrrhinus vittatus ‘‘Southern’’

Outgroups

Carollia subrufa

Sturnira erythromos

Uroderma magnirostrum

Vampyrodes caraccioli

tained Platyrrhinus

Vampyrodes as sister taxa,

supporting Owen’s (1987) hypothesis. Two char-

acters unite these taxa: one unambiguous syna-

pomorphy (fringe of hair on the edge of the uro-

patagium) and one parallel synapomorphy, which

is also present in Uroderma (accessory cusps on

P4). Lim (1993) found that the sister group to this

pair includes Ectophylla

Mesophylla

Vam-

pyressa

Chiroderma.

Wetterer et al. (2000) evaluated the phyloge-

netic relationships among the genera of the family

Phyllostomidae using 150 morphological, karyo-

logical, and molecular characters. In this analysis,

they also found Platyrrhinus and Vampyrodes to

be sister taxa, supported with a bootstrap value of

46. Their ACCTRAN (accelerated transformation

optimization) analysis indicated ﬁve synapomor-

phies for this grouping: uropatagial fringe pres-

ence, I1 occlusal margin generally straight or

slightly rounded, m1 paraconid present, pharyn-

geal region of tongue completely covered with pa-

pillae, cerebral vermis completely covering the

longitudinal ﬁssure between inferior colliculi, and

inferior colliculi exposed dorsally only along lat-

eral edges of cerebelar vermis. Their DELTRAN

(delayed transformation optimization) analysis

also identiﬁed three of these as synapomorphies:

uropatagial fringe presence, M1 hypocone pre-

sent, and pharyngeal region of tongue completely

covered with papillae. In their analysis Uroderma

was sister to the clade Platyrrhinus

Vampyro-

des, and Ectophylla

Vampyressa

Chiroderma

was sister to the clade Uroderma

Platyrrhinus

Vampyrodes.

Baker et al. (2000, 2003) evaluated the phylo-

genetic relationships of the family Phyllostomidae

at the generic level using nuclear and mitochon-

drial genes. In those analyses they also found Pla-

tyrrhinus and Vampyrodes to be sister taxa. More-

over, Mesophylla and Vampyressa were found to

be the sister of Platyrrhinus

Vampyrodes.

Materials and Methods

Taxonomic Sample and Outgroup

Determination

For this cladistic analysis, the taxonomy of

Koopman (1993) was followed with the exception

of P. dorsalis, P. lineatus, and P. umbratus. For

these species, Velazco and Solari(2003), who rec-

ognized three distinct morphs within P. dorsalis

(two not yet described; they are here denoted P.

dorsalis, P. dorsalis ‘‘Norte,’’ and P. dorsalis

‘‘Centro-Sur’’), was followed. They also recog-

nized P. nigellus as a valid species and considered

P. umbratus as a junior synonym of P. dorsalis.

After the character analysis, two different popu-

lations in P. helleri and P. vittatus were recog-

nized. For the purposes of the analysis herein,

they are denoted as P. helleri ‘‘Eastern,’’ P. hel-

leri ‘‘Western,’’ P. vittatus ‘‘Northern,’’ and P.

vittatus ‘‘Southern.’’ A list of all 14 ingroup and

4 outgroup taxa is given in Table 1.

Only one outgroup is necessary to root a phy-

logenetic tree (Nixon & Carpenter, 1993), but to

establish the polarity of ambiguous characters and

to test Platyrrhinus monophyly, four taxa were

used as outgroups: Carollia subrufa (Hahn, 1905),

Sturnira erythromos (Tschudii, 1844), Uroderma

magnirostrum Davis, 1968, and Vampyrodes car-

accioli (Thomas, 1889). Freeman (2000) consid-

ered Carollia as the most ancestral and least spe-

cialized genus in the tribe Stenodermatini. Wet-

terer et al. (2000) found that Carollia

Rhino-

phylla is the sister group of the subfamily

Stenodermatinae; they placed Sturnira as the most

basal taxon in the Stenodermatinae. Owen (1987),

Lim (1993), Baker et al. (2000, 2003), and Wet-

terer et al. (2000) all agreed that Vampyrodes was

the sister genus of Platyrrhinus. With respect to

the position of Uroderma, Owen (1987) placed it

in a clade together with Ectophylla and Artibeus,

Lim (1993) placed Uroderma in a clade together

with Artibeus, and Wetterer et al. (2000) placed

5VELAZCO: PLATYRRHINUS PHYLOGENY

Uroderma as the sister taxon of the clade Platyr-

rhinus

Vampyrodes. Baker et al. (2003) placed

Uroderma as sister to the clade Mesophylla

Ec-

tophylla

Platyrrhinus

Vampyrodes.

Sources of Data

Only adult specimens were examined based on

Morris (1972), Anthony (1988), and Pacheco and

Patterson (1992), and all the specimens belong to

scientiﬁc collections of museums (skins, skulls,

skeletons, and alcohol-preserved specimens). A

complete list of the specimens examined is pro-

vided in Appendix 1.

A total of 1,314 specimens were examined, be-

longing to the scientiﬁc collections of the follow-

ing museums; acronyms from museums holding

holotypes of Platyrrhinus species were also in-

cluded:

AMNH, American Museum Natural of History,

New York, New York, USA; BMNH, Natural

History Museum, London, England; EPN, Escue-

la Polite´cnica Nacional, Quito, Ecuador; FMNH,

Field Museum of Natural History, Chicago, Illi-

nois, USA; IND-M, La Unidad de Investigacio´n

‘‘Federico Medem’’—Inderena, Bogota´, Colom-

bia; LSUMZ, Louisiana State University, Muse-

um of Natural Science, Baton Rouge, Louisiana,

USA; MCZ, Museum of Comparative Zoology,

Harvard University, Cambridge, Massachusetts,

USA; MNHN, Muse´um National d’Histoire Na-

turelle, Paris, France; MSB, Museum of South-

western Biology, University of New Mexico, Al-

buquerque, New Mexico, USA; MUSM, Museo

de Historia Natural, Universidad Nacional Mayor

de San Marcos, Lima, Peru; MVZ, Museum of

Vertebrate Zoology, University of California,

Berkeley, California, USA; QCAZ, Pontiﬁcia

Universidad Cato´lica del Ecuador, Museo de

Zoologı´a—Mamı´feros, Quito, Ecuador; ROM,

Royal Ontario Museum, Toronto, Ontario, Cana-

da; TCWC, Texas Cooperative Wildlife Collec-

tion, Texas A&M Collection, College Station,

Texas, USA; UMMZ, University of Michigan,

Museum of Zoology, Ann Arbor, Michigan, USA;

USNM, National Museum of Natural History,

Smithsonian Institution, Washington, D.C., USA;

UV, Universidad del Valle, Cali, Colombia;

ZMB, Museum fu¨r Naturkunde der Humboldt-

Universita¨t zu Berlin, Berlin, Germany.

The number of specimens per species examined

in this study varied according to their availability

in these museum collections; each taxon was rep-

resented by at least 12 individuals.

Deﬁnition of the Characters and Ordering of

Character States

Autapomorphies were included in the phylo-

genetic analysis for the species of Platyrrhinus

because they are useful in diagnosis. In addition,

relevant characters in the relationships among the

outgroups were included to provide resolution. In

cases where variation inside the species is present,

the character was scored as polymorphic. Sim-

mons and Conway (2001) were followed for the

categorization of missing data.

The ﬁnal data set comprised a total of 60 char-

acters: 17 characters of the pelage and integu-

ment, 6 of the skull, 35 dental characters, and 2

characters of the postcranial skeleton (Appendix

2). Fifty-eight percent (35) of the characters were

binary, and 42% (25) of the characters were mul-

tistate.

Polarity

The ‘‘outgroup method’’ was used to establish

the polarity of the character states (Watrous &

Wheeler, 1981). On the basis of earlier studies,

outgroup species formed the sequence Carollia

subrufa, Sturnira erythromos, Uroderma magni-

rostrum, and Vampyrodes caraccioli. Carollia

subrufa was used to root the tree because it is

considered ancestral to the remaining three out-

group species (Freeman, 2000).

Methods of Phylogenetic Analysis

The matrix elaboration and the visualization of

the character mapping were performed using

MacClade 4.0 (Maddison & Maddison, 2000).

The phylogenetic analysis was performed using

PAUP version 4.0b6 (Swofford, 2001) on a Mac-

intosh PowerBook G4.

The branch and bound option of PAUP was

used to identify the most parsimonious trees with

the smallest number of steps and to calculate tree

statistics. The analysis was performed using all

characters unordered. A strict consensus was per-

formed because more than one parsimonious tree

was obtained. To perform the decay analysis (Bre-

mer, 1988), the program TreeRot version 2b (So-

6 FIELDIANA: ZOOLOGY

renson, 1999) was used. A branch and bound

bootstrap analysis was performed, with 10,000

replications, to evaluate the support of the result-

ing clades.

Character optimizations were calculated using

both the ACCTRAN and DELTRAN. The results

of these optimizations are presented in Appen-

dix 3.

Character Descriptions

Pelage and Integument

Character 1—Facial stripes. Absent (0); pre-

sent (1). The facial stripes are present as two lines

on each side of the face. The dorsomedial stripe

begins at the posterolateral edge of the noseleaf,

continues above the eye, and terminates close to

the pinna. The ventrolateral stripe begins in the

corner of the mouth and runs toward the tragus.

The facial stripes are present in all the species of

Platyrrhinus, Vampyrodes caraccioli, and Urod-

erma magnirostrum. In contrast, the facial stripes

are absent in Carollia subrufa and Sturnira ery-

thromos.

Owen (1987: character 4) uses the facial stripes

as part of a multistate character that he termed

‘‘Pelage patterns.’’ In this character Owen in-

cludes facial stripes, dorsal stripe, shoulders

patches, and striping on the wing. I agree with

Owen on his facial stripes scoring for the species

used in the present analysis. Lim (1993: character

1) and Wetterer et al. (2000: character 6) include

facial stripes in their phylogenetic analyses; char-

acter states and scoring used in this analysis co-

incide with theirs.

Character 2—Brightness of facial stripe. Dor-

somedial and ventrolateral stripes obviously sub-

equal (0); dorsomedial stripe more marked than

the ventrolateral stripe (1). In most Platyrrhinus

species, both facial stripes have the same tonality,

either dark or brilliant (e.g., P. aurarius, P. helleri

‘‘Eastern,’’ P. nigellus). In contrast, P. vittatus

‘‘Northern’’ and P. vittatus ‘‘Southern’’ have a

more luminous dorsomedial stripe. Carollia sub-

rufa and Sturnira erythromos were scored as ‘‘—

’’ because they do not have facial stripes. This is

the ﬁrst use of this character in a phylogenetic

analysis.

Character 3—Facial stripe coloration. Bril-

liant-white (0); dark (1). The facial stripes are

brilliant or white in P. brachycephalus, P. helleri

‘‘Eastern,’’ P. helleri ‘‘Western,’’ P. lineatus, P.

reciﬁnus, P. vittatus ‘‘Southern,’’ Vampyrodes

caraccioli, and Uroderma magnirostrum. In con-

trast, they are dark in P. aurarius, P. chocoensis,

P. dorsalis, P. dorsalis ‘‘Norte,’’ P. dorsalis

‘‘Centro-Sur,’’ P. infuscus, P. nigellus, and P. vit-

tatus ‘‘Northern.’’ Carollia subrufa and Sturnira

erythromos were scored as ‘‘—’’ because they

lack both facial stripes. This is the ﬁrst use of this

character in a phylogenetic analysis.

Character 4—Basal protuberance where the

genal vibrissae are implanted. Absent (0); pre-

sent (1). The genal vibrissae are implanted in the

cheek, ventral, and/or posterior to the eye. The

genal vibrissae are present in all the species of

Platyrrhinus. These vibrissae are implanted in a

basal protuberance in some Platyrrhinus (e.g., P.

brachycephalus, P. helleri ‘‘Eastern,’’ P. lineatus)

(Fig. 3A). In contrast, the genal vibrissae are im-

planted directly on the cheek in Carollia subrufa

and some species of Platyrrhinus (e.g., P. aurar-

ius, P. chocoensis) (Fig. 3B–C). This character

has not been used in previous studies.

Character 5—Number of vibrissae surround-

ing the margins of the noseleaf in a single ar-

ray. Nine vibrissae present (0); eight vibrissae

present (1); seven vibrissae present (2); six vi-

brissae present (3). Some species of Platyrrhinus

have seven vibrissae surrounding the margins of

the noseleaf in a single array (e.g., P. brachyce-

phalus, P. chocoensis, P. lineatus) (Fig. 3B). In

contrast, six vibrissae are presented to each side

of the noseleaf in some Platyrrhinus and Vam-

pyrodes (e.g., P. aurarius, P. dorsalis) (Fig. 3A).

Platyrrhinus helleri ‘‘Eastern’’ presents intraspe-

ciﬁc variation (seven and eight vibrissae present;

Fig. 3B–C). Sturnira erythromos have nine vi-

brissae surrounding the margins of the noseleaf.

Character 6—Number of vibrissae present on

the upper lip, ventral to the vibrissae that sur-

round the margins of the noseleaf. One vibrissa

present on each side of the face (0); two vibrissae

present (1). In some species of Platyrrhinus (e.g.,

P. aurarius, P. chocoensis, P. dorsalis), Sturnira

erythromos, Uroderma magnirostrum, and Vam-

pyrodes, a single vibrissa is present on the upper

lip, ventral to the vibrissae that surround the mar-

gins of the noseleaf (Fig. 3A). Only P. brachy-

cephalus and Carollia subrufa have two vibrissae

on the upper lip (Fig. 3B). Platyrrhinus helleri

‘‘Eastern,’’ P. infuscus, and P. nigellus present in-

traspeciﬁc variation in this character (one vibrissa

and two vibrissae present) (Fig. 3A–B). This char-

acter has not been used in any other study.

7VELAZCO: PLATYRRHINUS PHYLOGENY

. 3. Schematic view of the heads of three hypo-

thetical bats, showing details of basal protuberance (BP)

and pinna folds (PF). Scale bar

15 mm.

Character 7—Number of submental vibrissae

on each side of the chin. Absence of submental

vibrissae (0); two submental vibrissae present (1);

three submental vibrissae present (2); four sub-

mental vibrissae present (3); ﬁve submental vi-

brissae present (4). The submental vibrissae are

on the chin and lower lip and are usually arranged

in two notable parallel longitudinal rows (Pocock,

1914; Brown, 1971). Some Platyrrhinus (e.g., P.

aurarius, P. lineatus, P. nigellus), Uroderma

magnirostrum, and Vampyrodes (Fig. 4C, E) al-

ways have four submental vibrissae. Platyrrhinus

dorsalis ‘‘Centro-Sur’’ has two submental vibris-

sae (Fig. 4B). Platyrrhinus brachycephalus pre-

sent ﬁve submental vibrissae (Fig. 4D). Intraspe-

ciﬁc variation was noted (three and four submen-

tal vibrissae) in P. chocoensis (Fig. 4A, C, E).

Carollia subrufa lacks submental vibrissae. This

character has not been used in previous studies.

Character 8—Interramal vibrissae. Absence

(0); one interrramal vibrissa present (1); two in-

terramal vibrissae present (2). The interramal vi-

brissae form a median tuft on the gular region

between the two rami of the lower jaws and well

behind the mandibular symphysis (Pocock, 1914;

Brown, 1971). Platyrrhinus reciﬁnus, Sturnira er-

ythromos, and Uroderma magnirostrum lack in-

terramal vibrissae (Fig. 4A). An interramal vibris-

sa is present in some species of Platyrrhinus (e.g.,

P. brachycephalus, P. lineatus) and Vampyrodes

(Fig. 4B–D). Two interramal vibrissae are present

in P. infuscus, P. vittatus ‘‘Northern,’’ P. helleri

‘‘Western,’’ and Carollia subrufa (Fig. 4E). Intra-

speciﬁc variation is found in P. vittatus ‘‘South-

ern’’ (one and two interramal vibrissae present;

Fig. 4B–E). This character was used by Wetterer

et al. (2000: character 13).

Character 9—Inferior border of horseshoe.

Inferior border with no distinct boundary between

horseshoe and lip (0); inferior border partially

joined to the upper lip (1); inferior border com-

pletely free (2). The inferior border of the horse-

shoe is completely free of the upper lip in some

Platyrrhinus (e.g., P. brachycephalus, P. linea-

tus), Uroderma magnirostrum, and Vampyrodes

(Fig. 5C). Platyrrhinus chocoensis, P. dorsalis

‘‘Centro-Sur,’’ and P. infuscus have intraspeciﬁc

variation (inferior border completely free and par-

tially joined (Fig. 5B–C). In Carollia subrufa and

Sturnira erythromos, there is no distinct boundary

between horseshoe and the upper lip (Fig. 5A).

According to Wetterer et al. (2000: character 25),

the inferior border is completely free in all the

species of Platyrrhinus; however, I do not agree

with that assessment.

Character 10—Parallel folds located in the

pinna. Absent (0); poorly marked but distinguish-

able (1); well marked (2). Sturnira erythromos

lacks these folds (Fig. 3C). The folds in the pinna

are not very well deﬁned in some species of Pla-

tyrrhinus (e.g., P. dorsalis, P. lineatus, P. reciﬁ-

nus) and Vampyrodes (Fig. 3B). In contrast, P.

aurarius, P. chocoensis, P. dorsalis ‘‘Centro-

Sur,’’ P. nigellus, P. vittatus ‘‘Northern,’’ Carollia

subrufa, and Uroderma magnirostrum have well-

8 FIELDIANA: ZOOLOGY

. 4. Ventral view of the chin of ﬁve hypothetical bats, showing states of interramal(IV) and submental vibrissae

(SV). Scale bar

5 mm.

marked pinna folds (Fig. 3A). Platyrrhinus helleri

‘‘Eastern’’ and P. infuscus present intraspeciﬁc

variation, presenting both states (Fig. 3A–B). This

is the ﬁrst use of this character in a phylogenetic

analysis.

Character 11—Dorsal stripe. Absent (0); in-

distinct, thin, and obscured (1); deﬁnite but nar-

row (2); brilliant-white and wide (3). A dorsal

midline stripe is absent in Carollia subrufa and

Sturnira erythromos. A narrow dorsal stripe is

presented in some species of Platyrrhinus (e.g.,

P. chocoensis, P. dorsalis) and Uroderma mag-

nirostrum. Platyrrhinus lineatus, P. reciﬁnus, P.

vittatus ‘‘Northern,’’ P. vittatus ‘‘Southern,’’ and

Vampyrodes have the dorsal stripe that is white

and wide. In Platyrrhinus infuscus, the dorsal

stripe is thin and obscure. Platyrrhinus brachy-

cephalus, P. helleri ‘‘Eastern,’’ and P. nigellus all

show intraspeciﬁc variation in this character,some

individuals with a narrow dorsal stripe and others

with white and wide stripes. Owen (1987: char-

acter 4) used the dorsal stripe as part of a complex

multistate character termed ‘‘Pelage patterns,’’ in

which he included facial stripes, dorsal stripe, and

shoulder patches. I agree with Owen (1987) on

the scores for dorsal stripe in the species used in

the present analysis. Wetterer et al. (2000: char-

acter 7) used the dorsal stripe as a character with

three states: absent, present, and occasionally pre-

sent.

9VELAZCO: PLATYRRHINUS PHYLOGENY

. 5. Frontal view of the noseleaf in (A)Sturnira erythromos (MUSM 5310); (B.)Platyrrhinus infuscus (MUSM

9897); (C)P. nigellus (MUSM 16170). Scale bar

5 mm.

Character 12—Length of the dorsal fur.

Long,

8 mm (0); medium, average, 6.30–7.50

mm (1); short,

6.30 mm (2). Dorsal fur is of

average length in some species of Platyrrhinus

(e.g., P. helleri ‘‘Eastern,’’ P. reciﬁnus), Carollia

subrufa, Uroderma magnirostrum, and Vampyro-

des. In Platyrrhinus aurarius, P. dorsalis, P. dor-

salis ‘‘Norte,’’ P. nigellus, P. vittatus ‘‘Northern,’’

P. vittatus ‘‘Southern,’’ and Sturnira erythromos,

the dorsal fur is long. Platyrrhinus infuscus char-

acteristically has short dorsal fur. Platyrrhinus

chocoensis presents intraspeciﬁc variation, in

which some individuals have short dorsal fur,

while in others it is normal. This character was

used by Marques-Aguiar (1994: character 3) in

her study of the genus Artibeus.

Character 13—Bands of contrast in the dor-

sal fur. Bicolored (0); tricolored (1); tetracolored

(2). In most Platyrrhinus (e.g., P. brachycephalus,

P. chocoensis, P. dorsalis), Carollia subrufa, and

Uroderma magnirostrum, the dorsal fur is dis-

tinctly tricolored. In contrast, tetracolored dorsal

fur is presented in P. aurarius, P. reciﬁnus, and

Sturnira erythromos. Vampyrodes has bicolored

dorsal fur. Owen (1987: character 3) and Wetterer

et al. (2000: character 5) scored S. erythromos as

having tricolored dorsal fur; according to my ob-

servations, it has tetracolored dorsal fur. Wetterer

et al. (2000) scored all Platyrrhinus as having bi-

colored dorsal fur, as did Owen (1987) with the

exception of P. dorsalis which had tricolored dor-

sal fur. I agree with Owen’s score for P. dorsalis

but do not agree with the scores of the other Pla-

tyrrhinus species. Instead, I observed interspeciﬁc

variation in dorsal fur patterns among the species

of Platyrrhinus. I agree with Owen (1987) and

Wetterer et al. (2000) on the score of Vampyrodes.

Character 14—Bands of contrast in the ven-

tral fur. Unicolored (0); bicolored (1); tricolored

(2); tetracolored (3). A tricolored ventral fur is

present in some species of Platyrrhinus (e.g., P.

brachycephalus, P. infuscus), Carollia subrufa,

and Vampyrodes. In contrast, P. chocoensis, P

helleri ‘‘Eastern,’’ P. vittatus ‘‘Northern,’’ P. vit-

tatus ‘‘Southern,’’ and Uroderma magnirostrum

have a bicolored ventral fur. Platyrrhinus helleri

‘‘Western’’ presents intraspeciﬁc variation be-

cause some individuals present a bicolored ventral

fur and others a unicolored ventral fur. Sturnira

erythromos has tetracolored ventral fur. This char-

acter has not been used in any other study.

Character 15—Hair on the dorsum of the

feet. Sparse and short (0); intermediate in density

and length (1); dense and long (2). Dense and

long hair is present on the dorsal side of the feet

in some Platyrrhinus (e.g., P. lineatus, P. vittatus

‘‘Southern’’) and Sturnira erythromos. In Platyr-

rhinus aurarius, P. chocoensis, P. dorsalis, P.

helleri ‘‘Eastern,’’ and Vampyrodes, the hair on

the dorsal side of the feet is intermediate in den-

sity and length. Only P. infuscus, P. helleri

‘‘Western,’’ and Uroderma magnirostrum have

sparse and short hairs on the dorsal side of the

feet. Platyrrhinus brachycephalus presents intra-

speciﬁc variation in this character, some individ-

uals with sparse hairs, and others with an inter-

mediate distribution of hair on the dorsal side of

the feet. This character has not been used in any

other study.

Character 16—Posterior edge of the uropa-

tagium. ‘‘V’’ shaped (0); ‘‘U’’ shaped (1). The

uropatagium is ‘‘U’’ shaped in some Platyrrhinus

(e.g., P. brachycephalus, P. reciﬁnus), Carollia

subrufa, and Sturnira erythromos. In contrast,

10 FIELDIANA: ZOOLOGY

. 6. Ventral view of thebasicranium, showing de-

tails of the posterior margin of the hard palate: (A)Pla-

tyrrhinus brachycephalus (MUSM 13793); (B)P. helleri

‘‘Western’’ (MUSM 10725). Scale bar

8 mm.

Vampyrodes has ‘‘V’’-shaped uropatagium. Pla-

tyrrhinus dorsalis, P. helleri ‘‘Eastern,’’ P. helleri

‘‘Western,’’ P. infuscus, P. nigellus, P. vittatus

‘‘Northern,’’ and Uroderma magnirostrum have

intraspeciﬁc variation. This character was evalu-

ated only in alcohol-preserved specimens. This

character has not been used in any other study.

Character 17—Distribution of the fringe of

hair on the edge of the uropatagium. Absent

(0); usually hairy, occasionally sparsely (1);

densely haired (2). A densely haired uropatagium

fringe is present in some Platyrrhinus (e.g., P.

helleri ‘‘Eastern,’’ P. nigellus) and Sturnira ery-

thromos. In contrast, P. brachycephalus, P. cho-

coensis, P. dorsalis, P. infuscus, P. vittatus

‘‘Northern,’’ and Vampyrodes have a regular or

occasionally sparsely haired uropatagium fringe.

Carollia subrufa and Uroderma magnirostrum

lack this fringe of hair on the edge of the uropa-

tagium. I agree with Lim (1993: character 13) and

Wetterer et al. (2000: character 10) on the score

of Platyrrhinus, Carollia, Sturnira, Uroderma,

and Vampyrodes.

Skull

Character 18—Posterior border of the hard

palate. ‘‘U’’ shaped (0); ‘‘V’’ shaped (1). A ‘‘U’’-

shaped posterior border of the hard palate is pre-

sent in some species of Platyrrhinus (e.g., P. li-

neatus, P. reciﬁnus), Carollia subrufa, Sturnira

erythromos, and Uroderma magnirostrum (Fig.

6A). In contrast, P. aurarius, P. dorsalis, and P.

dorsalis ‘‘Norte’’ have a ‘‘V’’-shaped posterior

border of the hard palate (Fig. 6B). Platyrrhinus

brachycephalus, P. chocoensis, P. dorsalis ‘‘Cen-

tro-Sur,’’ P. helleri ‘‘Eastern,’’ P. helleri ‘‘West-

ern,’’ P. infuscus, P. nigellus, P. vittatus ‘‘North-

ern,’’ P. vittatus ‘‘Southern,’’ and Vampyrodes

have intraspeciﬁc variation for this character, with

some individuals possessing the ‘‘U’’-shaped pos-

terior border of the hard palate and other individ-

uals ‘‘V’’ shaped (Fig. 6A–B). Wetterer et al.

(2000: character 45) used this character, but they

scored Platyrrhinus, Vampyrodes, Uroderma, and

Sturnira as ‘‘—’’ because they considered that

there was a continuous range of intra- and inter-

speciﬁc variation among the shape of the posterior

border, making it not a useful character. Accord-

ing to my observations, I found some species with

intraspeciﬁc variation. Owen (1987: character 10)

used this character, and I do not agree with his

scores of Vampyrodes; he indicated that Vampy-

rodes only has a ‘‘U’’-shaped posterior border of

the hard palate, but I also found some individuals

having a ‘‘V’’-shaped posterior border of the hard

palate. Within Platyrrhinus, he did not ﬁnd intra-

speciﬁc variation, but according to my observa-

tions, Platyrrhinus demonstrates intraspeciﬁc var-

iation. Lim (1993: character 4) scored Platyrrhi-

nus, Vampyrodes, and Sturnira as having a hard

palate extended into interpterygoidal space, but he

did not score the shape of the posterior border of

11VELAZCO: PLATYRRHINUS PHYLOGENY

. 7. Dorsal view of two crania, showing details of the postorbital process (PP): (A)Platyrrhinus nigellus

(MUSM 16170); (B)P. infuscus (MUSM 9897). Scale bar

12 mm.

. 8. Lateral view of the temporal region, showing

details of the mastoid process (MP) and paraoccipital

process (PoP): (A)Platyrrhinus helleri ‘‘Western’’

(MUSM 10726); (B)P. nigellus (MUSM 16170); (C)P.

vittatus ‘‘Southern’’ (MUSM 14584); (D)Sturnira ery-

thromos (MUSM 5260). Scale bar

2 mm.

the hard palate in these genera. This character was

used by Marques-Aguiar (1994: character 21) and

Owen (1991: character 23) in their studies of the

systematics of the genus Artibeus and Dermanu-

ra, respectively.

Character 19—Postorbital process. Absent or

poorly developed (0); moderately developed (1);

well developed (2). A poorly developed postor-

bital process is present in some species of Platyr-

rhinus (e.g., P. chocoensis, P. dorsalis) and Stur-

nira erythromos (Fig. 7A). In Platyrrhinus aurar-

ius, P. helleri ‘‘Eastern,’’ P. infuscus, P. reciﬁnus,

P. vittatus ‘‘Northern,’’ P. vittatus ‘‘Southern,’’

and Vampyrodes, the postorbital process is mod-

erately developed (Fig. 7B). Platyrrhinus lineatus,

Carollia subrufa, and Uroderma magnirostrum

have a well-developed postorbital process. In Pla-

tyrrhinus dorsalis ‘‘Norte,’’ there is intraspeciﬁc

variation, with individuals having a poorly devel-

oped or moderately developed postorbital process

(Fig. 7A–B). This character was used by

Marques-Aguiar (1994: character 15) in her study

of the genus Artibeus.

Character 20—Paraoccipital process. Poorly

developed, almost imperceptible (0); moderately

developed (1); well developed (2). The paraoccip-

ital process appears moderately developed, about

half the size of the mastoid process, in some spe-

cies of Platyrrhinus (e.g., P. helleri ‘‘Eastern,’’ P.

nigellus), Carollia subrufa, Sturnira erythromos,

and Vampyrodes (Fig. 8B). In Platyrrhinus au-

rarius, P. infuscus, P. lineatus, P. vittatus ‘‘South-

ern,’’ and Uroderma magnirostrum, the paraoc-

cipital process is well developed, almost the same

size as the mastoid process (Fig. 8C–D). Platyr-

rhinus brachycephalus and P. helleri ‘‘Western’’

have the paraoccipital process poorly developed,

12 FIELDIANA: ZOOLOGY

. 9. Posterior view of the temporal region, show-

ing details of zygomatic arch (ZA) and fossa on the

squamosal end of the zygomatic arch: (A)Platyrrhinus

brachycephalus (MUSM 13793); (B)P. infuscus

(MUSM 9897); (C)P. dorsalis (FMNH 128141). Scale

bar

4 mm.

scarcely a third of the size of the mastoid process

(Fig. 8A). Intraspeciﬁc variation is present in P.

chocoensis, with the paraoccipital process poorly

to moderately developed (Fig. 8A–B). I do not

agree with Owen’s (1987: character 12) score for

P. brachycephalus, P. helleri, and P. nigellus;he

scored them as if they lacked a paraoccipital pro-

cess. Owen (1987) also scored P. dorsalis and P.

infuscus as having minimal development of the

paraoccipital process, but according to personal

observations, P. dorsalis has a moderately devel-

oped process (Fig. 8B), and P. infuscus has a

well-developed paraoccipital process (Fig. 8C).

This character has been used by Marques-Aguiar

(1994: character 20), Straney (1980: character J-

6), and Owen (1987: character 12) in their phy-

logenetic analyses of Artibeus, Phyllostominae,

and Dermanura, respectively.

Character 21—Apophysis in the paraoccipi-

tal process. Present (0); absent (1). All species of

Platyrrhinus, Carollia subrufa, and Vampyrodes

caraccioli lack an apophysis in the paraoccipital

process (Fig. 8A–C). In contrast, Sturnira ery-

thromos and Uroderma magnirostrum have an

apophysis in the paraoccipital process (Fig. 8D).

This character has not been used in any other

study.

Character 22—Fossa on the squamosal end

of the zygomatic arch, lateral to the glenoid

fossa. Absent or almost imperceptible (0); shallow

(1); deep (2). This fossa is absent or almost im-

perceptible in some species of Platyrrhinus (e.g.,

P. helleri ‘‘Eastern,’’ P. lineatus), Carollia sub-

rufa, Sturnira erythromos, Uroderma magniros-

trum, and Vampyrodes (Fig. 9A). Platyrrhinus

dorsalis ‘‘Norte,’’ P. infuscus, and P. reciﬁnus

have a shallow fossa (Fig. 9B). Platyrrhinus au-

rarius and P. dorsalis have a deep fossa (Fig. 9C).

Platyrrhinus chocoensis and P. dorsalis ‘‘Centro-

Sur’’ present intraspeciﬁc variation, some individ-

uals have the fossa on the squamosal end of the

zygomatic arch absent or almost imperceptible,

and others have this fossa shallow. Platyrrhinus

nigellus also demonstrate intraspeciﬁc variation,

with a shallow or deep fossa. This character has

not been used in any other study.

Character 23—Posterior clinoid processes.

Absent (0); present (1). This process is located on

the posterior face of the sella turcica in the sphe-

noid bone (Stromsten, 1947). Carollia subrufa,

Sturnira erythromos, and Uroderma magniros-

trum lack these processes (Fig. 10A). In contrast,

all the species of Platyrrhinus and Vampyrodes

possess posterior clinoid processes (Fig. 10B).

Dentition

Character 24—Upper median incisors. In

contact (0); without contact (1). The upper median

incisors are in contact in some species of Platyr-

rhinus (e.g., P. chocoensis, P. lineatus), Carollia

subrufa, Sturnira erythromos, and Vampyrodes. In

contrast, Uroderma magnirostrum has the upper

median incisors not in contact. Platyrrhinus bra-

chycephalus, P. helleri ‘‘Eastern,’’ P. helleri

‘‘Western,’’ P. infuscus, and P. reciﬁnus have in-

traspeciﬁc variation.

Character 25—Lobulation of the upper lat-

eral incisors. Monolobed (0); bilobed (1). Mon-

olobed upper lateral incisors are present in some

species of Platyrrhinus (e.g., P. chocoensis, P.

helleri ‘‘Eastern’’), Carollia subrufa, and Vam-

pyrodes. In contrast, P. dorsalis ‘‘Centro-Sur’’

13VELAZCO: PLATYRRHINUS PHYLOGENY

. 10. Detail of the posterior clinoid processes

(PCP), in a posterior view of cranium: (A) Absence of

PCP in Uroderma magnirostrum (MUSM 14029); (B)

Presence of the PCP in Platyrrhinus infuscus (MUSM

9897). Scale bar

8 mm.

. 11. Occlusal view of the maxillary toothrow:

(A)Sturnira erythromos (MUSM 5310); (B)Platyrrhin-

us infuscus (MUSM 9897). Scale bar

8 mm.

and Uroderma magnirostrum have bilobed upper

lateral incisors. Platyrrhinus brachycephalus, P.

dorsalis ‘‘Norte,’’ P. helleri ‘‘Western,’’ P. infus-

cus, P. lineatus, P. vittatus ‘‘Southern,’’ and Stur-

nira erythromos have intraspeciﬁc variation.

Character 26—Alignment of P3 in the tooth-

row. P3 aligned along the axis of the toothrow

(0); P3 aligned transversally to the toothrow (1).

Carollia subrufa and Sturnira erythromos have

the P3 aligned along the axis of the toothrow (Fig.

11A). In contrast, all the species of Platyrrhinus,

Uroderma magnirostrum, and Vampyrodes have

the P3 aligned transversally to the toothrow (Fig.

11B). This character has not been used in any

other study.

Character 27—Size of the P3 with regard to

P4. P3 slightly smaller than P4 (0); P3 about half

the size of P4 (1). All the species of Platyrrhinus,

Uroderma magnirostrum, and Vampyrodes pos-

sess a P3 slightly bigger than half the size of P4.

Carollia subrufa and Sturnira erythromos have a

P3 almost the same size as P4. This character has

not been used in any other study.

Character 28—Stylar cusps on the posterior

face of P4. P4 lacking stylar cusps or with one

stylar cusp present (0); two stylar cusps present

(1). All the species of Platyrrhinus, Uroderma

magnirostrum, and Vampyrodes have two stylar

cusps on the posterior face of P4 (Fig. 12). In

contrast, Carollia subrufa and Sturnira erythro-

mos do not possess two stylar cusps on P4, al-

though occasionally they exhibit one stylar cusp.

I agree with Lim (1993: character 6) in his scores

of Platyrrhinus species. I do not agree with Wet-

terer et al. (2000: character 57) on their scores for

Platyrrhinus because they indicated that this ge-

nus only occasionally has the second stylar cusp.

According to my observations, all the individuals

of Platyrrhinus always have two stylar cusps.

They also stated that Lim (1993: character 6) did

14 FIELDIANA: ZOOLOGY

. 12. Lateral view of the maxillary toothrow:

Platyrrhinus infuscus (MUSM 9897). Scale bar

8 mm.

not report the presence of the second stylar cusp

for Vampyrodes, when in fact he did report it.

Character 29—Sulcus on the posterior face of

P4. Absent (0); present (1). All the species of Pla-

tyrrhinus, Uroderma magnirostrum, and Vampy-

rodes have a sulcus on the posterior face of P4,

starting near the accessory cusps and ending at

the tip of P4. In contrast, Carollia subrufa and

Sturnira erythromos lack this sulcus. This char-

acter has not been used in any previous study.

Character 30—Sulcus on the anterior face of

P4. Absent (0); present (1). All the species of Pla-

tyrrhinus have a sulcus on the anterior face of P4,

starting near the cingulum and ending at the tip

of P4. In contrast, Carollia subrufa, Sturnira er-

ythromos, Uroderma magnirostrum, and Vampy-

rodes lack this sulcus on the anterior face of P4.

This character has not been used in any previous

study.

Character 31—Fossa on the hypoconal basin

of P4. Shallow (0); deep (1). The fossa of the

hypoconal basin of P4 is deep in some species of

Platyrrhinus (e.g., P. reciﬁnus, P. vittatus ‘‘South-

ern’’). In contrast, P. aurarius, P. chocoensis, P.

helleri ‘‘Eastern,’’ P. lineatus, Carollia subrufa,

Sturnira erythromos, and Uroderma magniros-

trum have a shallow fossa. Platyrrhinus brachy-

cephalus, P. dorsalis, P. dorsalis ‘‘Norte,’’ P.

dorsalis ‘‘Centro-Sur,’’ P. helleri ‘‘Western,’’ P.

nigellus, and Vampyrodes have intraspeciﬁc var-

iation. This character has not been used by other

researchers.

Character 32—M1 Parastyle. Absent (0); pre-

sent (1). The M1 parastyle is present in some spe-

cies of Platyrrhinus (e.g., P. helleri ‘‘Eastern,’’ P.

reciﬁnus). In contrast, other species of Playtrrhin-

us (e.g., P. aurarius, P. chocoensis), Carollia sub-

rufa, Sturnira erythromos, Uroderma magniros-

trum, and Vampyrodes lack the parastyle on M1.

Platyrrhinus lineatus has intraspeciﬁc variation.

Straney (1980: character: K 27) used this char-

acter in his phylogeny of the Phyllostominae.

Character 33—M1 Mesostyle. Absent (0); pre-

sent (1). The mesostyle on M1 is absent in some

species of Platyrrhinus (e.g., P. chocoensis, P.

dorsalis), Carollia subrufa, and Sturnira erythro-

mos. In contrast, P. vittatus ‘‘Southern,’’ Urod-

erma magnirostrum, and Vampyrodes possess a

mesostyle on M1. Platyrrhinus brachycephalus,

P. helleri ‘‘Eastern,’’ P. helleri ‘‘Western,’’ P. in-

fuscus, P. dorsalis ‘‘Norte,’’ and P. nigellus have

intraspeciﬁc variation.

Character 34—Labial cingulum at the base

of the M1 metacone. Absent (0); present (1); sty-

lar cusp on the labial cingulum of the M1 meta-

cone (2). Carollia subrufa, Sturnira erythromos,

and Uroderma magnirostrum lack the labial cin-

gulum on the base of the M1 metacone. Platyr-

rhinus brachycephalus, P. helleri ‘‘Eastern,’’ P.

helleri ‘‘Western,’’ P. reciﬁnus, and Uroderma

magnirostrum have the labial cingulum without a

stylar cusp. In contrast, a stylar cusp on the labial

cingulum of the M1 metacone is present in some

species of Platyrrhinus (e.g., P. chocoensis, P. in-

fuscus) and Vampyrodes. Platyrrhinus lineatus

and P. nigellus have intraspeciﬁc variation, as

some individuals have a stylar cusp on the labial

cingulum of the M1 metacone and others lack this

stylar cusp. This character has not been used by

other researchers.

Character 35—Sulcus on the posterior face of

the M1 paracone. Absent (0); present (1). All the

species of Platyrrhinus, Uroderma magnirostrum,

and Vampyrodes have a sulcus on the posterior

face of the M1 paracone. In contrast, Carollia

subrufa and Sturnira erythromos lack this sulcus

on M1.

Character 36—Lingual cingulum on the base

of the M1 metacone. Absent (0); lingual cingu-

lum present at the base of the M1 metacone (1);

stylar cusp on the lingual cingulum of the M1

metacone (2). Carollia subrufa, Sturnira erythro-

mos, and Vampyrodes lack the lingual cingulum

on the M1 metacone. A stylar cusp is present on

the cingulum of the lingual face of the M1 meta-

cone in some species of Platyrrhinus (e.g., P. in-

fuscus, P. lineatus). In contrast, P. helleri ‘‘East-

ern,’’ P. helleri ‘‘Western,’’ P. reciﬁnus, and

Uroderma magnirostrum lack this stylar cusp.

Platyrrhinus brachycephalus, P. chocoensis, P.

dorsalis, and P. nigellus have intraspeciﬁc varia-

tion, as some individuals have a stylar cusp on the

lingual cingulum of the M1 metacone and others

15VELAZCO: PLATYRRHINUS PHYLOGENY

. 13. Occlusal view of M1–M2: (A)Platyrrhinus

infuscus (MUSM 9897); (B)P. dorsalis ‘‘Norte’’

(MUSM 4946). Scale bar

2.5 mm.

lack this stylar cusp. This is the ﬁrst use of this

character in a phylogenetic analysis.

Character 37—Sulcus on the posterior face of

the paracone not joined to the cingulum of the

lingual face of the metacone on M1. Absent (0);

present (1). The majority of the Platyrrhinus spe-

cies (e.g., P. aurarius, P. brachycephalus, P. cho-

coensis) have this sulcus on the posterior face of

the paracone joined to the cingulum of the lingual

face of the metacone on M1 (Fig. 13A). In con-

trast, P. dorsalis ‘‘Norte’’ and Uroderma magni-

rostrum have the sulcus on the posterior face of

the paracone not joined to the cingulum of the

lingual face of the M1 metacone (Fig. 13B). Car-

ollia subrufa, Sturnira erythromos, and Vampy-

rodes caraccioli were scored as ‘‘—’’ because

they lack the cingulum on the lingual face of the

M1 metacone. This character has not been used

by other researchers.

Character 38—M1 metastyle. Absent (0); pre-

sent (1). The metastyle on M1 is present in the

majority of Platyrrhinus species (e.g., P. dorsalis,

P. dorsalis ‘‘Norte’’), Carollia subrufa, and Vam-

pyrodes. In contrast, Sturnira erythromos and

Uroderma magnirostrum lack the metastyle on

M1. Platyrrhinus brachycephalus, P. chocoensis,

P. helleri ‘‘Eastern,’’ and P. helleri ‘‘Western’’

have intraspeciﬁc variation.

Character 39—M1 protocone. Well-devel-

oped, large (0); moderately developed (1); small

and blunt (2). A well-developed M1 protocone is

present in Platyrrhinus brachycephalus. In con-

trast, P. chocoensis, P. dorsalis, P. dorsalis

‘‘Norte,’’ and P. helleri ‘‘Eastern’’ possess an M1

protocone that is small and blunt. Some species

of Platyrrhinus (e.g., P. aurarius, P. infuscus),

Carollia subrufa, Sturnira erythromos, Uroderma

magnirostrum, and Vampyrodes have a moderate-

ly developed protocone on M1. Platyrrhinus vit-

tatus ‘‘Northern’’ has intraspeciﬁc variation: some

individuals have the protocone with a moderate

development and others with a well-developed

protocone on M1. This character has not been

used by other researchers.

Character 40—M2 parastyle. Absent (0);

present (1). The parastyle on M2 is present in all

Platyrrhinus species and Vampyrodes. In contrast,

Carollia subrufa, Sturnira erythromos, and Uro-

derma magnirostrum lack a parastyle on M2. This

character has not been used in any other study.

Character 41—Labial cingulum of the M2

paracone. Absent (0); present (1). The labial cin-

gulum on the M2 paracone is present in some spe-

cies of Platyrrhinus (e.g., P. brachycephalus, P.

chocoensis) and Vampyrodes. In contrast, Carol-

lia subrufa, Sturnira erythromos, and Uroderma

magnirostrum lack the labial cingulum on the M2

paracone. Platyrrhinus dorsalis, P. dorsalis

‘‘Norte,’’ and P. dorsalis ‘‘Centro-Sur’’ have in-

traspeciﬁc variation. This character has not been

used in any other study.

Character 42—Stylar cusp on the lingual face

of the M2 paracone. Absent (0); present (1). A

stylar cusp on the lingual face of the M2 paracone

is present in some species of Platyrrhinus (e.g.,

P. infuscus, P. vittatus ‘‘Southern’’). In contrast,

other species of Platyrrhinus (e.g., P. aurarius, P.

brachycephalus), Carollia subrufa, Sturnira ery-

thromos, Uroderma magnirostrum, and Vampy-

rodes lack this stylar cusp on the lingual face of

the M2 paracone. Platyrrhinus chocoensis and P.

dorsalis have intraspeciﬁc variation. This char-

acter has not been used in any other study.

Character 43—M2 metastyle. Absent (0); pre-

sent (1). A metastyle on M2 is present in some

species of Platyrrhinus (e.g., P. brachycephalus,

P. chocoensis) and Uroderma magnirostrum. In

contrast, P. vittatus ‘‘Southern,’’ Carollia subrufa,

Sturnira erythromos, and Vampyrodes lack a me-

tastyle on M2. Platyrrhinus nigellus has intraspe-

16 FIELDIANA: ZOOLOGY

ciﬁc variation. This character has not been used

in any other study.

Character 44—Stylar cusp on the lingual face

of the M2 metacone. Absent (0); present (1). A

stylar cusp on the lingual face of the M2 meta-

cone is present in some species of Platyrrhinus

(e.g., P. infuscus, P. vittatus ‘‘Southern’’) and

Vampyrodes. In contrast, other species of Platyr-

rhinus (e.g., P. aurarius, P. chocoensis), Carollia

subrufa, Sturnira erythromos, and Uroderma

magnirostrum lack a stylar cusp on the lingual

face of the M2 metacone. Platyrrhinus dorsalis

‘‘Norte,’’ P. dorsalis ‘‘Centro-Sur,’’ and P. linea-

tus have intraspeciﬁc variation. This character has

not been used in any other study.

Character 45—Lingual cingulum of the M2

metacone. Absent (0); lingual cingulum of the

M2 metacone restricted to the metacone (1); lin-

gual cingulum of the M2 metacone continuous to

the paracone (2). Carollia subrufa, Sturnira ery-

thromos, and Vampyrodes lack the lingual cin-

gulum of the M2 metacone. The lingual cingulum

of the M2 metacone is continuous to the paracone

in Platyrrhinus chocoensis, P. dorsalis, and P. re-

ciﬁnus (Fig. 13A). In contrast, the lingual cingu-

lum of the M2 metacone is restricted to the meta-

cone in some species of Platyrrhinus (e.g., P. bra-

chycephalus, P. vittatus ‘‘Southern’’) and Uro-

derma magnirostrum (Fig. 13B). Platyrrhinus

helleri ‘‘Western,’’ P. infuscus, P. lineatus, and P.

nigellus have intraspeciﬁc variation in this char-

acter: some individuals have the lingual cingulum

of the M2 metacone restricted to the metacone,

whereas others have lingual cingulum of the M2

metacone continuous to the paracone. This char-

acter has not been used by other researchers.

Character 46—M2 hypoconal basin. Devel-

oped (0); rudimentary or small (1). A developed

hypoconal basin on M2 is present in all the spe-

cies of Platyrrhinus and Sturnira erythromos. In

contrast, in Carollia subrufa, Uroderma magni-

rostrum, and Vampyrodes, the M2 hypoconal ba-

sin is rudimentary or small.

Character 47—Two sulci on the lingual face

of the main cone of p4. Absent (0); present (1).

Two sulci on the lingual face of the main cone of

p4 are present in all the Platyrrhinus species,

Uroderma magnirostrum, and Vampyrodes. These

sulci originate at the base of the main cone and

continue to the tip of the main cone. In contrast,

Carollia subrufa and Sturnira erythromos lack

these sulci on p4. This character has not been used

in any other study.

Character 48—Cones on the posterior face of

p4 (one lingual and the other one labial). Two

cones present (0); one cone present on either the

labial or the lingual face (1); no cones present (2).

Two cones on the posterior face of p4 are present

in all the species of Platyrrhinus and Vampyrodes.

Sturnira erythromos and Uroderma magnirostrum

have only one cone. Carollia subrufa lack both

cones on the posterior face of p4. This character

has not been used by other researchers.

Character 49—Labial and lingual cingulum

of p4. Both cingula absent (0); only one cingulum

present, either the lingual or the labial (1); both

cingula present (2). Both cingula on p4 are pre-

sent in the majority of Platyrrhinus species (e.g.,

P. dorsalis, P. vittatus ‘‘Southern’’). In Platyr-

rhinus aurarius, P. chocoensis, Sturnira erythro-

mos, Uroderma magnirostrum, and Vampyrodes,

only one cingulum is present on p4. Carollia sub-

rufa lack cingula on p4. This character has not

been used by other researchers.

Character 50—Stylid cusps on the anterior

or anterolingual face of the main cone of p4.

Absent (0); one stylid cusp present (1); two stylid

cusps present (2). Two stylid cusps on the anterior

face of the main cone of p4 are present in Pla-

tyrrhinus brachycephalus. In Platyrrhinus vittatus

‘‘Northern’’ and Vampyrodes, one and two stylid

cusps, respectively, are present on the anterolin-

gual face. Platyrrhinus helleri ‘‘Eastern’’ has a

single stylar cusp on the anterior face of p4. In

contrast, other Platyrrhinus species (e.g., P. au-

rarius, P. chocoensis), Carollia subrufa, Sturnira

erythromos, and Uroderma magnirostrum lack

stylid cusps on the anterior face of the main cone

of p4. This is the ﬁrst use of this character in a

phylogenetic study.

Character 51—Stylid cusp on the posterior

face of the main cone of p4. Absent (0); present

(1). One or two stylid cusps are present on the

posterior face of the main cone of p4 in some

species of Platyrrhinus (e.g., P. helleri ‘‘Eastern,’’

P. helleri ‘‘Western’’). In contrast, other species

of Platyrrhinus (e.g., P. aurarius, P. chocoensis),

Carollia subrufa, Sturnira erythromos, Uroderma

magnirostrum, and Vampyrodes lack stylid cusps

on the posterior face of the main cone of p4. This

character has not been used by other researchers.

Character 52—m1 paraconid. Present (0); ab-

sent (1). Most Platyrrhinus species (e.g., P. au-

rarius, P. chocoensis), Carollia subrufa, Uroder-

ma magnirostrum, and Vampyrodes lack a para-

conid on m1. In contrast, Sturnira erythromos has

a paraconid on m1. Platyrrhinus nigellus has in-

traspeciﬁc variation. Wetterer et al. (2000: char-

17VELAZCO: PLATYRRHINUS PHYLOGENY

acter 68) used this character in their phylogeny of

the Phyllostomidae, but I do not agree on their

score for Platyrrhinus. They scored it as a poly-

morphism (with the paraconid occasionally pre-

sent). However, my observations indicate that the

majority of Platyrrhinus lack the paraconid on

m1, as does Vampyrodes.

Character 53—m1 cingulum. Absent (0); only

the labial cingulum present (1); both labial and

lingual cingula present (2). Both cingula on m1

are present in all the species of Platyrrhinus. In

contrast, Carollia subrufa and Sturnira erythro-

mos lack both cingula on m1. Uroderma magni-

rostrum and Vampyrodes have only the labial cin-

gulum. This character has not been used in any

other study.

Character 54—Stylid cusps on the anterior

face of the m1 protoconid. Absent (0); present

(1). A stylid cusp on the anterior face of the m1

protoconid is present in all species of Platyrrhin-

us. In contrast, Carollia subrufa, Sturnira ery-

thromos, Uroderma magnirostrum, and Vampy-

rodes lack this stylid cusp on the anterior face of

the m1 protoconid. This character has not been

used in any other study.

Character 55—m1 metaconid. Well developed

(0); poorly developed (1); absent (2). A well-de-

veloped m1 metaconid is present in Platyrrhinus

infuscus, P. lineatus, and Sturnira erythromos.

However, other species of Platyrrhinus (e.g., P.

aurarius, P. reciﬁnus), Uroderma magnirostrum,

and Vampyrodes lack a metaconid on m1. Platyr-

rhinus dorsalis ‘‘Norte,’’ P. vittatus ‘‘Northern,’’

P. vittatus ‘‘Southern,’’ and Carollia subrufa have

a poorly developed m1 metaconid. Platyrrhinus

nigellus has intraspeciﬁc variation, with some

specimens lacking the m1 metaconid and others

with it poorly developed. I do not agree with the

score of present that Wetterer et al. (2000: char-

acter 69) assigned to Vampyrodes because, ac-

cording to my observations, Vampyrodes lacks a

metaconid on m1.

Character 56—m2 hypoconid. Present (0); ab-

sent (1). The m2 hypoconid is present in Carollia

subrufa and Sturnira erythromos. In contrast, the

majority of the species of Platyrrhinus (e.g., P.

brachycephalus, P. nigellus), Uroderma magni-

rostrum, and Vampyrodes lack the hypoconid on

m2. Platyrrhinus helleri ‘‘Eastern’’ has intraspe-

ciﬁc variation. This character has not been used

in other studies.

Character 57—Stylid cusp between the meta-

conid and protoconid on m2. Absent (0); present

(1). A stylid cusp between the metaconid and pro-

toconid on m2 is present in some species of Pla-

tyrrhinus (e.g., P. brachycephalus, P. vittatus

‘‘Southern’’) and Vampyrodes. In contrast, P. cho-

coensis, P. infuscus, P. reciﬁnus, Carollia subru-

fa, Sturnira erythromos, and Uroderma magniros-

trum lack this stylid cusp between the metaconid

and protoconid on m2. This character has not

been used in any other study.

Character 58—Cingula on m2. Both cingula

labial and lingual absent (0); labial cingulum pre-

sent (1); both cingula present (2). Both cingula on

m2 are present in the majority of species of Pla-

tyrrhinus (e.g., P. brachycephalus, P. infuscus)

and Uroderma magnirostrum. In contrast, Vam-

pyrodes has only the labial cingulum, and Car-

ollia subrufa and Sturnira erythromos lack both

cingula on m2. Platyrrhinus helleri ‘‘Western’’

has intraspeciﬁc variation, with some individuals

with both cingula and others with only the labial

cingulum. This character has not been used by

other researchers.

Postcranial Skeleton

Character 59—Third and ﬁfth metacarpal

arrangement. Length of M III

M V (0); length

of M III

M V (1); length of M III

M V (2).

Platyrrhinus aurarius, P. dorsalis ‘‘Norte,’’ P. li-

neatus, P. nigellus, P. vittatus ‘‘Northern,’’ Stur-

nira erythromos, and Vampyrodes have the third

metacarpal shorter that the ﬁfth metacarpal. In

Platyrrhinus helleri ‘‘Eastern,’’ P. helleri ‘‘West-

ern,’’ P. reciﬁnus, and Uroderma magnirostrum,

the third metacarpal is longer than the ﬁfth meta-

carpal. The third metacarpal is subequal to the

ﬁfth in P. vittatus ‘‘Southern.’’ Platyrrhinus cho-

coensis, P. dorsalis, P. dorsalis ‘‘Centro-Sur,’’

and P. infuscus exhibit intraspeciﬁc variation: the

third metacarpal can be subequal to or shorter

than the ﬁfth metacarpal. Platyrrhinus brachyce-

phalus and Carollia subrufa also have intraspe-

ciﬁc variation: the third metacarpal can be sube-

qual to or longer than the ﬁfth metacarpal. Wet-

terer et al. (2000: character 84) used this character

in their phylogeny of the Phyllostomidae; they in-

dicated that in all species of Platyrrhinus, the

third metacarpal was subequal in length to the

ﬁfth metacarpal. Simmons (1996: character 8)

used this character in her phylogeny of the genus

Micronycteris.

Character 60—Insertion of the posterior

edge of the plagiopatagium. Onto the ankle re-

gion (0); onto the ﬁrst metatarsal (1). The pos-

18 FIELDIANA: ZOOLOGY

terior edge of the plagiopatagium inserts at the

ﬁrst metatarsal in all the species of Platyrrhinus

(e.g., P. helleri ‘‘Eastern,’’ P. reciﬁnus), Urod-

erma magnirostrum, and Vampyrodes. The pos-

terior edge of the plagiopatagium inserts at the

ankle region in Carollia subrufa and Sturnira er-

ythromos. Owen (1987: character 6) and Wetter-

er et al. (2000: character 84) used this character,

and I agree with the scores assigned to Platyr-

rhinus and the outgroup. Marques-Aguiar (1994:

character 7) used this character in her phylogeny

of the genus Artibeus, as did Straney (1980:

characters G 9–13) in his phylogeny of the sub-

family Phyllostominae.

Results

Parsimony analysis of the data resulted in two

most parsimonious trees of 162 steps each (CI

0.54; CI excluding uninformative characters

0.52; RI

0.58). A strict consensus of these trees

with the bootstrap and decay values is shown in

Figure 14. Monophyly of Platyrrhinus was sup-

ported with bootstrap and decay values of 84%

and 3%, respectively.

Two main clades were observed within Platyr-

rhinus. One is comprised of the smaller species

P. lineatus and P. reciﬁnus and is supported

weakly by a bootstrap value of 33% and a decay

value of 1. In this clade, the basal species P. line-

atus and P. reciﬁnus were unresolved, whereas

the clade formed by P. helleri ‘‘Eastern’’

brachycephalus

P. helleri ‘‘Western’’ was sup-

ported by a bootstrap value of 72% and a decay

value of 1. The clade formed by P. brachyceph-

alus

P. helleri ‘‘Western’’ also was supported

by a bootstrap value of 72% and a decay value of

1. The second main clade comprised of the re-

maining nine species of Platyrrhinus was com-

pletely resolved and was supported by a bootstrap

value of 43% and a decay value of 2. Platyrrhinus

chocoensis, P. dorsalis, and P. aurarius occupied

the three most basal branches, followed by P. ni-

gellus

P. dorsalis ‘‘Centro-Sur,’’ P. dorsalis

‘‘Norte,’’ P. infuscus, P. vittatus ‘‘Northern,’’ and

P. vittatus ‘‘Southern.’’

There was strong support (bootstrap

98% and

decay

6) for Vampyrodes caraccioli as sister

taxon of Platyrrhinus. The analysis also supported

(bootstrap

100% and decay

10) Uroderma

magnirostrum as sister taxa of the clade com-

prised of Platyrrhinus

Vampyrodes. The most

basal branches of the tree were occupied by Car-

ollia subrufa (subfamily Carolliinae) and Sturnira

erythromos (most basal genus of the Stenoder-

matinae).

Discussion and Conclusions

The present phylogenetic analysis strongly sup-

ports Platyrrhinus monophyly but only weakly

supports the interspeciﬁc relationships (Fig. 14).

Owen’s (1987) phylogenetic hypothesis also sup-

ported Platyrrhinus monophyly but did not in-

clude all the species, nor did it provide support

values. Three synapomorphies are described for

the ﬁrst time for Platyrrhinus: sulcus on the an-

terior face of the P4, both labial and lingual cin-

gula present on the m1, and stylid cusps present

on the anterior face of the m1 protoconid.

Two main divisions of Platyrrhinus were evi-

dent in both analyses. However, they were weak-

ly supported, so I do not suggest recognizing

them as separate subgenera. The ﬁrst clade in-

cludes all the smaller species of Platyrrhinus (P.

brachycephalus

P. helleri ‘‘Eastern’’

P. hel-

leri ‘‘Western’’), P. lineatus, and P. reciﬁnus and

is supported by six characters: poorly marked but

distinguishable pinna folds (character 10), dense

and long hair on the dorsum of the feet (15), a

densely haired uropatagium edge (17), M1 par-

astyle present (32), labial cingulum at the base

of the M1 metacone present (34), and both labial

and lingual cingula present on p4 (49). The pre-

sent hypothesis strongly contradicts with Owen’s

interspeciﬁc phylogenetic hypothesis. Owen’s

(1987) preferred hypothesis placed P. brachy-

cephalus and P. helleri (the only ‘‘small’’ spe-

cies of Platyrrhinus recognized at that time) in

different clades. In the present study, P. brachy-

cephalus and the two forms of P. helleri were

placed in the same clade.

The second clade, containing the nine larger

species of Platyrrhinus, is supported by seven

synapomorphies: dark facial stripes (character 3),

absence of the basal protuberance where the genal

vibrissae are implanted (4), folds in the pinna well

marked (10), dorsal stripe deﬁnite but narrow

(11), posterior border of the hard palate ‘‘V’’-

shaped (18), postorbital process absent or poorly

developed (19), and M1 protocone small and

blunt (39).

The present analysis supports the hypothesis

of Velazco and Solari (2003), recognizing P.

19VELAZCO: PLATYRRHINUS PHYLOGENY

. 14. Strict consensus tree of the analysis with the characters unordered. Decay values are given above each

branch; bootstrap values are below.

dorsalis as a complex of three different species

(two not yet described). The analysis placed P.

dorsalis close to P. chocoensis, which also is a

species that occurs on the western slope of the

Andes. Platyrrhinus dorsalis ‘‘Norte’’ and P.

dorsalis ‘‘Centro-Sur’’ (the two undescribed spe-

cies) were placed closer to P. nigellus and P.

infuscus, species that are distributed on the east-

ern slope of the Andes. No differences were

found between the paratypes of P. umbratus and

the specimens of P. dorsalis. Therefore, I treat

P. umbratus as a junior synonym of P. dorsalis.

Platyrrhinus dorsalis ‘‘Norte’’ and P. dorsalis

‘‘Centro-Sur’’ exhibit several autopomorphies

that differentiate them from P. dorsalis and each

other. Consequently, these two taxa are named

and described here. Platyrrhinus lineatus and P.

nigellus demonstrate clear morphological differ-

ences and are located in different clades, sup-

porting their validity as species.

The branches representing P. helleri ‘‘Eastern’’

and P. helleri ‘‘Western’’ have sufﬁcient distin-

guishing characteristics to be considered different

species. The distribution of P. helleri ‘‘Eastern’’

is the widest of the genus, comprising most of

Central and South America and encompassing

20 FIELDIANA: ZOOLOGY

type localities of P. helleri (Peters, 1866), P. za-

rhinus (Allen, 1891), and P. zarhinus incarum

(Thomas, 1912). Comparisons between the de-

scriptions of these species and P. helleri ‘‘East-

ern’’ suggest that P. helleri is the appropriate bi-

nomial name for P. helleri ‘‘Eastern,’’ with P.

zarhinus and P. zarhinus incarum representing ju-

nior synonyms. Platyrrhinus helleri ‘‘Western,’’

which is found in the Choco´ region in Colombia

and Ecuador, does not have a name available and

is described here.

Platyrrhinus vittatus ‘‘Northern’’ and P. vitta-

tus ‘‘Southern’’ are the largest forms in this genus.

Platyrrhinus vittatus ‘‘Northern’’ is restricted to

the lowlands in Costa Rica, Panama, Atlantic and

Paciﬁc Colombia, and Venezuela. Platyrrhinus

vittatus ‘‘Southern’’ is distributed mainly in the

highlands of Ecuador, Peru, and Bolivia. The ho-

lotype of P. vittatus is from Puerto Cabello, Vene-

zuela, and is distinguishable both qualitatively

and morphometrically from P. vittatus ‘‘South-

ern.’’ Platyrrhinus vittatus ‘‘Southern’’ is larger

than P. vittatus ‘‘Northern’’ and has qualitative

morphological and morphometric differences.

Platyrrhinus vittatus ‘‘Southern’’ is a taxon with-

out an available name and therefore is described

here.

Like previous phylogenetic hypotheses for the

family Phyllostomidae that placed Platyrrhinus

and the monotypic genus Vampyrodes as sister

taxa (Baker et al., 2000, 2003; Lim, 1993; Owen,

1987 [only discrete characters]; Smith, 1976; Van

Den Bussche, 1992; Wetterer et al., 2000), the

present study places Vampyrodes as a sister taxa

of Platyrrhinus.

Taxonomic Diagnoses of the Platyrrhinus

Species

A taxonomic diagnosis for 14 species of Pla-

tyrrhinus used in the phylogenetic analysis is pre-

sented. This includes descriptions of four new

species. The diagnoses are based on the ACCT-

RAN and DELTRAN optimizations of the data.

Unambigously derived conditions are in italics.

States with ambiguous optimizations are not ital-

icized. All observations are based on adult indi-

viduals, external and craniodental measurements

are in millimeters, and body mass (weight) is in

grams. The ﬁrst ﬁve measurements listed here

were obtained from skin tags or ﬁeld notes made

by the collector of each specimen; the other di-

mensions were measured with a digital caliper to

the nearest 0.01 mm. Measurements are deﬁned

as follows:

Weight (W): Body mass in grams.

Total length (TL): Distance from the snout to

the tip of the last caudal vertebra.

Hind foot length (HL): Distance from the an-

terior edge of the base of the calcar to the tip of

the claw of the longest toe.

Ear length (E): Distance from the notch to the

ﬂeshy tip of the pinna.

Forearm length (FA): Distance from the olec-

ranon process to the carpals.

Tibia length (Tibia): Distance from the prox-

imal end of the tibia to the posterior base of the

calcar.

Greatest length of skull (GLS): Distance from

the most posterior point of the occiput to the an-

teriormost point of the premaxilla (excluding in-

cisors).

Condiloincisive length (CIL): Distance be-

tween the posterior-most point of the occipital

condyles and the anterior-most point on the upper

incisors.

Condilocanine length (CCL): Distance be-

tween the posterior-most point of the occipital

condyles and the anterior-most point on the upper

canines.

Postorbital breadth (PB): Least breadth across

the frontals posterior to the postorbital processes

or bulges.

Zygomatic breadth (ZB): Greatest breadth

across zygomatic arches.

Braincase breadth (BB): Greatest breadth of

the braincase, excluding the mastoid and paraoc-

cipital processes.

Mastoid breadth (MB): Greatest breadth

across the mastoid region.

Maxillary toothrow length (MTRL): From

the anterior-most edge of the canine crown to the

posterior-most edge of the crown of M3.

Breadth across molars (BAM): Greatest

breadth across the outer edges of the crowns of

the upper molars.

Measurements of male and females were com-

bined in species comparisons because no differ-

ences in size were apparent between males and

females in any species of Platyrrhinus. Velazco

and Solari (2003) evaluated sexual dimorphism in

the Peruvian populations of P. dorsalis ‘‘Norte,’’

P. dorsalis ‘‘Centro-Sur,’’ and P. nigellus and

concluded that these three species did not dem-

onstrate signiﬁcant sexual dimorphism.

21VELAZCO: PLATYRRHINUS PHYLOGENY

ABLE

2. Measurements of the type series of Pla-

tyrrhinus albericoi.

Holotype

MUSM

19149

Paratype

FMNH

170145

Sex Female Female

(pregnant)

Weight

Total length

Hind foot length

Ear length

Forearm length

100

Tibia length

Greatest length of skull

Condyloincisive length

Condylocanine length

Postorbital beadth

23.93

32.84

32.63

31.89

7.65

23.71

32.51

31.63

30.82

7.39

Zygomatic breadth

Braincase breadth

Mastoid breadth

Maxillary toothrow length

Breadth across molars

20.85

13.38

15.85

13.89

15.23

20.68

13.58

16.12

13.44

15.57 F

. 15. Distributional map of Platyrrhinus albericoi.

Systematics

Family Phyllostomidae Gray, 1825

Subfamily Stenodermatinae Gervais, 1856

Genus Platyrrhinus Saussure, 1860

Platyrrhinus albericoi, new species. Figures 1, 16

(bottom)

Vampyrops vittatus Gardner and Carter, 1972b: 74–

75 (part)

Vampyrops vittatus Koopman, 1978: 11

Vampyrops vittatus Swanepoel and Genoways, 1979:

66, 104 (part)

Vampyrops vittatus Barquez and Olrog, 1980: 53–54

Vampyrops vittatus Anderson et al., 1982: 7–8

Vampyrops vittatus Anderson, 1985: 7

Vampyrops vittatus Alberico, 1990: 349 (part)

Platyrrhinus vittatus Anderson, 1993: 8, 24, 69

Platyrrhinus vittatus Anderson, 1997: 36, 244, 589

(part)

Platyrrhinus vittatus Albuja, 1999: 134–135 (part)

Platyrrhinus vittatus Emmons et al., 2001: 255

Platyrrhinus vittatus Romo, 2001: 258

Platyrrhinus vittatus Solari et al., 2001: 112, 263

Platyrrhinus vittatus Salazar-Bravo et al., 2003: 15

Platyrrhinus vittatus ‘‘Southern’’ (this paper)

YPE

ATERIAL

—The holotype, MUSM 19149,

is an adult female collected by Sergio Solari (orig-

inal ﬁeld number SS 2067) on 14 April 2001 at

San Pedro, Paucartambo-Pilcopata road, 1480 m

in elevation, Province of Paucartambo, Depart-

ment of Cuzco, Peru, approximately 13

W. The type locality is in the Cultural

Zone of the Manu Biosphere Reserve. The skin,

skull, and alcohol preserved carcass were in good

condition. Frozen tissues are deposited at the

Field Museum of Natural History (FMNH

172107).

The paratype is an adult female (FMNH

170145) from Suecia, km 138.5 Carretera Shin-

tuya, Province of Paucartambo, Department of

Cuzco, Peru. The holotype and paratype as well

as other specimens from the known distributional

range are listed in Appendix 1. Dimensions of

each specimen of the type series of Platyrrhinus

albericoi are provided in Table 2.

ISTRIBUTION

—Platyrrhinus albericoi is cur-

rently known from the Eastern slope of the Andes

in Ecuador, Peru, and Bolivia. Currently, the

northern limit of its distribution is the Province of

Pichincha (Ecuador), and the southern limit is the

Department of Cochabamba (Bolivia). However,

this taxon is expected to occur in southern Colom-

bia (Fig. 15). The elevational range comprise from

1480 to 2500 m.

TYMOLOGY

—The speciﬁc epithet is chosen to

honor Michael Alberico, who devoted his scien-

tiﬁc career to the study of Colombian mammals.

IAGNOSIS

—Size large (FA

62–63 mm; CIL

31.6–32.6 mm; W

55–68 g); facial stripes

brilliant-white; folds in the pinna poorly marked

but distinguishable;fringe of hair on the edge of

the uropatagium dense;paraoccipital process

well developed; M1 mesostyle present; M2 me-

22 FIELDIANA: ZOOLOGY

ABLE

3. Selected measurements

a,b

of members of the genus Platyrrhinus.

Platyrrhinus

aurarius Platyrrhinus

brachycephalus Platyrrhinus

chocoensis Platyrrhinus

dorsalis Platyrrhinus

helleri

30.5

0.7

30–31 (2)

78.5

4.9

75–82 (2)

15.0

0.8

14–17 (12)

13.6

2.8

10–20 (20)

58.3

3.8

52–68 (33)

11.0

1.0

9–13 (37)

30.5

2.1

29–32 (2)

76.5

2.1

75–78 (2)

14.5

0.7

14–15 (2)

—

75.5

6.4

71–80 (2)

13.2

0.7

12–15 (11)

14.5

2.7

11–19 (10)

57.7

3.9

52–65 (15)

10.5

0.9

8–12 (21)

Tibia

20.7

0.6

20–22 (12)

52.7

1.1

51–54 (12)

21.30

0.76

20.00–22.00 (12)

16.3

1.2

13–19 (33)

37.6

1.8

33–42 (37)

15.36

0.93

13.18–17.60 (33)

19.0

1.4

18–20 (2)

48.5

0.7

48–49 (2)

19.65

1.72

18.43–20.86 (2)

18.8

0.7

18–20 (11)

48.9

1.3

46–50 (11)

20.32

1.34

17.39–21.88 (11)

15.7

1.4

12–17 (14)

37.8

1.5

35–40 (21)

15.04

0.69

13.60–16.00 (21)

GLS

CIL

CCL

27.23

0.38

26.53–27.87 (12)

26.18

0.39

25.62–27.01 (12)

25.55

0.36

25.12–26.35 (12)

20.44

0.50

19.50–21.53 (47)

19.08

0.53

18.30–20.26 (47)

18.64

0.50

17.75–19.86 (47)

27.33–0.49

26.77–27.69 (3)

25.59

0.84

24.64–26.22 (3)

25.18

0.76

24.33–25.80 (3)

26.42

0.47

25.46–27.40 (13)

25.06

0.48

24.33–25.75 (13)

24.59

0.48

23.82–26.43 (13)

21.21

0.75

19.04–22.50 (67)

19.95

0.80

18.03–21.28 (66)

19.49

0.79

17.65–20.80 (67)

6.65

0.12

6.39–6.83 (12)

16.89

0.33

16.38–17.57 (9)

11.85

0.19

11.55–12.28 (12)

5.39

0.22

4.92–6.15 (48)

12.29

0.55

11.46–13.64 (38)

9.31

0.30

8.76–9.94 (47)

6.55

0.22

6.33–6.77 (3)

16.91

0.34

16.57–17.26 (3)

11.82

0.08

11.74–11.89 (3)

6.32

0.12

6.13–6.59 (13)

16.13

0.67

14.88–17.10 (12)

11.52

0.33

10.74–11.93 (13)

5.44

0.15

5.17–5.88 (67)

12.56

0.49

11.49–13.54 (55)

9.38

0.27

8.62–9.89 (66)

MTRL

BAM

14.01

0.34

13.42–14.58 (12)

11.07

0.26

10.71–11.55 (12)

12.45

0.33

11.88–12.90 (12)

10.66

0.32

10.04–11.44 (47)

7.37

0.27 (41)

6.82–7.94 (41)

8.75

0.37

8.11–9.72 (48)

13.75

0.43

13.26–14.06 (3)

10.94

0.29

10.70–11.25 (3)

11.55

0.12

11.45–11.69 (3)

13.34

0.33

12.76 –13.91 (13)

10.68

0.33

10.17–11.34 (13)

11.49

0.49

10.88–12.60 (13)

10.88

0.38

10.04–11.61 (66)

7.88

0.40

6.96–8.59 (63)

9.02

0.45

8.14–9.80 (65)

tastyle absent; length of the third metacarpal sub-

equal to the ﬁfth metacarpal.

ESCRIPTION

—The largest known species of

Platyrrhinus (FA

62–63 mm; GLS

32.5–32.8

mm; Tables 2–3). Dorsal fur dark brown, ventral

fur light brown, dorsal and ventral hair tricolored;

dorsomedial facial stripes more marked than the

ventrolateral; dorsal hairs

8 mm long on the

back; narrow dorsal stripe brighter than the facial

ones; folds in the pinna poorly marked but distin-

guishable; six vibrissae surrounding the margins

of the noseleaf in a single array; four submental

vibrissae present; one or two interramal vibrissae

present; noseleaf longer than wide, inferior border

of the horseshoe completely free of the upper lip;

dense and long hair on the dorsum of the feet;

‘‘U’’-shaped posterior edge of the uropatagium

with a densely haired fringe, uropatagium extends

along the midline 4.5–7 mm long; insertion of the

posterior edge of the plagiopatagium onto the ﬁrst

metatarsal; third metacarpal subequeal in length

with ﬁfth metacarpal.

Almost imperceptible fossa on the squamosal

end of the zygomatic arch; postorbital process

moderately developed; paraoccipital process well

developed; upper median incisors convergent and

in contact; sulcus on the posterior face of the up-

per canines; two stylar cusps present on the pos-

terior face of the P4; stylar cusp present on the

cingulum of the lingual face of the M1 metacone;

M1 metastyle and mesostyle present; M1 proto-

cone moderately developed; stylar cusp present on

the lingual face of the M2 paracone; M2 metastyle

absent; M2 parastyle present; stylar cusp present

on the lingual face of the M2 metacone; lingual

cingulum of the M2 metacone restricted to the

metacone; two sulci present on the lingual face of

the main cone of p4; both labial and lingual cin-

gula present on p4; stylid cusps absent on the an-

terior or anterolingual face of the main cone of

p4; m1 paraconid absent; stylid cusp present on

the anterior face of the m1 protoconid; poorly de-

veloped m1 metaconid; m2 hypoconid absent;

stylid cusp between the metaconid and protoconid

23VELAZCO: PLATYRRHINUS PHYLOGENY

ABLE

3. Continued.

Platyrrhinus

infuscus Platyrrhinus

lineatus Platyrrhinus

nigellus Platyrrhinus

reciﬁnus Platyrrhinus

vittatus

44.5

5.1

36–59 (23)

88.4

6.4

77–105 (36)

15.5

0.9

14–18 (57)

21.9

3.1

18–28 (20)

70.1

4.7

60–78 (22)

12.3

0.9

10–1 (28)

21.6

3.1

19–30 (13)

65.9

3.3

60–74 (32)

12.1

0.9

10–15 (33)

17.7

0.8

17–19 (6)

91.3

1.5

89–93 (6)

12.2

0.7

11–13 (6)

62.3

2.5

60–65 (3)

92.6

7.3

83–100 (5)

14.9

1.5

13–17 (7)

Tibia

22.4

1.1

20–24 (36)

57.9

1.6

54–62 (52)

23.75

1.13

20.80–26.00 (49)

19.4

1.4

15–2 (22)

47.1

0.8

46–48 (22)

19.51

0.82

18.12–21.00 (20)

18.5

1.1

16–21 (31)

43.0

1.6

40–47 (33)

16.94

0.73

15.62–18.46 (30)

19.2

0.7

18–20 (6)

42.5

0.5

42–43 (6)

16.28

0.76

15.20–17.50 (6)

24.0

1.0

23–25 (5)

59.8

1.2

57–61 (7)

21.74

0.90

20.43–22.82 (7)

GLS

CIL

CCL

29.74

0.73

27.96–31.22 (63)

28.32

0.65

26.39–29.89 (61)

27.70

0.63

25.93–29.01 (63)

24.01

0.46

23.08–24.94 (29)

22.52

0.39

21.98–23.45 (24)

21.89

0.39

21.36–22.79 (24)

24.43

0.51

23.51–25.88 (35)

23.04

0.54

21.94–24.50 (34)

22.56

0.54

21.57–23.98 (35)

23.32

0.29

23.03–23.76 (6)

22.26

0.26

21.99–22.61 (6)

21.73

0.21

21.55–22.06 (6)

30.57

0.82

28.90–31.39 (7)

29.63

1.09

28.12–31.29 (8)

28.91

1.03

27.48–30.41 (8)

6.88

0.20

6.32–7.34 (65)

18.39

0.48

17.42–19.33 (60)

12.69

0.28

11.96–13.35 (62)

6.31

0.18

5.98–6.61 (30)

14.42

0.35

13.80–15.28 (26)

10.77

0.24

10.77–11.25 (29)

6.12

0.17

5.61–6.41 (36)

14.30

0.35

13.45–15.48 (35)

10.72

0.19

10.37–11.20 (36)

5.74

0.21

5.46–6.10 (6)

14.20

0.33

13.55–14.45 (6)

10.44

0.21

10.18–10.80 (6)

7.49

0.24

7.23–7.89 (8)

19.07

0.43

18.41–19.67 (8)

13.17

0.26

12.81–13.50 (8)

MTRL

BAM

15.15

0.38

14.16–15.97 (63)

12.17

0.28

11.53–12.79 (63)

13.79

0.39

12.88–14.52 (63)

12.22

0.33

11.76–12.89 (25)

8.91

0.19

8.57–9.30 (30)

10.13

0.21

9.62–10.83 (31)

12.32

0.32

11.66–13.18 (35)

9.54

0.30

8.96–10.16 (31)

10.55

0.29

10.04–11.09 (36)

11.94

0.30

11.57–12.41 (6)

8.96

0.26

8.62–9.39 (6)

10.41

0.30

9.90–10.71 (6)

15.43

0.38

14.68–15.96 (7)

12.72

0.45

12.02–13.30 (7)

14.43

0.57

13.59–15.37 (8)

Weight is in grams, all other measurements are in millimeters.

Summary statistics (mean and standard deviation [above], observed range and sample size [below]) of measure-

ments for each species (see Appendix 1 for a list of the specimens measured).

of m2 present; both labial and lingual cingula pre-

sent on m2.

OMPARISONS

—Platyrrhinus albericoi is found

sympatrically with P. dorsalis ‘‘Norte,’’ P. dor-

salis ‘‘Centro-Sur,’’ and P. nigellus. Platyrrhinus

albericoi is easily distinguishable from these three

species by its larger forearm length and greatest

length of skull: P. albericoi (FA

62–63 mm;

GLS

32.5–32.8 mm), P. dorsalis ‘‘Centro-Sur’’

(FA

45–51 mm; GLS

25.0–26.8 mm), and

P. nigellus (FA

40–47 mm; GLS

23.5–25.9

mm). Platyrrhinus albericoi has been traditionally

confused with P. vittatus, so subsequent compar-

isons are restricted to these two large species.

Externally, Platyrrhinus albericoi can be distin-

guished from P. vittatus by darker dorsal fur and

brighter and wider facial stripes in P. albericoi.

Moreover, folds in the pinna are poorly marked

but distinguishable in P. albericoi, whereas in P.

vittatus the folds in the pinna are well marked.

The dorsal stripe is white and narrow in P. alber-

icoi, whereas in P. vittatus the dorsal stripe is

white and wide. The posterior edge of the uro-

patagium always is ‘‘U’’ shaped in P. albericoi,

whereas it can be ‘‘U’’ or ‘‘V’’ shaped in P. vit-

tatus. A densely haired uropatagium fringe is pre-

sent in P. albericoi, whereas the uropatagium

fringe is usually hairy, occasionally sparsely in P.

vittatus. The third metacarpal is greater in length

than the ﬁfth metacarpal in P. albericoi but is sub-

equal in length in P. vittatus.

Cranially, Platyrrhinus albericoi and P. vittatus

are very similar. The only apparent differences are

that the cranium of P. albericoi is more robust

than that of P. vittatus, and the paraoccipital pro-

cess is better developed in P. albericoi.

Dentally, Platyrrhinus albericoi and P. vittatus

differ as follows: M1 mesostyle present in P. al-

bericoi and absent in P. vittatus; moderately de-

veloped M1 protocone in P. albericoi, whereas in

24 FIELDIANA: ZOOLOGY

P. vittatus the protocone on M1 tends to be better

developed; M2 metastyle absent in P. albericoi

and present in P. vittatus; stylid cusps absent on

the anterior or anterolingual face of the main cone

of p4 in P. albericoi, whereas one stylid cusp on

the anterolingual face of the main cone of p4 is

present in P. vittatus (Fig. 16).

Platyrrhinus aurarius (Handley and Ferris, 1972)

Vampyrops aurarius Handley and Ferris, 1972. Type

locality: km 125, 85 km SSE El Dorado, State of

Bolivar, Venezuela.

OLOTYPE

—Vampyrops aurarius USNM

387163, an adult male preserved as skin and skull,

was collected by Merlin D. Tuttle and Arden L.

Tuttle on 18 May 1966 at km 125, 85 km SSE El

Dorado, State of Bolivar, Venezuela, elevation

1000 m.

ISTRIBUTION

—Endemic to the Guianan Shield

(southern Venezuela, Guyana, and Suriname),

from 140 to 1250 m (Fig. 17).

IAGNOSIS

—Size medium (FA

51–54 mm;

CIL

25.6–27.0 mm; W

30–31 g; Table 3);

dorsal fur tetracolored;postorbital process mod-

erately developed;paraoccipital process well de-

veloped; only the labial cingulum present on p4.

Platyrrhinus brachycephalus (Rouk and Carter,

1972)

Vampyrops brachycephalus Rouk and Carter, 1972.

Type locality: 3 mi S Tingo Maria, Department of

Hua´nuco, Peru.

Vampyrops latus Handley and Ferris, 1972. Type lo-

cality: San Juan, Province of Oxapampa, Depart-

ment of Pasco, Peru (USNM 364408).

Vampyrops latus saccharus Handley and Ferris, 1972.

Type locality: Manacal, 5 km S and 25 km E Ca-

ru´pano, State of Sucre, Venezuela (USNM

408411).

OLOTYPE

—Vampyrops brachycephalus TCWC

12193, adult male preserved as skin and skull,

was collected by D. C. Carter (original ﬁeld num-

ber 5513) on 28 August 1964 at 3 mi S Tingo

Maria, Department of Hua´nuco, Peru, elevation

2400 ft (731 m).

ISTRIBUTION

—Known from the Amazon Basin

(Brazil, Bolivia, Colombia, Ecuador, French Gui-

ana, Guyana, Peru, Suriname, and Venezuela) and

the mountain forests in Ecuador and Peru, from

90 to 1295 m (Fig. 17).

Diagnosis—Size small (FA

33–42 mm; CIL

18.3–20.3 mm; W

10–20 g) (Table 3); one

vibrissa present on the upper lip, ventral to the

vibissae that surround the margin of the noseleaf;

ﬁve submental vibrissae present on each side of

the chin; ventral fur tricolored; fringe of hair on

the edge of the uropatagium usually hairy, occa-

sionally sparsely;a well-developed M1 proto-

cone;two stylid cusps present on the anterior face

of the main cone of p4.

Platyrrhinus chocoensis Alberico and Velasco,

1991

Platyrrhinus chocoensis Alberico and Velasco, 1991.

Type locality: Quebrada El Platinero, 12 km W Ist-

mina (by road), Department of Choco´, Colombia.

OLOTYPE

—Platyrrhinus chocoensis UV 3817,

adult male preserved as skin and skull, was col-

lected by M. S. Alberico (original ﬁeld number,

MSA 1316) on 16 April 1984 at Quebrada El Pla-

tinero, 12 km W Istmina (by road), Department

of Choco´, Colombia, elevation 100 m.

ISTRIBUTION

—Found in the lowlands of the

Choco´ biogeographic region of Colombia, to

northwestern Ecuador, from35 to 305 m (Fig. 18).

IAGNOSIS

—Size medium (FA

48–49 mm;

CIL

24.6–26.2 mm; W

29–32 g; Table 3);

ventral fur bicolored; M1 protocone small and

blunt; only the labial cingulum present on the p4;

stylid cusp between the metaconid and protoconid

of the m2 present.

Platyrrhinus dorsalis (Thomas, 1900)

Vampyrops dorsalis Thomas, 1900. Type locality:

Paramba, Province of Imbabura, Ecuador.

Vampyrops umbratus Lyon, 1902. Type locality: San

Miguel, Department of La Guajira, Colombia

(MCZ B8180).

Vampyrops oratus Thomas, 1914. Type locality:

‘‘Galifari’’ Picacho de Galipa´n, Cerro del Avila,

Distrito Federal, Venezuela (BMNH 14.7.27.1).

Vampyrops aquilus Handley and Ferris, 1972. Type

locality: On the head of the Rı´o Pucro, Cerro Malı´,

Province of Darie´n, Panama (USNM 338025).

OLOTYPE

—Vampyrops dorsalis BMNH

99.12.5.1, juvenile male preserved as skin and

skull, was collected by R. Miketta (original ﬁeld

number 61) on 14 April 1899 at Paramba, Prov-

ince of Imbabura, Ecuador, elevation 1100 m.

ISTRIBUTION

—From southern Panama, along

both slopes of the Andes in Colombia, and only

the western slope in Ecuador, and northern Vene-

zuela, from 230 to 2012 m (Fig. 18).

IAGNOSIS

—Size medium (FA

46–50 mm;

CIL

24.3–25.7 mm; Table 3); folds in the pinna

25VELAZCO: PLATYRRHINUS PHYLOGENY

. 16. Labial views of the left p2 and p4 illustrating taxonomic differences in the presence or absence of

accessory cusps on the anterolingual face of the main cone of p4. (Top)Platyrrhinus vittatus (LSUMZ 25465)

with one accessory cusp (arrow). (Bottom)Platyrrhinus albericoi (FMNH 170145) without accessory cusp

(arrow).

absent; M1 protocone small and blunt; only the

labial cingulum present on p4; stylar cusp present

on the lingual face of the M2 metacone; both la-

bial and lingual cingula present on p4.

Platyrrhinus helleri (Peters, 1866)

Vampyrops helleri Peters, 1866. Type locality: Mex-

ico.

Vampyrops zarhinus Allen, 1891. Type locality: Obis-

po, Canal Zone, Panama (MCZ 3211).

Vampyrops zarhinus incarum Thomas, 1912. Type lo-

cality: Pozuzo, Department of Pasco, Peru (BMNH

12.1.15.1).

YNTYPE

—Vampyrops helleri ZMB 3276, adult,

preserved as skin with the skull not removed, was

collected by Heller in 1850 in Mexico.

ISTRIBUTION

—From Mexico (Oaxaca and Ve-

racruz) to Peru, Bolivia, and Amazonian Brazil,

plus Trinidad and Tobago, from 160 to 1295 m

(Fig. 19).

IAGNOSIS

—Size small (FA

35–40 mm; CIL

18.0–21.3 mm; W

11–19 g; Table 3); ventral

fur bicolored; hair on the dorsum of the feet in-

termediate in density and length; M1 protocone

small and blunt;one stylid cusp present on the

anterior face of the main cone of p4.

26 FIELDIANA: ZOOLOGY

. 17. Distributional map of Platyrrhinus aurarius and P. brachycephalus.

Platyrrhinus infuscus (Peters, 1880)

Vampyrops infuscus Peters, 1880. Type locality: Gru-

ta de Ninabamba, Hacienda Ninabamba, Province

of Hualgayoc, Department of Cajamarca, Peru.

Vampyrops fumosus Miller, 1902. Type locality: Hui-

tanaa˜, upper Purus River, State of Amazonas, Bra-

zil (USNM 105530).

Vampyrops intermedius Marinkelle, 1970. Type lo-

cality: Mina de Upin, near Restrepo, Department of

Meta, Colombia (Universidad de los Andes, Co-

lombia 14885).

OLOTYPE

—The holotype collected by L. Tac-

zanowski from Gruta de Ninabamba, Hacienda

Ninabamba, Province of Hualgayoc, Department

of Cajamarca, Peru, and deposited in the Warsaw

Museum (Polska Akademia Nauk, Instytut Zool-

ogiczny) was apparently destroyed by ﬁre, per-

haps as early as 1936 (Gardner & Carter, 1972b).

In 1972, Gardner and Carter designated a neo-

type: TCWC 12199, adult male, preserved as

skin and skull, collected by D. C. Carter (original

ﬁeld number 5477) on 26 August 1964 at 2 mi

N Tingo Maria, Province of Leoncio Prado, De-

partment of Hua´nuco, Peru, elevation 2000 ft

(610 m).

ISTRIBUTION

—Colombia to Peru, Bolivia, and

NW Brazil, from 183 to 1900 m (Fig. 20).

IAGNOSIS

—Size large (FA

54–62 mm; CIL

26.4–29.9 mm; W

36–59 g; Table 3); dorsal

stripe indistinct, thin and obscured; dorsal fur

short,

6.30 mm;sparse and short hair on the

dorsum of the feet; uropatagium fringe usually

hairy, occasionally sparsely; paraoccipital process

well developed; M1 parastyle present;m1 meta-

27VELAZCO: PLATYRRHINUS PHYLOGENY

. 18. Distributional map of Platyrrhinus chocoensis and P. dorsalis.

conid developed;stylid cusp absent between the

metaconid and protoconid of m2.

Platyrrhinus ismaeli, new species Figure 21, 23

(top)

cf. Vampyrops aurarius Lemke et al., 1982: 230–231

Vampyrops sp. C Alberico, 1990: 349–352

Platyrrhinus dorsalis Pacheco and Patterson, 1991:

101, 104, 111, 112, 113, 114, 121

Platyrrhinus dorsalis Albuja, 1999: 135–136 (part)

Platyrrhinus ‘‘dorsalis norte’’ Solari et al., 2001: 263

Platyrrhinus dorsalis ‘‘Norte’’ Velazco and Solari,

2003

Platyrrhinus dorsalis ‘‘Norte’’ (this paper)

YPE

ATERIAL

—The holotype, MUSM 4946,

is an adult collected by Bruce D. Patterson (orig-

inal ﬁeld number BDP 2484) on 2 May 1987 at

19 km E of Balsas, 1945 m (6380 ft) in elevation,

Province of Chachapoyas, Department of Ama-

zonas, Peru. The skin, skull, and skeleton were in

good condition. Frozen tissues are deposited at the

Field Museum of Natural History (FMNH

129138).

Paratypes include ﬁve adult specimens: one fe-

male (FMNH 129134) and one male (FMNH

129136) from the type locality; and two females

(FMNH 129139, 129146) and one male (FMNH

28 FIELDIANA: ZOOLOGY

. 19. Distributional map of Platyrrhinus helleri.

129143) from Hacienda Limo´n, W of Balsas,

Province of Celendin, Department of Cajamarca,

Peru. The holotype and paratypes along with other

specimens from the known distributional range

are listed in Appendix 1. Measurements of each

specimen of the type series of Platyrrhinus is-

maeli are provided in Table 4.

ISTRIBUTION

—Platyrrhinus ismaeli (Fig. 21) is

currently known from both slopes of the Andes in

Colombia, Ecuador, and Peru. The northern limit

of the distribution is the Department of Huila (Co-

lombia), and the southern limit is the Department

of San Martin (Peru). The elevation is from 1230

to 2950 m (Fig. 22).

TYMOLOGY

—Platyrrhinus ismaeli is named in

honor of Ismael Ceballos Bendezu´, an interna-

tionally well-known naturalist from Cuzco, Peru,

in recognition of his important contributions to the

study of Peruvian bats.

IAGNOSIS

—Size medium (FA

50–56 mm;

CIL

26.6–28.2 mm; W

30–51 g); folds in

the pinna absent; sulcus on the posterior face of

the paracone not joined to the cingulum of the

lingual face of the metacone on M1; m1 meta-

conid poorly developed.

ESCRIPTION

—A medium-sized species of Pla-

tyrrhinus (FA

50–56 mm; GLS

27.8–30.1

mm) with dark brown dorsal fur, grayish ventral

29VELAZCO: PLATYRRHINUS PHYLOGENY

. 20. Distributional map of Platyrrhinus infuscus and P. lineatus.

fur, and tricolored dorsal and ventral hair. Dor-

somedial and ventrolateral facial stripes obviously

subequal and darkish, dorsal hairs 8–10 mm long

on the back; narrow dorsal stripe brighter than the

facial ones; folds in the pinna poorly marked but

distinguishable; six vibrissae surrounding the

margins of the noseleaf in a single array; four sub-

mental vibrissae present; one interramal vibrissa

present; noseleaf longer than wide, inferior border

of the horseshoe completely free of the upper lip;

dense and long hair on the dorsum of the feet;

‘‘U’’-shaped posterior edge of the uropatagium

with a densely haired fringe, uropatagium extends

along the midline 4.5–7 mm long; insertion of the

posterior edge of the plagiopatagium onto the ﬁrst

metatarsal; third metacarpal shorter than ﬁfth

metacarpal; fossa on the squamosal end of the zy-

gomatic arch shallow; upper median incisors con-

vergent and in contact; sulcus present on the pos-

terior face of the upper canines; two stylar cusps

present on the posterior face of P4; labial cingu-

lum present at the base of the metacone of M1;

stylar cusp present on the cingulum of the lingual

face of the M1 metacone; sulcus on the posterior

face of the paracone not joined to the cingulum

of the lingual face of the metacone on M1; M1

metastyle present; M1 protocone small and blunt;

stylar cusp present on the lingual face of the M2

paracone; M2 metastyle present; two sulci present

on the lingual face of the main cone of p4; m1

paraconid absent; m1 entoconid present; stylid

cusp present on the anterior face of the m1 pro-

30 FIELDIANA: ZOOLOGY

ABLE

4. Measurements of the type series of Platyrrhinus ismaeli.

Holotype

MUSM

4946

Paratype

FMNH

129134

Paratype

FMNH

129136

Paratype

FMNH

129139

Paratype

FMNH

129143

Paratype

FMNH

129146

Sex

Weight

Total length

Hind foot length

Ear length

Male

Female

Male

Female

Male

Female

Forearm length

Tibia length

Greatest length of skull

Condyloincisive length

Condylocanine length

21.68

27.94

27.10

26.22

19.52

27.77

26.57

26.07

21.68

28.15

27.47

26.88

21.27

28.14

27.06

26.49

20.56

28.47

26.83

26.14

21.31

28.17

26.62

26.06

Postorbital breadth

Zygomatic breadth

Braincase breadth

Mastoid breadth

Maxillary toothrow length

Breadth across molars

6.46

17.13

12.00

13.99

11.35

12.63

6.27

16.93

11.57

13.63

11.43

12.65

6.57

18.21

12.29

14.72

11.66

12.81

6.43

17.31

12.03

14.04

11.75

13.08

6.36

17.33

12.02

13.98

11.33

12.45

6.65

16.41

11.70

14.06

11.48

12.41

. 21. Frontal view of Platyrrhinus ismaeli; adult male photographed at Hacienda Limo´n, Cajamarca, Peru, by

B. D. Patterson.

toconid; poorly developed m1 metaconid; m2 hy-

poconid absent; stylid cusp present between the

metaconid and protoconid of m2; both labial and

lingual cingula present on m2. Velazco (2001) and

Velazco and Solari (2003) did not ﬁnd secondary

sexual dimorphism in populations of P. ismaeli

from Peru in either qualitative morphologic or

morphometric characters.

31VELAZCO: PLATYRRHINUS PHYLOGENY

. 22. Distributional map of Platyrrhinus ismaeli.

OMPARISONS

—In several localities Platyr-

rhinus ismaeli is found sympatrically with P. al-

bericoi and P. nigellus, and it occurs with P. dor-

salis in one locality at El Parque Nacional Nat-

ural de la Cueva de Los Guacharos, Department

of Huila, in Colombia. Platyrrhinus ismaeli is

easily distinguished from P. albericoi and P. ni-

gellus by its forearm length and greatest length

of skull: P. albericoi (FA

62–63 mm; GLS

32.5–32.8 mm) and P. nigellus (FA

40–47

mm; GLS

23.5–25.9 mm; Tables 2–4). It dif-

fers from P. dorsalis in its GLS, condiloincisive

length, condilocanine length, and maxillary

toothrow length (Tables 3–4). Platyrrhinus is-

maeli has traditionally been confused with P.

dorsalis, so subsequent comparisons are restrict-

ed to these two species.

Externally, Platyrrhinus ismaeli can be distin-

guished from P. dorsalis by two characters. Pla-

tyrrhinus ismaeli has dense and long hair on the

dorsal side of the feet, whereas the hair in the

dorsum of the feet is sparsely distributed and with

some long hairs in P. dorsalis. The hairs on the

edge of the uropatagium are densely haired and

longer than 2.1 mm in P. ismaeli, whereas P. dor-

salis presents a regular or occasionally sparsely

32 FIELDIANA: ZOOLOGY

haired uropatagium fringe, with hairs shorter than

1.5 mm.

Cranially, Platyrrhinus ismaeli and P. dorsalis

are very similar. Platyrrhinus ismaeli has a more

robust cranium with a wider rostrum. Moreover,

P. ismaeli has a shallow fossa on the squamosal

end of the zygomatic arch, lateral to the glenoid

fossa, whereas P. dorsalis has a deep fossa.

Dentally, Platyrrhinus ismaeli and P. dorsalis

exhibit some differences. Whereas in P. ismaeli

the sulcus on the posterior face of the paracone

not joined to the cingulum of the lingual face of

the metacone on M1, this discontinuity is absent

in P. dorsalis (Fig. 23). The lingual cingulum of

the M2 metacone is restricted to the metacone in

P. ismaeli, whereas in P. dorsalis the lingual cin-

gulum of the M2 metacone continuous to the M2

paracone. Finally, a poorly developed metaconid

on m1 is present in P. ismaeli, whereas P. dorsalis

lacks the m1 metaconid.

OMMENTS

—In his systematic review of Pla-

tyrrhinus from the northwestern South America,

Alberico (1990) identiﬁed Platyrrhinus sp.Casa

potentially new species from Colombia. Platyr-

rhinus sp.C(FA

50.9–53.2 mm) was collected

at several localities in the Cueva de los Guacharos

National Park near the headwaters of the Mag-

dalena River at midelevations (ca. 1500 m) in Co-

lombia. Platyrrhinus ismaeli has been collected at

Cueva de los Guacharos National Park, and the

range of variation of the FA size in Platyrrhinus

sp. C falls within the range of P. ismaeli (FA

50–56 mm). Therefore, it is likely that Platyr-

rhinus sp. C is conspeciﬁc with P. ismaeli.

Platyrrhinus lineatus (E

´. Geoffroy, 1810)

Phyllostoma lineatum E

´. Geoffroy St.-Hilaire, 1810.

Type locality: Asuncio´n, Paraguay.

Vampyrops lineatus sacrillus Thomas,1924. Type lo-

cality: Rio Doce, State of Espı´rito Santo, Brazil

(BMNH 23.12.12.9).

OLOTYPE

—Phyllostoma lineatum MNHN 953,

adult, preserved as skin and skull, was collected

by Felix de Azara in Asuncio´n, Paraguay.

ISTRIBUTION

—Lowlands from eastern and

southeastern Brazil, northern Uruguay, northeast-

ern Argentina, Paraguay, and Bolivia, from 122

to 971 m (Fig. 20).

IAGNOSIS

—Size medium (FA

46–48 mm;

CIL

22.0–23.4 mm; W

18–28 g; Table 3);

dense and long hair on the dorsum of the feet;

postorbital process well developed; paraoccipital

process well developed;m1 metaconid developed.

Platyrrhinus masu, new species. Figure 24

Vampyrops dorsalis Sanborn, 1951: 10

Vampyrops dorsalis Barquez and Orlog, 1980: 54

Vampyrops dorsalis Anderson et al., 1982: 7

Vampyrops dorsalis Anderson, 1985: 7

Platyrrhinus dorsalis Anderson, 1993: 8, 23, 69

Platyrrhinus dorsalis Anderson et al., 1993: 14, 24

Platyrrhinus dorsalis Anderson, 1997: 36, 37, 239,

240 (part)

Platyrrhinus dorsalis Emmons et al., 2001: 255

Platyrrhinus dorsalis Romo, 2001: 258

Platyrrhinus cf. dorsalis Solari et al., 2001: 112

Platyrrhinus ‘‘dorsalis sur’’ Solari et al., 2001: 263

Platyrrhinus dorsalis Salazar-Bravo et al., 2003: 15

Platyrrhinus dorsalis ‘‘Centro-Sur’’ Velazco and So-

lari (2003)

Platyrrhinus dorsalis ‘‘Centro-Sur’’ (this paper)

YPE

ATERIAL

—The holotype, FMNH

123917, is an adult female collected by David E.

Willard (original ﬁeld number DEW 425) on 18

October 1981 at Consuelo, km 165, 17 km by

road west of Pilcopata, Province of Paucartambo,

Department of Cuzco, Peru, approximately

S, 71

W. The type locality is in the

Cultural Zone of the Manu Biosphere Reserve.

The holotype is a specimen preserved in alcohol

with the skull removed and cleaned. The skull and

body were in good condition.

Paratypes include six adult specimens: one fe-

male (FMNH 172100) from San Pedro, Paucar-

tambo-Pilcopata road, Province of Paucartambo,

Department of Cuzco, Peru; one male (MVZ

166595) from Kiteni, Rı´o Urubamba, Province of

La Convencio´n, Department of Cuzco, Peru; two

females (MUSM 14562, 14566) from Campamen-

to Llactahuaman, Kimbiri, Province of La Con-

vencio´n, Department of Cuzco, Peru; and two

males (MUSM 18265, 18266) from Cordillera de

Carpish, Chinchao, Province of Hua´nuco, Depart-

ment of Hua´nuco, Peru. The holotype, paratypes,

and other specimens from the known distribution-

al range are listed in Appendix 1. Measurements

of each specimen of the type series of Platyrrhi-

nus masu are provided in Table 5.

ISTRIBUTION

—Platyrrhinus masu (Fig. 24) is

currently known from the eastern slope of the An-

des in Bolivia and Peru. The northern limit of the

distribution is the Department of Hua´nuco (Peru)

and the southern limit is the Department of La

Paz (Bolivia). The elevational range extends from

650 to 3350 m (Fig. 25).

TYMOLOGY

—From the Quechua word

‘‘masu,’’ meaning bat. Most of the collection lo-

calities of Platyrrhinus masu have Quechua as the

main language.

33VELAZCO: PLATYRRHINUS PHYLOGENY

. 23. Occlusal views of the left M1 and M2 illustrating taxonomic differences in the presence or absence of

a connection between the sulcus on the posterior face of the paracone on M1 and the cingulum of the lingual face

of the metacone on M1. (Top)Platyrrhinus ismaeli (MUSM 4946) not connected (arrow). (Bottom)Platyrrhinus

dorsalis (FMNH 128141) connected (arrow).

IAGNOSIS

—Size medium (FA

45–51 mm;

CIL

24.0–25.5 mm; W

23–33 g; Table 6);

two submental vibrissae on each side of the chin;

dorsal stripe brilliant-white and wide; dorsal hairs

on the back 6.3–7.5 mm; upper lateral incisors

bilobed.

ESCRIPTION

—A medium-sized species of Pla-

tyrrhinus (FA

45–51 mm; GLS

25.0–26.8

mm; Table 6). Dorsal fur dark brown, ventral fur

grayish, dorsal and ventral hair tricolored; dorso-

medial and ventrolateral facial stripes obviously

subequal and darker; dorsal hairs 6.3–7.5 mm

long on the back; narrow dorsal stripe brighter

that the facial ones, starting at the end of the dor-

somedial facial stripes and ending on the rump;

dorsal stripe deﬁnite but narrow; folds in the pin-

na well marked; six vibrissae surrounding the

margins of the noseleaf in a single array; two sub-

mental vibrissae present on each side of the chin;

one interramal vibrissae present; noseleaf longer

than wide; dense and long hair on the dorsum of

the feet; ‘‘U’’-shaped posterior edge of the uro-

patagium with a densely haired fringe, uropata-

gium extends along the midline 4.5–7 mm long;

insertion of the posterior edge of the plagiopata-

gium onto the ﬁrst metatarsal; third metacarpal

34 FIELDIANA: ZOOLOGY

ABLE

5. Measurements of the type series of Platyrrhinus masu.

Holotype

FMNH

123917

Paratype

FMNH

172100

Paratype

MVZ

166595

Paratype

MUSM

14562

Paratype

MUSM

14566

Paratype

MUSM

18265

Paratype

MUSM

18266

Sex

Weight

Total length

Hind foot length

Ear length

Female

—

Female

Male

Female

Male

Forearm length

Tibia length

Greatest length of skull

Condyloincisive length

Condylocanine length

19.58

26.03

24.87

24.39

19.03

26.42

24.77

24.19

20.41

26.14

25.52

24.87

19.13

26.70

25.40

24.69

19.28

26.84

25.13

24.54

20.15

26.53

25.35

24.87

19.63

26.41

25.51

25.04

Postorbital breadth

Zygomatic breadth

Braincase breadth

Mastoid breadth

Maxillary toothrow length

Breadth across molars

6.19

16.02

11.33

13.19

10.37

11.82

6.50

16.52

11.62

13.09

10.33

12.11

6.57

16.11

11.93

13.71

10.53

11.55

6.30

16.78

11.74

13.60

10.69

12.58

6.39

15.93

11.56

13.50

10.59

12.02

6.14

16.31

11.70

13.81

10.96

12.17

6.10

16.96

11.76

13.91

11.11

12.78

. 24. Frontal view of Platyrrhinus masu (FMNH 174760); adult male photographed at La Esperanza in the

Cultural zone of the Manu Biosphere Reserve, Cuzco, Peru, by B. D. Patterson.

shorter than or subequal to ﬁfth metacarpal; post-

orbital process absent or poorly developed; zy-

gomatic arches slightly divergent; paraoccipital

process well developed; upper median incisors

convergent and in contact; upper lateral incisors

bilobed; sulcus present on the posterior face of the

upper canines; two stylar cusps present on the

posterior face of P4; labial cingulum present at

35VELAZCO: PLATYRRHINUS PHYLOGENY

. 25. Distributional map of Platyrrhinus masu.

the base of the metacone on M1; sulcus present

on the posterior face of the M1 paracone; stylar

cusp present on the cingulum of the lingual and

labial face of the M1 metacone; M1 metastyle

present; moderately developed M1 protocone; sty-

lar cusp present on the lingual face of the M2

paracone; M2 metastyle present; lingual cingulum

of the M2 metacone restricted to the metacone;

M2 hypoconal basin developed; two sulci present

on the lingual face of the main cone of p4; two

cones present on the posterior face of p4; both

labial and lingual cingula present on p4; both la-

bial and lingual cingula present on m2; m1 para-

conid absent; m1 entoconid present; stylid cusp

on the anterior face of the m1 protoconid; m1

metaconid absent; m2 hypoconid present; stylid

cusp present between the metaconid and proto-

conid of m2; both labial and lingual cingula pre-

sent on m2. Velazco (2001) and Velazco and So-

lari (2003) did not ﬁnd secondary sexual dimor-

phism in either qualitative morphologic or mor-

phometric characters.

OMPARISONS

—Platyrrhinus masu is found

sympatrically with P. albericoi and P. nigellus in

several localities. Morphometrically, P. masu is

easily distinguished from P. albericoi, but there

is overlap in some measurements of masu and ni-

gellus:P. albericoi (FA

62–63 mm; CIL

31.6–32.6 mm; W

55–68 g), P. nigellus (FA

40–47 mm; CIL

21.9–24.5 mm; W

19–30

g), and P. masu (FA

45–51 mm; CIL

23.0–

25.5 mm; W

23–33 g). Platyrrhinus masu has

been traditionally confused with P. dorsalis; how-

ever, it also exhibits overlap of measurements

36 FIELDIANA: ZOOLOGY

ABLE

6. Selected measurements

a,b

of Platyrrhinus ismaeli and P. masu.

Platyrrhinus ismaeli Platyrrhinus masu

Weight

Total length

Hind foot length

37.1

5.2

30–51 (19)

86.7

5.3

78–98 (20)

15.5

1.6

13–18 (20)

27.0

2.7

23–33 (21)

74.3

3.2

70–82 (25)

13.6

0.9

12–16 (26)

Ear length

Forearm length

Tibia length

21.2

0.7

20–22 (19)

52.7

1.4

50–56 (20)

21.97

0.87

18.34–21.96 (20)

19.2

1.4

16–21 (25)

48.4

1.5

45–51 (26)

19.44

0.80

17.35–20.86 (26)

Greatest length of skull

Condyloincisive length

Condylocanine length

28.53

0.56

27.77–30.08 (20)

27.19

0.43

26.57–28.33 (20)

26.57

0.44

26.06–27.59 (20)

26.02

0.53

24.96–26.84 (26)

24.87

0.44

23.98–25.52 (26)

24.36

0.41

23.45–25.04 (26)

Postorbital breadth

Zygomatic breadth

Braincase breadth

6.48

0.20

6.17–6.86 (20)

17.21

0.49

16.25–18.21 (20)

11.97

0.23

11.57–12.29 (20)

6.33

0.18

5.99–6.61 (26)

16.13

0.34

15.62–16.96 (25)

11.54

0.21

11.19–11.93 (26)

Mastoid breadth

Maxillary toothrow length

Breadth across molars

14.07

0.30

13.43–14.72 (20)

11.84

0.34

11.33–12.48 (20)

13.08

0.46

12.40–13.83 (20)

13.46

0.22

13.09–13.91 (26)

10.57

0.22

10.17–11.11 (25)

12.05

0.28

11.55–12.78 (25)

Weight is in grams, all other measurements are in millimeters.

Summary statistics (mean and standard deviation [above], observed range and sample size [below]) of measure-

ments for each species (see Appendix 1 for a list of the specimens measured).

with P. ismaeli and P. nigellus (Tables 3, 6).

Therefore, comparisons will focus on differentia-

tion of these four species.

Externally, Platyrrhinus masu can be distin-

guished from these species by three submental vi-

brissae on each side of the chin in P. masu,

whereas P. dorsalis, P. ismaeli, and P. nigellus

have four submental vibrissae on each side of the

chin; inferior border of the noseleaf partially

joined to the upper lip in some individuals and

completely free in others in P. masu, whereas in

P. dorsalis, P. ismaeli, and P. nigellus, the infe-

rior border of the noseleaf is completely free in

all individuals; parallel folds in the pinna well

marked in P. masu and P. nigellus, whereas in P.

dorsalis and P. ismaeli, the parallel folds are

poorly marked but distinguishable; dorsal stripe

white and wide in P. masu and P. nigellus, where-

as in P. dorsalis and P. ismaeli, the dorsal stripe

is deﬁnite but narrow; dorsal fur length 6.3–7.5

mm in P. masu, whereas P. dorsalis, P. ismaeli,

and P. nigellus have the dorsal fur length

8 mm;

dense and long hair on the dorsum of the feet in

P. ismaeli, P. masu, and P. nigellus, whereas P.

dorsalis has the hair on the dorsum of the feet

intermediate in density and length; and fringe of

hair on the edge of the uropatagium densely

haired in P. ismaeli, P. masu, and P. nigellus,

whereas P. dorsalis has the uropatagium fringe

usually hairy, occasionally sparsely.

Dentally, Platyrrhinus masu can be distin-

guished from these species by the following char-

acters: upper lateral incisors bilobed in P. masu,

whereas P. dorsalis and P. nigellus have the upper

lateral incisors monolobed; P. dorsalis, P. masu,

and P. nigellus present the sulcus on the posterior

face of the paracone of M1 joined to the cingulum

of the lingual face of the metacone on M1, where-

as P. ismaeli present these two structures not

joined; M1 protocone moderately developed in P.

masu and P. nigellus, whereas P. dorsalis and P.

ismaeli have the M1 protocone small and blunt;

37VELAZCO: PLATYRRHINUS PHYLOGENY

ABLE

7. Measurements of the type species of Platyrrhinus matapalensis.

Holotype

FNH 81079 Paratype

FMNH 81080 Paratype

FMNH 81081 Paratype

MUSM 10725 Paratype

MUSM 10726

Sex Male Male Male Male Female

(pregnant)

Weight

Total length

Hind foot length

Ear length

Forearm length

—

Tibia length

Greatest length of skull

Condyloincisive length

Condylocanine length

Postorbital breadth

17.34

22.04

20.77

20.30

5.43

15.50

22.00

20.92

20.40

5.54

17.09

21.46

20.22

19.82

5.40

14.69

22.24

21.34

20.82

5.67

14.43

21.07

20.12

19.65

5.47

Zygomatic breadth

Braincase breadth

Mastoid breadth

Maxillary toothrow length

Breadth across molar

12.99

9.41

10.99

8.31

9.14

13.06

9.49

11.31

8.54

9.46

—

9.47

10.54

8.22

9.06

—

9.75

11.11

8.56

9.39

12.57

9.35

10.61

8.13

9.01

stylar cusp present on the lingual face of the M2

paracone in P. ismaeli and P. masu, whereas in

P. nigellus, that stylar cusp is lacking; lingual cin-

gulum of the M2 metacone restricted to the meta-

cone in P. ismaeli and P. masu, whereas in P.

dorsalis, the lingual cingulum of the M2 meta-

cone continues to the paracone; and lack of the

m1 metaconid in P. dorsalis and P. masu, where-

as in P. ismaeli, the m1 metaconid is poorly de-

veloped.

Platyrrhinus matapalensis new species. Figure

27 (top)

Vampyrops helleri Sanborn, 1955: 412 (part)

Vampyrops helleri Tuttle, 1970: 73 (part)

Vampyrops helleri Koopman, 1978: 12 (part)

Platyrrhinus helleri Rodriguez, 1998: 73

Platyrrhinus helleri Albuja, 1999: 139–140 (part)

Platyrrhinus helleri ‘‘Western’’ (this paper)

YPE

ATERIAL

—The holotype, FMNH 81079,

is an adult male collected by Celestino Kalinows-

ki (original ﬁeld number 1524) on 3 July 1954 at

Matapalo, 54 m in elevation, Province of Zaru-

milla, Department of Tumbes, Peru, approximate-

ly 3

S, 80

W. The skin and skull were

in good condition.

Paratypes include four adult specimens: two

males (FMNH 81080, 81081) from the type lo-

cality; and one female (MUSM 10726) and one

male (MUSM 10725) from Quebrada Naranjal-

Campo Verde, Z.R. Tumbes, 620–680 m in ele-

vation, Province of Zarumilla, Department of

Tumbes, Peru. All these specimens are adult and

are listed in Appendix 1 with other specimens

from the known distributional range. Measure-

ments of each specimen of the type series of Pla-

tyrrhinus matapalensis are provided in Table 7.

ISTRIBUTION

—Platyrrhinus matapalensis is

currently known from the western slope of the

Andes in Ecuador and Peru. The northern limit of

the distribution is the province of Esmeraldas (Ec-

uador) and the southern limit is the Department

of Tumbes (Peru). The elevational range extends

from 54 to 680 m (Fig. 26).

TYMOLOGY

—The speciﬁc name matapalensis

refers to the type locality Matapalo (Tumbes,

Peru), where the taxon was ﬁrst collected.

IAGNOSIS

—Size small (FA

37–39 mm; CIL

20.1–21.3 mm; W

16–20 g); two interramal

vibrissae; dorsal stripe deﬁnite but narrow;ven-

tral fur bicolored, occasionally unicolored;dense

and long hair on the dorsum of the feet.

ESCRIPTION

—A small-sized species of Platyr-

rhinus (FA

37–39 mm; GLS

21.1–22.2 mm).

Dorsal fur light brown, ventral fur brownish, dor-

sal hair tricolored, ventral hair bicolored in some

individuals and in others unicolored; dorsomedial

and ventrolateral facial stripes obviously subequal

and brighter, dorsal hairs 6.3–7.5 mm long on the

back; narrow dorsal stripe with the same colora-

tion as the facial ones; folds in the pinna poorly

marked but distinguishable; seven vibrissae sur-

rounding the margins of the noseleaf in a single

array; four submental vibrissae present; two inter-

ramal vibrissae present; noseleaf longer than

wide, inferior border of the horseshoe completely

38 FIELDIANA: ZOOLOGY

. 26. Distributional map of Platyrrhinus matapalensis.

free of the upper lip; hairs on the dorsum of the

feet sparse and short; ‘‘U’’ or ‘‘V’’ shaped pos-

terior edge of the uropatagium with a fringe

densely haired, uropatagium extends along the

midline

4.5 mm long; insertion of the posterior

edge of the plagiopatagium onto the ﬁrst metatar-

sal; ﬁfth metacarpal shorter than third metacarpal;

postorbital and paraoccipital processes poorly de-

veloped; fossa on the squamosal end of the zygo-

matic arch almost imperceptible; sulcus present on

the posterior face of the upper canines; two stylar

cusps present on the posterior face of P4; both la-

bial and lingual cingulum present at the base of the

metacone of M1; sulcus on the posterior face of

the paracone joined to the cingulum of the lingual

face of the M1 metacone; M1 parastyle present;

M1 metastyle occasionally present; M1 protocone

moderately developed; stylar cusp absent on the

lingual face of the M2 paracone; labial cingulum

of the M2 paracone present; M2 metastyle present;

stylar cusp absent on the lingual face of the M2

paracone; both labial and lingual cingula present

on p4; stylid cusps absent on the anterior or anter-

olingual face of the main cone of p4; stylid cusp

present on the posterior face of the main cone of

p4; m1 paraconid absent; m1 metaconid absent; m2

hypoconid absent; stylid cusp between the meta-

conid and protoconid of m2 present; both labial

and lingual cingula present on m2, but occasionally

only the labial cingulum is present.

OMPARISONS

—Platyrrhinus matapalensis is

found sympatrically with P. chocoensis and P.

dorsalis. Morphometrically, P. matapalensis is

easily distinguished from both as follows: P. cho-

coensis (FA

48–49 mm; CIL

24.6–26.2 mm;

29–32 g), P. dorsalis (FA

46–50 mm;

CIL

24.3–25.7 mm), and P. matapalensis (FA

37–39 mm; CIL

20.1–21.3 mm; W

16–

20 g). Platyrrhinus matapalensis has been con-

fused with P. helleri (Sanborn, 1955; Tuttle, 1970;

39VELAZCO: PLATYRRHINUS PHYLOGENY

Koopman, 1978); however, P. matapalensis could

also be mistaken for P. brachycephalus because

there is overlap in the measurements (Table 3, 7).

Therefore, the comparisons focus on P. matapa-

lensis, P. brachycephalus, and P. helleri because

these are the smaller species in the genus and are

subject to potential confusion.

Externally, Platyrrhinus matapalensis can be dis-

tinguished from the other two species by the follow-

ing: one vibrissa present on the upper lip of P. ma-

tapalensis, ventral to the vibrissae that surround the

margins of the noseleaf in, whereas P. brachyce-

phalus has two vibrissae on the upper lip; four sub-

mental vibrissae on each side of the chin in P. hel-

leri and P. matapalensis, whereas P. brachycephal-

us has ﬁve submental vibrissae on each side of the

chin; two interramal vibrissae present in P. mata-

palensis, whereas P. brachycephalus and P. helleri

have one interramal vibrissa; ventral fur unicolored

or bicolored in P. matapalensis, whereas P. brachy-

cephalus has the ventral fur tricolored and P. helleri

bicolored; sparse and short hair on the dorsum of

the feet in P. matapalensis, whereas in P. helleri,

the hair on the dorsum of the feet is intermediate in

density and length; and fringe of hair on the edge

of the uropatagium densely haired in P. helleri and

P. matapalensis, whereas P. brachycephalus has the

fringe of hair on the edge of the uropatagium usually

hairy, occasionally sparsely.

Cranially, Platyrrhinus matapalensis can be

distinguished from the other two species by the

following characters: a poorly developed postor-

bital process in P. brachycephalus and P. mata-

palensis, whereas in P. helleri, the postorbital pro-

cess is moderately developed; and paraoccipital

process almost imperceptible in P. brachycephal-

us and P. matapalensis, whereas in P. helleri, the

paraoccipital process is moderately developed.

Dentally, Platyrrhinus matapalensis can be dis-

tinguished from the other two species by the fol-

lowing: M1 protocone moderately developed in P.

matapalensis, whereas in P. brachycephalus, the

M1 protocone is well developed, and in P. helleri,

the M1 protocone is small and blunt, and lack of

stylid cusps on the anterior face of the main cone

of p4 in P. matapalensis, whereas P. brachyce-

phalus has two stylid cusps on the anterior face

of the main cone of p4, and P. helleri has only

one stylid cusp (Fig. 27).

Platyrrhinus nigellus (Gardner and Carter, 1972)

Vampyrops nigellus Gardner and Carter, 1972a. Type

locality: Huanhuachayo, Department of Ayacucho,

Peru.

OLOTYPE

—Vampyrops nigellus LSUMZ

16415, adult male, preserved as skin and skull,

was collected by A. L. Gardner (original ﬁeld

number ALG 11684) on 6 May 1971 at Huan-

huachayo, Department of Ayacucho, Peru, eleva-

tion 1660 m.

ISTRIBUTION

—Along the Andes from western

Venezuela south to Colombia, Ecuador, Peru, and

Bolivia, from 620 to 2757 m (Fig. 28).

IAGNOSIS

—Size medium (FA

40–47 mm;

CIL

21.9–24.5 mm; W

19–30 g; Table 3);

facial stripes darker; absence of a basal protuber-

ance, where the genal vibrissae are implanted;

postorbital process moderately developed; paraoc-

cipital process moderately developed; a deep fos-

sa on the squamosal end of the zygomatic arch,

lateral to the glenoid fossa; m1 metaconid poorly

developed.

Platyrrhinus reciﬁnus (Thomas, 1901)

Vampyrops reciﬁnus Thomas, 1901. Type locality:

Recife, State of Pernambuco, Brazil.

OLOTYPE

—Vampyrops reciﬁnus BMNH

81.2.16.4, adult male, preserved in alcohol with

the skull removed and cleaned, was collected by

W. A. Forbes at Recife, State of Pernambuco, Bra-

zil.

ISTRIBUTION

—Eastern Brazil, elevation 200 m

(Fig. 28).

IAGNOSIS

—Size small to medium (FA

42–

43 mm; CIL

22.0–22.6 mm; W

17–19 g;

Table 3); interramal vibrissae absent;dorsal fur

tetracolored; dense and long hair on the dorsum

of the feet; an almost imperceptible fossa on the

squamosal end of the zygomatic arch, lateral to

the glenoid fossa;a deep fossa on the hypoconal

basin of P4;stylid cusp absent between the meta-

conid and the protoconid of m2.

Platyrrhinus vittatus (Peters, 1860)

Artibeus vittatus Peters, 1860. Type locality: Puerto

Cabello, State of Carabobo, Venezuela.

OLOTYPE

—Artibeus vittatus ZMB 568, adult

male, preserved in alcohol with the skull removed

and cleaned, was collected by von Appun at

Puerto Cabello, State of Carabobo, Venezuela.

ISTRIBUTION

—Costa Rica, Panama, western

and northern Colombia, and northern Venezuela,

from 640 to 1400 m (Fig. 28).

IAGNOSIS

—Size large (FA

57–61 mm; CIL

28.1–31.3 mm; W

60–65 g; Table 3); par-

40 FIELDIANA: ZOOLOGY

. 27. Labial views of the left p2 and p4 illustrating taxonomic differences in the number of stylid cusps on

the anterior face of the main cone of p4. (Top)Platyrrhinus matapalensis (FMNH 81079) lack stylid cusps (arrow).

(Center)Platyrrhinus helleri (FMNH 127127) with one stylid cusp present (arrow). (Bottom)Platyrrhinus brachy-

cephalus (FMNH 139584) with two stylid cusps present (arrow).

41VELAZCO: PLATYRRHINUS PHYLOGENY

. 28. Distributional map of Platyrrhinus nigellus, P. reciﬁnus, and P. vittatus.

allel folds in the pinna well marked; dorsal stripe

brilliant-white and wide; fringe of hair on the

edge of the uropatagium usually hairy, occasion-

ally sparsely;paraoccipital process moderately

developed; one stylid cusp present on the antero-

lingual face of the main cone of p4. In older in-

dividuals (AMNH 15100–15101, 34232; ZMB

568 [Holotype]) with a moderate degree of tooth

wear, this stylid cusp appears as a nub on the an-

terolingual face of the main cone.

OMMENTS

—In his systematic review of Pla-

tyrrhinus from northwestern South America, Al-

berico (1990) identiﬁed Platyrrhinus sp.Basa

potentially new species from Colombia. Platyr-

rhinus sp.B(FA

56.9–59.4 mm) was collected

only at the northern end of the Andes in Colom-

bia, in the Department of Antioquia at 500 m; this

taxon shows the same range of variation in FA

size as P. vittatus (sensu this paper) and was

found within the geographic range of P. vittatus.

Some populations of P. vittatus (sensu Alberico,

1990) belong to P. albericoi, suggesting that Pla-

tyrrhinus sp. B is conspeciﬁc with P. vittatus.

From the three potentially new species identi-

ﬁed by Alberico (1990), two are conspeciﬁc with

Platyrrhinus ismaeli and P. vittatus; the third

probably constitutes a new species. Platyrrhinus

sp.A(FA

53.0–57.0 mm) is found along the

western base of the western Andes of Colombia

from the department of Choco´, south almost to

the Department of Narin˜o, from 230 to 1000 m in

elevation. Based on both geographical range and

FA size, this taxon does not overlap with any of

the other 14 Platyrrhinus taxa.

42 FIELDIANA: ZOOLOGY

Commonly, between 8 and 10 nominal species

are recognized for Platyrrhinus (Alberico, 1990;

Ferrell & Wilson, 1991; Jones & Carter, 1976,

1979; Koopman, 1993, 1994; Nowak, 1999;

Owen, 1987; Swanepoel & Genoways, 1979).

Based on the results obtained in this work, the

genus Platyrrhinus contains at least 14 species.

Therefore, it currently ranks as the most speciose

genus in the family Phyllostomidae.

Acknowledgments

For the loan of specimens or for their hospital-

ity while visiting their respective institutions, I

thank Luis Albuja (EPN), Michael D. Carleton

(USNM), Judy Chupasko (MCZ), Luis Coloma

(QCAZ), Joseph A. Cook and William L. Gannon

(MSB), Judith Eger and Burton Lim (ROM),

Mark S. Hafner (LSUMZ), Philip Myers

(UMMZ), Vı´ctor Pacheco (MUSM), Bruce D.

Patterson (FMNH), James L. Patton (MVZ), and

Nancy B. Simmons and Robert S. Voss (AMNH).

The visits to the AMNH and FMNH were sup-

ported by the AMNH Collection Study Grant

(AMNH) and the Barbara E. Brown Fund for

Mammal Research (FMNH). I am also very grate-

ful to Robert Asher (ZMB), who kindly took some

pictures of the holotype of Platyrrhinus vittatus.

Betty Strack (FMNH) assisted with the SEM pho-

tography. Sean Bober (FMNH) shared his exper-

tise in GIS.

Useful comments on various drafts of this man-

uscript were provided by Christine L. Hice,

Burton K. Lim, Victor Pacheco, Bruce D. Patter-

son, and two anonymous reviewers.

I am very grateful to the Departamento de Mas-

tozoologı´a at the Museo de Historia Natural

(UNMSM) and the Division of Mammals of the

Field Museum of Natural History for providing

space, technical, and scientiﬁc support.

During the preparation of this manuscript, I was

supported by a curatorial assistantship funded by

a NSF grant (DEB-9870191) to Bruce D. Patter-

son, D. F. Stotz, and J. W. O. Ballard and by the

Barbara E. Brown Fund for Mammal Research.

Part of this work was submitted to the Universi-

dad Nacional Mayor de San Marcos, Lima, Peru,

in partial fulﬁllment of the requirements of my

Magister degree.

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ELASCO

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Appendix 1: Specimens Examined

The following list includes all specimens ex-

amined in this study, with their respective locali-

ties. See Materials and Methods for abbreviations.

Individuals or series marked with an asterisk were

used in the elaboration of the Tables 2 to 7.

Platyrrhinus albericoi:Ecuador: Pichincha,

Nanegal, Gavila´n de Orongo (QCAZ 1979, 2003);

Peru: Ayacucho, Yuraccyacu (MUSM 993); Cuz-

co, La Convencio´n, Kimbiri, Camp. Wayrapata

(MUSM 14599); Cuzco, La Convencio´n, Kimbiri,

Camp. Llactahuaman (FMNH 177477, MUSM

14583–14584, 14586, 14588–14589); Cuzco,

Paucartambo, San Pedro (MUSM 19149*); Cuz-

co, Paucartambo, Suecia, km 138.5 Carretera

Shintuya (FMNH 170145*); Cuzco, Paucartambo,

Pillahuata (FMNH 172108); Pasco, Oxapampa,

Pozuzo, Palmira (MUSM 10973).

Platyrrhinus aurarius:Guyana: Potaro-Sipa-

runi, Mount Ayanganna, Toe Slope Camp (ROM

114679*); Potaro-Siparuni, Mount Ayanganna,

First Plateau Camp (ROM 114702*); Venezuela:

Amazonas, Cerro de la Neblina, Base del Pico

Maguire (FMNH 137294–137298*, 137299–

137303, 137304–137307*, 137308–137311);

Amazonas, Cerro de la Neblina, Left Bank del

Rı´o Baria (FMNH 137312, 137313*); Amazonas,

km W Pico Zuloaga (AMNH 261225–

261229); Bolı´var, Cerro Guaiquinima (AMNH

235339–235342); Bolı´var, La Escalera (AMNH

265572–265584).

Platyrrhinus brachycephalus:Bolivia: Cocha-

bamba, 4 km SE Villa Tunari (UMMZ 126755);

El Beni, Itenez, Buenavista (FMNH 115012*); El

Beni, Mamore, San Joaquin (FMNH 96092*,

115016–115018*); El Beni, Vaca Diez, La Esper-

anza (FMNH 115003*, 115005*); La Paz, Santa

Ana de Madidi (AMNH 262353); Pando, Santa

Rosa (AMNH 262520); Brazil: Amazonas, Ta-

batinga (AMNH 170623); Colombia: Arauca,

Rı´o Arauca (FMNH 92302–92303, 92326*,

92327); Meta, La Macarena Parque, Refugio

(FMNH 58739*); Meta, Villavicencio (AMNH

207872–207873, 207875); Meta, Villavicencio,

Finca El Buque (FMNH 121261*); Putumayo, Es-

tacio´n de Bombeo, Guamues (FMNH 113657);

Putumayo, San Antonio, Rı´o Guamues (FMNH

114018–114020); Ecuador: Napo, Marian

(FMNH 124990*); Napo, San Jose´ (AMNH

64033); Napo, San Jose de Payamino (FMNH

124991–124992*); Napo, Zancudo (FMNH

124985–124986*); Orellana, Canto´n Aguarico,

Pozo Exploratorio PCSA-2 (EPN 985199,

985201); Pastaza, Sarayacu (AMNH 67649);

French Guiana: Cayenne (FMNH 21745–

21746); Peru: Amazonas, Condorcanqui, Santa

Rosa de Nieva (MUSM 12789); Amazonas, San

Juan (Bagua Grande) (MUSM 1010); Cuzco, La

Convencio´n, Camisea, Armihuari (MUSM 13770,

13774–13775, 13778–13779, 13786, 13808);

Cuzco, La Convencio´n, Camisea, Cashiriari

(MUSM 13788–13790, 13821, 13833); Cuzco, La

Convencio´n, Kiteni (MUSM 1012; UMMZ

160637); Cuzco, La Convencio´n, Camisea, Kon-

kariari (MUSM 14788); Cuzco, La Convencio´n,

Camisea, Pagoreni (MUSM 13791–13792); Cuz-

co, La Convencio´n, Camisea, San Martı´n (MUSM

13793–13794, 13844); Cuzco, La Convencio´n,

Camisea, Segakiato (MUSM 14793); Cuzco, Pau-

cartambo, Consuelo, km 165, 17 km by road W

Pilcopata (FMNH 123920*, 123923*); Cuzco,

Quispicanchi, Collpa de San Lorenzo (FMNH

93580*, 93582–93585*); Cuzco, Quispicanchi,

Huajyumbe (FMNH 84381*); Hua´nuco, Leoncio

Prado, Tingo Maria, Rı´o Azul (FMNH 98008*);

Loreto, Alto Amazonas, Rı´o Morona, Quebrada

Pushaga (FMNH 89091*); Loreto, Alto Amazon-

as, Yurimaguas (FMNH 19651*); Loreto,Quebra-

da Sucusari, Camp. Explornapo (MUSM 6750);

Loreto, Requena, Jenaro Herrera (MUSM 5927);

Loreto, Loreto, Rı´o Samiria, Base Atun (FMNH

122874–122876*, 122878*); Loreto, Loreto, Rı´o

Tigre, 1 km above Rı´o Tigrillo (FMNH 122880*);

Loreto, Mariscal Ramon Castilla, Rı´o Amazonas,

mouth of Rı´o Peruate (FMNH 89096*); Loreto,

Mariscal Ramon Castilla, Rı´o Yavari Mirim, Que-

brada Esperanza (FMNH 89095*); Loreto, May-

nas, Rı´o Nanay, Santa Luisa (FMNH 87074*);

Madre de Dios, Albergue Rı´o Madre de Dios, 12

km E Puerto Maldonado (MUSM 1013); Madre

de Dios, Manu, Alto Rı´o Madre de Dios, Haci-

46 FIELDIANA: ZOOLOGY

enda Amazonia (FMNH 139639*, FMNH

125930*, 125938*; MUSM 8850); Madre de

Dios, Hacienda Erika, Rı´o Alto Madre de Dios,

opposite Salvacio´n (UMMZ 160636, 160638);

Madre de Dios, Reserva Cuzco Amazo´nico, 15

km NE Puerto Maldonado (MUSM 6268–6270,

7150); Madre de Dios, Manu, Pakitza (MUSM

6769, 12558, 12560, 12562, 12564); Madre de

Dios, Manu, Rı´o Palotoa (MUSM 9844); Madre

de Dios, Manu, Rı´o Palotoa, left bank, 12 km up-

stream from mouth (FMNH 139581*, 139584–

139585*, 139587*); Madre de Dios, Rı´o Tambo-

pata (MUSM 191); Madre de Dios, Rı´o Tambo-

pata, Explorer’s Inn (MUSM 1002, 1006–1007);

Pasco, Oxapampa, Huancabamba, Comunidad

Nativa Castillo (MUSM 394–396, 398, 400, 430,

496–499); Puno, Carabaya, Coasa, mouth of Que-

brada Ursulinda with the Rı´o Candamo (MUSM

15882); Ucayali, Coronel Portillo, Pucallpa

(FMNH 64316–64317*); Ucayali, Coronel Porti-

llo, Yarinacocha (FMNH 98009*); Ucayali, Padre

Abad, B. N. von Humboldt (MUSM 8527); Su-

riname: Nickerie, Wageningen (UMMZ 175688);

Paramaribo, Peu Et Contant (UMMZ 175768);

Venezuela: Tachira, San Juan de Colon (FMNH

21117*).

Platyrrhinus chocoensis:Colombia: Cauca,

Alto Micay, Betania (FMNH 113745*, 113822–

113823, 113825–113927, 113830–113832); Cau-

ca, Quebrada Guangui (AMNH 235774–235779);

Choco´, Quebrada Docordo (AMNH 233186–

233187); Valle del Cauca, Concesio´n Bajo Cali-

ma, Cuartel BV83 (FMNH 140696–140697*);

Valle del Cauca, Zabaletas (FMNH 85838;

UMMZ 169039–169040, 169048); Ecuador: Es-

meraldas, Borbo´n, Rı´o Cayapas, Community of

Zapote (QCAZ 2063); Esmeraldas, Borbo´n, Rı´o

Cayapas, Community of Viruela (QCAZ 2175);

Esmeraldas, San Lorenzo, Estacio´n La Chiquita

(QCAZ 2375, 2378, 3281–2382); Esmeraldas, 3

km S San Miguel (EPN 84361).

Platyrrhinus dorsalis:Colombia: Cauca, Alto

Micay, Betania (FMNH 113372, 113380, 113385,

113394–113396, 113821*, 113824*, 113828–

113829*, 113831*, 113833–113835*); Cauca,

Charguayaco (FMNH 113538–113539*); Cauca,

Popaya´n (FMNH 90327*); Huila, Las Cuevas

Parque, Upper Cabana (FMNH 58740*); La Gua-

jira, San Antonio (MCZ B-8300); Magdalena,

Palomino (MCZ B-8301); Valle del Cauca, El Sil-

encio (UMMZ 169038); Ecuador: Carchi, Pailo´n,

NW Parroquia de Chical (EPN 871732, 871748,

871778, 871843–871848, 871868); Carchi, Par-

roquia Tobar Donoso, El Pailo´n (EPN 871747,

871765); Esmeraldas, Borbo´n, Rı´o Cayapas, Rı´o

Chimbocal, Comuna Corriente Grande (QCAZ

2157); Esmeraldas, Eloy Alfaro, Pichiyacu, Rı´o

Cayapas, upstream Borbo´n (QCAZ 1468); Pichin-

cha, Nanegal (QCAZ 1975); Pichincha, Tambillo,

25 km S Quito (EPN 7465, 7467); Panama: Da-

rien, Parque Nacional Darien, Rancho Frio

(FMNH 128141*).

Platyrrhinus helleri:Belize: Cayo district, Ba-

nana Bank (FMNH 58264*); Cayo district, Ba-

nana Bank Ranch (FMNH 58265–58268*); Cayo

district, Macaw Bank (FMNH 106804*); Toledo

district, Columbia Forest Preserve, Forestry Camp

(FMNH 58270*); Toledo district, Columbia For-

est Preserve, Mahogany Plant, N Forestry Camp

(FMNH 58269*); Toledo district, Forest Home

(FMNH 128073*); Bolivia: Beni, 6 km W Cas-

arabe (AMNH 255919, 255921–255922, 255927);

Beni, Gral. Jose´ Ballivia´n, Serrania Pilon, 27 km

NRı´o Quiquibey by road to San Borja (AMNH

261058, 261062, 261064–261069, 261071–

261074); Beni, Itenes, Buenavista (FMNH

115001*, 115009–115011*, 115012, 115013*);

Beni, Mamore, San Joaquı´n (FMNH 96092,

115002, 115006, 115007–115008*, 115016–

115018, 115019*); Beni, opposite Costa Marques,

Brazil (AMNH 209502); Beni, Puerto Almace´n

(AMNH 255909, 255912); Beni, Vaca Diez, La

Esperanza (FMNH 115003–115005, 115014*);

Cochabamba, 3 km SW Villa Tunari (AMNH

244629–244630); Cochabamba, 50 km NW Villa

Tunari (UMMZ 126754, 126756); La Paz, Santa

Ana de Madidi (AMNH 261628–261631,

261634–261639, 261642–261647, 262351,

262354–262355); La Paz, Sararia, 2 hr (river)

NW Puerto Linares (UMMZ 126757–126758);

Pando, La Cruz (AMNH 262507); Pando, Las

Piedras (AMNH 262514); Pando, Rı´o Nareuda

(AMNH 248880, 249062); Pando, Santa Rosa

(AMNH 262518, 262522–262523); Pando, W

Bank Rı´o Beni, opposite Hamburgo (AMNH

262510) ; Santa Cruz, 23 km S Camp Los Fierros,

Parque Nacional Noel Kempff Mercado (AMNH

264068–264070); Santa Cruz, 4.5 km N Buen Re-

tiro (AMNH 260228); Santa Cruz, Camp Los

Fierros (AMNH 263616); Santa Cruz, Estancia

Cachuela Esperanza (AMNH 260219–260221,

260223–260224, 260226–260227); Santa Cruz,

San Rafael de Amboro (AMNH 261663, 262350);

Brazil: Para´, Bele´m, Mocambo/Embrapa (FMNH

126601*, 126602, 126603*, 126604–126605,

126606*); Rondoˆnia, Pedras Negras (AMNH

209517); Colombia: Cauca, Alto Micay, Betania

(FMNH 113321, 113330, 113344, 113348,

47VELAZCO: PLATYRRHINUS PHYLOGENY

113883–113884, 113885*, 113886, 113887–

113889*, 114023–114026); Cauca, Quebrada

Guangui (AMNH 235780–235788); Cauca, Rı´o

Saija, Quebrada Huanqui (FMNH 104841,

104843); Choco´, Quebrada Docordo (AMNH

233188); Cundinamarca, Mesitas de Colegio

(AMNH 207863, 207871); Cundinamarca, Sasai-

ma (FMNH 49156*); Magdalena, Cacagualito

(FMNH 13202*); Meta, Villavicencio (FMNH

121262*); Valle del Cauca, Rı´o Zabaletas

(UMMZ 168982–168984, 168987, 168993–

168995, 168999); Costa Rica: Alajuela, Guayabo

(FMNH 18045–18046*); Guanacaste, Guanacaste

2 km S, 12 km E Bolson (FMNH 123152); Pun-

tarenas, 2 km S San Vito, Finca Las Cruces

(FMNH 123153); Puntarenas, Dominical (UMMZ

112024, 112031); Ecuador: Napo, Estacio´n La

Selva, 85 km SE Coca (QCAZ 2518–2519);

Napo, Orellana, Comuna Indillana, mouth of Rı´o

Indillana and Rı´o Napo, P. N. Yasunı´ (QCAZ

969); Pastaza, Rı´o Cushueme, Cushueme, 150 km

SE Puyo (FMNH 104753, 104759); Guatemala:

Alta Vera Paz, Los Ra´pidos (AMNH 214242); Pe-

ten, Tikal National Park (FMNH 58584*); Santa

Rosa, Avellana (AMNH 243774, 243923); Santa

Rosa, 13 km N Avellana, Finca Camalote

(AMNH 243775); Santa Rosa, 2 km ESE Ixpaco

(AMNH 244337); Santa Rosa, 5 km N Avellana,

Chiquihuitan (AMNH 245325); Honduras: Cor-

tes, Santo Domingo, aprox. 5.5 km ESE Cuyamel

(AMNH 265119–265120); Lempira, Tepusuna

(FMNH 47615–47616*); Mexico: Oaxaca, Tutla

(FMNH 51856*); Veracruz, Tuxtla Mts (FMNH

127127–127130*); Veracruz, Tuxtla Mts, 0.5 km

E cerro Balzapote (FMNH 127115–127120*,

127125–127126*, 127131–127132*, 127134–

127136*); Veracruz, Tuxtla Mts, 0.5 km W cerro

Balzapote (FMNH 127121–127124*); Panama:

Darie´n, Cana (LSUMZ 25485–25486*); Peru:

Amazonas, Bongara, Rı´o Utcubamba, between

Churuja and Pedro Ruiz (FMNH 129149–

129150*; MUSM 4950–4952); Amazonas, Luya,

Rı´o Utcubamba, 11 km by road NW Pedro Ruiz

(FMNH 129152–129153*); Cuzco, La Conven-

cio´n, Camisea, Armihuari (MUSM 13799, 13803,

13807); Cuzco, La Convencio´n, Camisea, Cashi-

riari (MUSM 13811, 13824–13825); Cuzco, La

Convencio´n, Camisea, Las Malvinas (MUSM

14794); Cuzco, La Convencio´n, San Martı´n

(MUSM 13846–13847, 13849); Cuzco, Quispi-

canchi, Collpa de San Lorenzo (FMNH 93586–

93587*); Junı´n, Alto Yurinaqui (MUSM 1009);

Loreto, Mariscal Ramon Castilla, Rı´o Yavari Mir-

im, Quebrada Esperanza (FMNH 89094*); Lore-

to, Maynas, Rı´o Nanay, Santa Luisa (FMNH

87080*); Loreto, Requena, Jenaro Herrera

(MUSM 871, 4217, 5526, 5598, 5928); Loreto,

Requena, Jenaro Herrera, 1.4 km N Centro de In-

vestigacio´n Jenaro Herrera (MUSM 5500); Lore-

to, Rı´o Yarapa, Albergue Yacumana, SE Iquitos

(MUSM 9434); Madre de Dios, Albergue, Rı´o

Madre de Dios, 12 km E Puerto Maldonado

(MUSM 1008); Madre de Dios, Manu, Aguas

Calientes, Rı´o Alto Madre de Dios, 1 km below

Shintuya (UMMZ 160635); Madre de Dios,

Manu, Hacienda Amazonia (FMNH 125951*,

139631*; MUSM 8855, 9863–9864); Madre de

Dios, Manu, Pakitza (MUSM 6768, 12555–

12557, 12559, 12563, 12565–12575); Madre de

Dios, Manu, Rı´o Palotoa (MUSM 9884); Madre

de Dios, Manu, Rı´o Palotoa, left bank, 12 km up-

stream from mouth (FMNH 139648*); Madre de

Dios, Rı´o Tambopata, Explorer’s Inn (MUSM

1003); Pasco, Oxapampa, Pozuzo (MUSM 10942,

10946); Pasco, Villa Ame´rica (MUSM 1005);

Ucayali, Padre Abad, Bosque Nacional Alexander

von Humboldt (MUSM 8633–8636, 8528–8531);

Trinidad and Tobago: Saint George, Arima Val-

ley (AMNH 149624); Saint George, Chaguara-

mas, U.S. Naval Station (AMNH 183164); Saint

George, Maracas Valley, Water Fall road (AMNH

176283); Saint George, Port of Spain, Saint Clair,

2 Scott (AMNH 176284); Saint George, 4 mi by

road N Arima (AMNH 246222–246224); Saint

Patrick, Point Fortin (AMNH 183858–183859);

Venezuela: Aragua, Rancho Grande (AMNH

144386); Guarico, 8 km N 13 km W San Jose´de

Guaribe (AMNH 247644); Tachira, San Juan de

Colo´n (FMNH 20539–20540*); Zulia, 9 km N

Rio Catatumbo (AMNH 244039); Zulia, Puerto

Delicias (AMNH 244038).

Platyrrhinus infuscus:Bolivia: Beni, Gral. Jose´

Ballivia´n, Serranı´a Pilo´n, 27 km by road N Rı´o

Quiribay (AMNH 261076); Cochabamba, 4 km

SE Villa Tunari (UMMZ 126761); La Paz, 6.6 km

downstream Canavari, Valle del Rı´o Coroico

(AMNH 246621); La Paz, Serrania Bellavista, 35

km N Canavari (AMNH 246622–246623); Pando,

Rı´o Nareuda (AMNH 248882–248883); Santa

Cruz, 3 km N and 13.5 km W San Rafael de Am-

boro, Rı´o Saguayo (AMNH 261623); Santa Cruz,

San Rafael de Amboro (AMNH 261664); Colom-

bia: Meta, La Macarena Parque, Camp. Izawa

(FMNH 58741*, 58745*); Meta, La Macarena

Parque, Cano, Refugio Cabana (FMNH 58742–

58743*, 58744); Meta, La Macarena Parque,

Cano Cabana, Cabana Duda (FMNH 58746*);

Meta, La Macarena Parque, Refugio (FMNH

48 FIELDIANA: ZOOLOGY

58747*); Meta, Villavicencio, Villavicencio

(FMNH 51733*); Meta, Villavicencio, Small

Cano (FMNH 58748); Putumayo, Estacio´n de

Bombeo, Guamues (FMNH 113699, 113752,

113898); Putumayo, San Antonio, Rı´o Guamues

(FMNH 113413, 113415, 113417, 113899–

113900, 114128–114129); Putumayo, Rı´o Me-

caya (FMNH 71525, 72123*); Ecuador: Napo,

Loreto, P. N. Napo-Galeras, W side, line 28

(QCAZ 1533–1534); Napo, Loreto, P. N. Napo-

Galeras, W side, line 30 (QCAZ 1406–1407);

Napo, Rı´o Cotapino, Oriente (FMNH 47587*);

Napo, Rı´o Suno (AMNH 67929); Napo, San Jose´

(AMNH 67907–67912, 67918–67919, 67922,

67925–67926); Napo, San Jose´ de Payamino

(FMNH 124989*); Pastaza, Montalvo, (FMNH

41429*); Pastaza, Rı´o Copotaza (FMNH 53503*);

Pastaza, Rı´o Pindo Yaco (FMNH 43137–43139*);

Sucumbı´os, Gonzalo Pizarro, Los Cedros,

Bosque, 10 km S Lumbaquı´ (QCAZ 521–523);

Tungurahua, Ban˜os, Oriente (FMNH 47588*);

Tungurahua, Palmera (AMNH 67658–67664);

Peru: Amazonas, 43 km NE Chiriaco (MUSM

143); Amazonas, Luya, Rı´o Utcubamba, 15 km

by road NW Pedro Ruiz (FMNH 129158–

129159*, 129161*; MUSM 4953–4954); Cuzco,

La Convencio´n, Camisea, Armihuari (MUSM

13850); Cuzco, La Convencio´n, Camisea, Pago-

reni (MUSM 13858–13859); Cuzco, La Conven-

cio´n, Camisea, San Martı´n (MUSM 13862); Cuz-

co, La Convencio´n Camisea, Segakiato (MUSM

14798); Cuzco, Paucartambo, Consuelo, km 165,

17 km by road W Pilcopata (FMNH 123931–

123932*; MUSM 9923–9924); Cuzco, Paucar-

tambo, Tono, 5 km S Rı´o Tono and 18 km W

Patria (MUSM 9950); Cuzco, Quispicanchi, Coll-

pa de San Lorenzo (FMNH 93545*); Cuzco,

Quispicanchi, Hacienda Cadena (FMNH 68447*,

68449*, 78691–78692*, 78694*); Cuzco, Quis-

picanchi, Huajyumbe (FMNH 68450–68454*,

75153–75154*, 78693*, 84406*); Cuzco, Quis-

picanchi, San Juan Grande (FMNH 75155–

75156*); Junı´n, Alto Yurinaqui (MUSM 994); Ju-

nin, Chanchamayo, Tarma, Vitoc Valley (FMNH

51525*); Loreto, Mouth of Rı´o Curaray (AMNH

71696–71701, 71708); Loreto, Loreto, Rı´o Tigre,

1 km above Rı´o Tigrillo (FMNH 122902*); Ma-

dre de Dios, Hacienda Erika, Rı´o Alto Madre de

Dios opposite Salvacio´n (UMMZ 160628–

160629); Madre de Dios, Manu, Rı´o Alto Madre

de Dios, Hacienda Amazonia (FMNH 125990*,

126010*, 126019*, 139663–139665*, 139674*,

139676*, 139685*, 139695–139696*, 139700–

139702*, 139710*; MUSM 994, 9847, 9897,

9902–9904, 9910, 9928); Madre de Dios, Manu,

Aguas Calientes, Rı´o Alto Madre de Dios, 1 km

below Shintuya (UMMZ 160630–160632); Madre

de Dios, Manu, Cerro de Pantiacolla (FMNH

122134*, 139737*, 139739–139742*, 139745*,

139748–139749*; MUSM 9918, 9920–9921);

Madre de Dios, Manu, Itahuania (FMNH

84405*); Madre de Dios, Manu, Pakitza (MUSM

12576); Madre de Dios, Reserva Cuzco Amazo´n-

ico, 15 km NE Puerto Maldonado (MUSM 6272–

6273); Pasco, Oxapampa, Pozuzo (MUSM 10960,

10969); Pasco, Oxapampa, Pozuzo, Palmira

(MUSM 10941, 10947); Puno, Carabaya, Coasa,

mouth of Quebrada Ursulinda with Rı´o Candamo

(MUSM 15824); Ucayali, Afﬂuent of Mamanchi-

ta (MUSM 370).

Platyrrhinus ismaeli:Colombia: Huila, Las

Cuevas Parque, Cueva India (FMNH 58733–

58735*, 58737–58738*); Huila, Cuevas de los

Guacharos National Park, 225 m W Upper Ca-

ban˜a (IND 1993*); Huila, Cuevas de los Guacha-

ros National Park, Entrance to Indian Cave (IND

2244*); Huila, Cueva de Los Guacharos National

Park, on top Guacharos Cave (IND 2308*); Huila,

Cuevas de los Guacharos National Park, Upper

Bridge on Rı´o Suaza (FMNH 58732*, 58736*);

Ecuador: Azuay, Hacienda Sector de Challtapac,

close to Rı´o Giro´n (QCAZ 2237); Loja, Loja, San

Pedro de Vilcabamba, 3 km N Vilcabamba

(QCAZ 1171–1174, 1177, 1180); Morona Santi-

ago, San Jose´ Alto, Canto´n Paute, Rı´o San Vi-

cente (EPN 912940); Napo, Loreto, P. N. Napo-

Galeras, W side, line 28 (QCAZ 1291); Napo, San

Rafael Cascada (FMNH 124988*); Peru: Ama-

zonas, 19 km E Balsas (FMNH 129133–129134*,

129136–129137*; MUSM 4944–4945, 4946*);

Cajamarca, 12 km SSW by road to San Miguel

(MUSM 4949); Cajamarca, Celendin, Hacienda

Limo´n, W Balsas (FMNH 129139*, 129143*,

129145–129146*; MUSM 4947–4948); Cajamar-

ca, San Ignacio (MUSM 12884–12889); Lamba-

yeque, Ferren˜afe, Bosque Chin˜ama (MUSM 925–

926); Piura, Ayabaca, ca. 44 km ESE by road

from Ayabaca, ladera Cerro Mayordomo (MUSM

996–998); San Martı´n, Huallaga (MUSM 16167–

16169); San Martı´n, La Playa, 28 km NE Pataz

(MUSM 7283); San Martı´n, Las Palmas, 32 km

NE Pataz (MUSM 7285–7293); San Martı´n, Las

Papayas (MUSM 7294).

Platyrrhinus lineatus:Bolivia: Beni, Rı´o Ma-

more´ (AMNH 210805–210807, 210809); Beni,

Rı´o Tijamuchi (AMNH 262352); Santa Cruz, 4

km N and 1 km W Santiago de Chiquitos (AMNH

260231); Santa Cruz, 12 km S and 8 km E Santa

49VELAZCO: PLATYRRHINUS PHYLOGENY

Cruz (AMNH 255931–255932); Santa Cruz, Cer-

cado, Cercado (FMNH 50990*); Santa Cruz, Chi-

quitos, Santiago (FMNH 105844–105862*,

105899–105908, 105909*, 105910–105913);

Santa Cruz, Chiquitos, 3 km S and 3 km W San-

tiago de Chiquitos (AMNH 261042–261052,

261054–261056); Santa Cruz, Robore´ (AMNH

260232–260233); Santa Cruz, Santa Rosa de la

Roca (AMNH 263618); Brazil: Bahia, Municı´pio

de Valenc¸a, Guaibim (MVZ 185601); Ceara´, Serra

de Ibiapaba (FMNH 19516); Espı´rito Santo, Mun-

icı´pio de Santa Teresa, Estac¸a˜o Biolo´gica Santa

Lu´cia (MVZ 185602); Mato Grosso do Sul, Bel-

vedere de Urucum de Corumba´ (AMNH 37019–

37021); Mato Grosso do Sul, Maracaju´ (AMNH

134923–134926); Mato Grosso do Sul, Rio Va-

caria, Fazenda Capa˜o Bonito (FMNH 47963–

47966, 47968–47977, 47980, 47983–47994,

48008); Mato Grosso do Sul, Urucum (AMNH

36993–37002, 37004–37005, 37007–37018,

39017–39019, 39026–39029; FMNH 26775,

26777–26781*, 26783–26784*, 30032–30048);

Paraı´ba, Joa˜o Pessoa, Mata do Buraquinho (MVZ

185596); Pernambuco, Municı´pio de Tamandare´,

Coqueiral just outside S border Ibama headquar-

ters at Tamandare´ (MVZ 185598); Rio de Janeiro,

Municı´pio Duas Barras, Town Duas Barras, Gar-

den at house of Mario & Tereza Habib (MVZ

185603); Sa˜o Paulo, Sa˜o Paulo, (AMNH 207091);

Sa˜o Paulo, Valparaı´so (FMNH 41645*); Para-

guay: Alto Paraguay, Fuerte Olimpo (FMNH

145258*); Alto Paraguay, W Bank Rı´o Paraguay,

Estancia Puerto Ramos, 5 km SSE Bahı´a Negra

(AMNH 265411–265413); Canendiyu, Igatimi,

(AMNH 234285); Central, Asuncio´n, District Ca-

tedral (AMNH 205183–205185); Central, Asun-

cio´n, Recoleta (AMNH 248311–248325, 248503;

UMMZ 124321–124332, 125422–125443,

125876–125877, 125879–125901, 133734); Guai-

ra´, villarrica (AMNH 148662–148666); Itapu´a,

Trinidad (AMNH 36523); La Cordillera, 1.6 km

by road S Tobatı´ (UMMZ 125878); Paraguarı´,

Parque Nacional Ybycuı´ (UMMZ 133732); Para-

guarı´, Saltos de Pirareta (UMMZ 133733); Para-

guarı´, Sapucai (AMNH 23770–23773; FMNH

48791*).

Platyrrhinus masu:Bolivia: Cochabamba, 50

km NW Villa Tunari (UMMZ 126759–126760);

La Paz, Serranı´a Bellavista (AMNH 246610–

246614); La Paz, Serranı´a Bellavista, 47 km by

road N Caranavi (UMMZ 158068); Peru: Cuzco,

3 km E Amaybamba (MUSM 989–992); Cuzco,

La Convencio´n, Kimbiri, Camp. Llactahuaman

(MUSM 14559–14560, 14562*, 14565, 14566*);

Cuzco, La Convencio´n, Kimbiri, Camp. Wayra-

pata (MUSM 14568, 14570, 14572); Cuzco, La

Convencio´n, Rı´o Urubamba, Kiteni (MVZ

166595*); Cuzco, Paucartambo, 72 km by road

NE Paucartambo (UMMZ 160627, 160633); Cuz-

co, Paucartambo, Challabamba, P. V. Acjanaco

(FMNH 170112*; MUSM 8851); Cuzco, Paucar-

tambo, Consuelo, km 165, 17 km by road W Pil-

copata (FMNH 123917*; MUSM 9854); Cuzco,

Paucartambo, La Esperanza (FMNH 174759*,

174760); Cuzco, Paucartambo, Pillahuata

(MUSM 11793, 11794, 11795*); Cuzco, Paucar-

tambo, Quitacalzon, Carretera Paucartambo-Pil-

copata km 163 (MUSM 8852); Cuzco, Paucar-

tambo, San Pedro (FMNH 172100* ; MUSM

8853–8854, 11792); Cuzco, Paucartambo, Suecia,

km 138.5 road to Shintuya (FMNH 170113*);

Cuzco, Quispicanchi, Camante (FMNH 68455*);

Cuzco, Quispicanchi, Collpa de San Lorenzo

(FMNH 93594*); Cuzco, Quispicanchi, Hacienda

Cadena (FMNH 93588*); Madre de Dios, Manu,

Cerro de Pantiacolla (FMNH 122136*); Madre de

Dios, Manu, Cerro de Pantiacolla, upstream Rı´o

Palotoa (FMNH 139597–139599*, 139601–

129603*, 139606–139607*; MUSM 9849–9853,

9868, 9966); Madre de Dios, Manu, Rı´o Alto Ma-

dre de Dios, Hacienda Amazonia (FMNH

139590–139591*, 139593*; MUSM 9848);

Hua´nuco, Hua´nuco, Chinchao, Cordillera de Car-

pish (MUSM 18265*, 18266*); Pasco, Oxapam-

pa, San Alberto (MUSM 10272–10273); Pasco,

Pasco, Paucartambo, Auquimarca, Anexo Santa

Isabel (MUSM 15881–15884).

Platyrrhinus matapalensis:Ecuador: Bolı´var,

Barraganete, 3 km SW Echeandı´a (EPN 80.4.1,

80458); Esmeraldas, Borbo´n, Rı´o Cayapas, Rı´o

Chimbocal, Comuna Corriente Grande (QCAZ

2155); Esmeraldas, Borbo´n, Rı´o Cayapas, Com-

munity of Zapote (QCAZ 2066); Esmeraldas, Co-

munidad Valle del Sade, 8 km E mouth of the Rı´o

Sade in the Rı´o Esmeraldas (EPN 85904); Es-

meraldas, Eloy Alfaro, Borbo´n, Rı´o Santiago, Es-

tero Marı´a, Comuna Selva Alegre (QCAZ 1921);

Esmeraldas, Eloy Alfaro, San Miguel, Rı´o Caya-

pas, R. E. Cotacachi-Cayapas (QCAZ 508); Guay-

as, Naranjal, La Unio´n (QCAZ 2124); Guayas, El

Triunfo, El Piedrero (QCAZ 2624); Los Rı´os,

Quevedo, Rı´o Palenque (QCAZ 509); Pichincha,

Nanegal, Chacapata, Recinto Playa Rica (QCAZ

1900); Peru: Tumbes, Zarumilla, Z.R. Tumbes,

Quebrada Los Naranjos, Campo Verde (MUSM

10725–10726*); Tumbes, Zarumilla, Matapalo

(FMNH 81079–81081*).

Platyrrhinus nigellus:Bolivia: La Paz, 20 km

50 FIELDIANA: ZOOLOGY

NNE Caranavi (UMMZ 127174); La Paz, Serran-

ı´a Bellavista, 35 km N Canavari (AMNH 246616–

246620); Santa Cruz, 4.5 km N and 1.5 km E

Cerro Amboro, Rı´o Pitasama (AMNH 261665);

Colombia: Cesar, San Sebastia´n (FMNH

69484*); Narin˜o, El Carmen (FMNH 113891–

113892, 113894–113897); Ecuador: Azuay, Val-

le Yunguilla (FMNH 53504*); Loja, Loja, Mas-

anamaca, 12 km S Vilcabamba (QCAZ 1210);

Morona Santiago, P. N. Sangay Sardinayacu (EPN

964718); Napo, Loreto, P. N. Napo-Galeras, W

side, line 28 (QCAZ 1525); Napo, Loreto, P. N.

Napo-Galeras, W side, line 30 (QCAZ 1388–

1389); Napo, San Rafael Cascada (FMNH

124987*); Napo, Tena, Atacapi, Cordillera de Los

Guacamayos (QCAZ 1264); Pichincha, Nanegal,

Gavila´n de Orongo (QCAZ 1974); Pichincha, Es-

tacio´n Forestal La Favorita, close to Chiriboga

(EPN 799–10); Pichincha, Reserva Rı´o Guajalito

(QCAZ 3159); Peru: Amazonas, Cordillera del

Co´ndor (MUSM 10628); Ayacucho, Huanhu-

achayo (AMNH 233644–233684); Cajamarca,

San Ignacio (MUSM 12890–12892); Cuzco, La

Convencio´n, Kimbiri, Camp. Wayrapata (MUSM

14579*); Cuzco, La Convencio´n, Kimbiri, Camp.

Llactahuama´n (MUSM 14561, 14575*, 14576–

14578); Cuzco, Paucartambo, Bosque de las Nu-

bes, km 150 by road Paucartambo-Pilcopata

(MUSM 8857); Cuzco, Paucartambo, Consuelo,

km 165 by road Paucartambo-Shintuya (FMNH

123947*, 174777*, 174779*, 174781*, 174783*;

MUSM 9970–9971, 9975); Cuzco, Paucartambo,

Quitacalzo´n, road Paucartambo-Pilcopata km 163

(MUSM 8858, 8860); Cuzco, Paucartambo, San

Pedro (MUSM 11796); Cuzco, Quispicanchi,

Collpa de San Lorenzo (FMNH 93589–93593*,

93595–93597*, 93607*); Cuzco, Quispicanchi,

Hacienda Cadena (FMNH 93598–93606*); Madre

de Dios, Manu, Cerro de Pantiacolla, headwaters

of Rı´o Palotoa (FMNH 139577–139578*,

139580*; MUSM 9969); Madre de Dios, Manu,

Hacienda Amazonia (FMNH 126031–126032*;

MUSM 9955); Hua´nuco, Hua´nuco, Chinchao,

Cordillera de Carpish (MUSM 18263*, 18264*,

18273*); Pasco, Oxapampa, Pozuzo, Palmira

(MUSM 10943–10944, 10971); San Martı´n,

Huallaga, La Morada (MUSM 16170–16172,

16176); San Martı´n, Las Palmas, 32 km NE Pataz

(MUSM 7295–7296).

Platyrrhinus reciﬁnus:Brazil: Sa˜o Paulo

(USNM 545002); Sa˜o Paulo, Guaratuba (USNM

542612); Sa˜o Paulo, Iguape˜ (USNM 542613); Sa˜o

Paulo, Municı˜pio Ilhabela, Ilha de Sa˜o Sebastia˜o,

Parque Estadual Ilhabela (MVZ 185604*,

185605, 185606*, 185607–185608, 185609–

185611*); Sa˜o Paulo, Municı´pio Saleso´polis, Mu-

seu de Zoologia USP, Estac¸a˜o Biolo´gica Borace´ia

(MVZ 185901*).

Platyrrhinus vittatus:Colombia: Magdalena,

Valparaı´so (AMNH 15100–15101*); Narin˜o, Ri-

caurte (AMNH 34232*); Costa Rica: Puntarenas,

Monteverde (UMMZ 116681–116682*); Pana-

ma: Darie´n, ca. 6 km NW Cana, E slope Cerro

Pirre (LSUMZ 25464–25466*); Darie´n, Cerro

Mali (AMNH 238203–238209); Darien, Cerro Ta-

carcuna, Rı´o Pucuro (AMNH 238210–238218);

Venezuela: Carabobo, Puerto Cabello (ZMB

568).

Carollia subrufa:Mexico: Chiapas, 7.5 mi SW

by road Ixtapa (AMNH 249087); Oaxaca, Juchi-

ta´n, San Jero´nimo (AMNH 36037–36038); Oa-

xaca, Juchita´n, 10 mi S Tapanatepec (AMNH

177625–177626); Oaxaca, Tehuantepec, Limo´n

(AMNH 208238–208240, 167028, 171642–

171644, 171648); Oaxaca, Tehuantepec, Cerro

San Pedro (AMNH 146150); Oaxaca, 60 mi Te-

huantepec (AMNH 189729).

Sturnira erythromos:Peru: Amazonas, ca. 20

km by road W Leymebamba (MUSM 4908–

4910); Ayacucho, Yuraccyacu (MUSM 5269);

Cuzco, Paucartambo, Morro Leguia, km 135 road

Paucartambo-Shintuya (MUSM 8866–8868);

Cuzco, Paucartambo, Challabamba, P. V. Acjana-

co (MUSM 8876–8877); Cuzco, Paucartambo,

Pillahuata (MUSM 9980); Cuzco, Paucartambo,

San Pedro (MUSM 11799); Hua´nuco, Hua´nuco,

Cordillera Carpish (MUSM 172); Hua´nuco, Un-

chog pass between Chunchubamba and Hacienda

Paty, NNW Acomayo (MUSM 5260); La Liber-

tad, Mashua, E of Tayabamba, road to Ongo´n

(MUSM 5285); La Libertad, Utcubamba, road to

Ongo´n (MUSM 5310).

Uroderma magnirostrum:Peru: Cuzco, La

Convencio´n, Camisea, Armihuari (MUSM

14019); Cuzco, La Convencio´n, Camisea, Kon-

kariari (MUSM 14841–14842); Cuzco, La Con-

vencio´n, Camisea, Pagoreni (MUSM 14026);

Cuzco, La Convencio´n, Camisea, San Martı´n

(14027–14037); Cuzco, La Convencio´n, Camisea,

Segakiato (14843–14844); Ucayali, Padre Abad,

Bosque Nacional Alexander von Humboldt

(MUSM 8630–8631).

Vampyrodes caraccioli:Costa Rica: Puntar-

enas, 2 km SW Rincon de Osa (MSB 26965–

26969); Ecuador: Pichincha, Estacio´n Forestal

La Favorita, close to Chiriboga (EPN 799–12,

799–15); Sucumbı´os, Shushuﬁndi, Limoncocha

(QCAZ 514); Peru: Cuzco, La Convencio´n, Cam-

51VELAZCO: PLATYRRHINUS PHYLOGENY

isea, Armihuari (MUSM 14070–14072); Cuzco,

La Convencio´n, Camisea, Pagoreni (MUSM

14073); Cuzco, La Convencio´n, Camisea, San

Martı´n (MUSM 14075–14078); Loreto, Rı´o Sa-

miria, Tacshacocha (MUSM 1017); Madre de

Dios, Manu, Cerro de Pantiacolla (MUSM

10102); Madre de Dios, Manu, Pakitza (MUSM

739, 12625–12624); Madre de Dios, Manu, Ha-

cienda Amazonia (MUSM 10100–10101, 10106);

Pasco, Oxapampa, Pozuzo (MUSM 10997); Pas-

co, Oxapampa, Pozuzo, Rı´o Negro (MUSM

10996); Ucayali, Atalaya, Sepahua (MUSM

12809).

Appendix 2: Data Matrix

This data matrix includes all the taxa examined

(ingroup and outgroup) and all the characters used

in the phylogenetic analysis. Polymorphisms are

shown between brackets.

Carollia subrufa

0–02 10202 01120 10021 10000 00000 00000 0–

110 00000 10200 01001 000{01}0

Sturnira erythromos

0–10 02000 00232 12001 0000{01} 00000 00000

0–010 00000 00110 00000 00020

Uroderma magnirostrum

10012 03022 21110 {01}0022 00011 11110

00101 11010 00101 11110 01102 10201

Vampyrodes caraccioli

10013 03121 31021 01{01}11 10100 11110

{01}0121 0–111 10010 11011 01102 11121

Platyrrhinus aurarius

10103 03122 20221 12112 12100 11111 00021

20111 10101 01010 01212 11221

Platyrrhinus brachycephalus

10012 14121 {23}112{01} 11{01}00

101{01}{01} 11111 {01}1{01}11 {12}0{01}01

10101 01022 11212 112{01}1

Platyrrhinus chocoensis

10102 0{23}1{12}2 2{12}111 11{01}0{01}

1{01}100 11111 00021 {12}0{01}21 1{01}102

01010 01212 102{12}1

Platyrrhinus dorsalis

10103 03121 20121 {01}1101 12100 11111

{01}0021 {12}0121 {01}{01}112 01020 01212

112{12}1

Platyrrhinus dorsalis ‘‘Norte’’—Platyrrhinus is-

maeli sp. nov.

10103 03121 20122 121{01}1 1110{01} 11111

{01}0{01}21 21121 {01}11{01}1 01020 01211

11221

Platyrrhinus dorsalis ‘‘Centro-Sur’’—Platyrrhin-

us masu sp. nov.

10103 011{12}2 31122 12{01}01 1{01}101

11111 {01}0021 20111 {01}11{01}1 01020

01212 112{12}1

Platyrrhinus helleri ‘‘Eastern’’—Platyrrhinus hel-

leri

1001{12} {01}312{12} {23}1111 {01}2{01}11

101{01}0 11111 01{01}11 10{01}21 10101

01021 11212 {01}1201

Platyrrhinus helleri ‘‘Western’’—Platyrrhinus

matapalensis sp. nov.

10012 03221 211{01}0 {01}2{01}00

101{01}{01} 11111 {01}1{01}11 10{01}11

1010{12} 01020 11212 11{12}01

Platyrrhinus infuscus

10103 {01}32{12}{12} 12120 {01}1{01}12

111{01}{01} 11111 11{01}21 20111 1111{12}

01020 01210 102{12}1

Platyrrhinus lineatus

10012 03121 31122 12022 1010{01} 11111

0{01}0{12}1 20111 101{01}{12} 01020 01210

11221

Platyrrhinus nigellus

10103 {01}3122 {23}0122 {01}2{01}01

1{12}100 11111 {01}0{01}{12}1 {12}0111

10{01}0{12} 01020 0{01}21{12} 11221

Platyrrhinus reciﬁnus

10012 03021 31222 12011 111{01}0 11111

11011 10111 10102 01020 01212 10201

Platyrrhinus vittatus ‘‘Northern’’—Platyrrhinus

vittatus

11103 03222 30112 {01}1{01}11 10100 11111

10021 201{01}1 11111 01021 01211 11221

Platyrrhinus vittatus ‘‘Southern’’—Platyrrhinus

albericoi sp. nov.

11003 03{12}21 20112 12{01}12 1010{01}

11111 10121 20111 11011 01020 01211 11211

Appendix 3: Optimization of

Characters

The following apomorphies support the mono-

phyletic clades shown in Figure 29. Both ACCT-

RAN and DELTRAN optimizations are given for

each clade. Diagnosis for each character is given

as follow: (character number; consistency index)

character state description, state number

→

state

number. Unambiguous transformations are indi-

cated by a double arrow ‘‘

⇒

,’’ transformations

52 FIELDIANA: ZOOLOGY

. 29. Strict consensus tree with nodes numbered as a reference for Appendix 3, which presents apomorphies

of the clades.

that occur in only one optimization are indicated

by a single arrow ‘‘

→

.’’

Platyrrhinus (Node 1)—ACCTRAN: (30;

1.000) Sulcus present on the anterior face of P4,

⇒

1; (33; 0.333) M1 mesostyle absent, 1

→

(36; 0.667) Stylar cusp present on the lingual

cingulum of the M1 metacone, 0

⇒

2; (45;

0.500) lingual cingulum of the M2 metacone

continuous to the paracone, 0

⇒

2; (46; 0.500)

M2 hypoconal basin developed, 1

⇒

0; (49;

0.500) both labial and lingual cingula present on

p4, 1

⇒

2; (53; 1.000) both labial and lingual

cingula present on m1, 1

⇒

2; (54; 1.000) stylid

cusps present on the anterior face of the m1 pro-

toconid, 0

⇒

1. DELTRAN: (30; 1.000) Sulcus

on the anterior face of P4, 0

⇒

1; (36; 0.667)

Stylar cusp present on the lingual cingulum of

the M1 metacone, 0

⇒

2; (43: 0.333) M2 metas-

tyle present, 0

→

1; (45; 0.500) lingual cingulum

of the M2 metacone continuous to the paracone,

⇒

2; (46; 0.500) M2 hypoconal basin devel-

oped, 1

⇒

0; (53; 1.000) both labial and lingual

cingula present on m1, 1

⇒

2; (54; 1.000) stylid

cusps on the anterior face of the m1 protoconid,

⇒

Node 2—ACCTRAN: (3; 0.500) dark facial

stripes, 0

⇒

1; (4; 0.500) Absence of a basal pro-

tuberance, where the genal vibrissae are implant-

ed, 1

⇒

0; (10; 0.333) folds in the pinna well

marked, 1

→

2; (11; 0.429) dorsal stripe deﬁnite,

but narrow, 3

→

2; (18; 1.000) posterior border

of the hard palate ‘‘V’’-shaped, 0

→

1; (19; 0.286)

postorbital process absent or poorly developed, 1

⇒

0; (39; 0.400) M1 protocone small and blunt,

→

2. DELTRAN: (3; 0.500) dark facial stripes,

⇒

1; (4; 0.500) Absence of a basal protuber-

ance, where the genal vibrissae are implanted, 1

⇒

0; (11; 0.429) dorsal stripe deﬁnite, but narrow,

53VELAZCO: PLATYRRHINUS PHYLOGENY

→

2; (19; 0.286) postorbital process absent or

poorly developed, 2 0.

Node 3—ACCTRAN: (5; 0.667) six vibrissae

surrounding the margins of the noseleaf in a sin-

gle array, 2

⇒

3; (12; 0.500) dorsal fur

8 mm,

⇒

0; (22; 0.500) A deep fossa on the squamosal

end of the zygomatic arch, lateral to the glenoid

fossa, 0

⇒

2. DELTRAN: (5; 0.667) six vibrissae

surrounding the margins of the noseleaf in a sin-

gle array, 2

⇒

3; (12; 0.500) dorsal fur

8 mm,

⇒

0; (18; 1.000) posterior border of the hard

palate ‘‘V’’-shaped, 0

→

1; (22; 0.500) A deep

fossa on the squamosal end of the zygomatic arch,

lateral to the glenoid fossa, 0

⇒

Node 4—ACCTRAN: (17; 0.286) A densely

haired uropatagium fringe, 1

⇒

2; (39; 0.400) M1

protocone moderately developed, 2

→

1; (45;

0.500) lingual cingulum of the M2 metacone re-

stricted to the metacone, 2

⇒

1. DELTRAN: (17;

0.286) A densely haired uropatagium fringe, 1

⇒

2; (45; 0.500) lingual cingulum of the M2 meta-

cone restricted to the metacone, 2

⇒

Node 5—ACCTRAN: (15; 0.286) dense and

long hair on the dorsum of the feet, 1

⇒

2; (22;

0.500) A shallow fossa on the squamosal end of

the zygomatic arch, lateral to the glenoid fossa, 2

→

1; (31; 0.500) A deep fossa on the hypoconal

basin of P4, 0

→

1. DELTRAN: (15; 0.286) dense

and long hair on the dorsum of the feet, 1

⇒

(49; 0.500) both labial and lingual cingula of p4

present, 1

→

Node 6—ACCTRAN: (42; 1.000) stylar cusp

present on the lingual face of the M2 paracone, 0

⇒

1; (44; 0.333) stylar cusp present on the lingual

face of the M2 metacone, 0

→

1. DELTRAN: (22;

0.500) A shallow fossa on the squamosal end of

the zygomatic arch, lateral to the glenoid fossa, 2

→

1; (42; 1.000) stylar cusp present on the lingual

face of the M2 paracone, 0

⇒

Node 7—ACCTRAN: (10; 0.333) folds in the

pinna poorly marked but distinguishable, 2

→

(19; 0.286) postorbital process moderately devel-

oped, 0

→

1; (55; 0.400) poorly developed m1

metaconid, 2

⇒

1. DELTRAN: (55; 0.400) poorly

developed m1 metaconid, 2

⇒

Node 8—ACCTRAN: (8; 0.400) two interra-

mal vibrissae present, 1

⇒

2; (17; 0.286) edge of

the uropatagium, usually hairy, 2

→

1; (20; 0.333)

A well-developed paraoccipital process, 1

→

DELTRAN: (8; 0.400) two interramal vibrissae

present, 1

⇒

2; (19; 0.286) postorbital process

moderately developed, 0

→

1; (31; 0.500) A deep

fossa on the hypoconal basin of P4, 0

→

1; (44;

0.333) stylar cusp present on the lingual face of

the M2 metacone, 0

→

Node 9—ACCTRAN: (2; 1.000) dorsomedial

facial stripe more marked than the ventrolateral

stripe, 0

⇒

1; (14; 0.333) ventral fur bicolored, 2

⇒

1; (22; 0.500) An almost imperceptible fossa

on the squamosal end of the zygomatic arch, lat-

eral to the glenoid fossa, 1

⇒

0. DELTRAN: (2;

1.000) dorsomedial facial stripe more marked

than the ventrolateral stripe, 0

⇒

1; (14; 0.333)

ventral fur bicolored, 2

⇒

1; (22; 0.500) An al-

most imperceptible fossa on the squamosal end of

the zygomatic arch, lateral to the glenoid fossa, 1

⇒

Node 10—ACCTRAN: (15; 0.286) dense and

long hair on the dorsum of the feet, 1

→

2; (17;

0.286) A densely haired uropatagium edge, 1

⇒

2; (32; 0.500) M1 parastyle present, 0

→

1; (34;

1.000) presence of a labial cingulum at the base

of the M1 metacone, 2

→

1. DELTRAN: (10;

0.333) folds in the pinna poorly marked but dis-

tinguishable, 2

→

1; (17; 0.286) A densely haired

uropatagium edge, 1

⇒

2; (49; 0.500) both labial

and lingual cingula on p4 present, 1

→

Node 11—ACCTRAN: (45; 0.500) lingual cin-

gulum of the M2 metacone restricted to the meta-

cone, 2

⇒

1; (51; 1.000) stylid cusp present on

the posterior face of the main cone of p4, 0

⇒

DELTRAN: (45; 0.500) lingual cingulum of the

M2 metacone restricted to the metacone, 2

⇒

(51; 1.000) stylid cusp present on the posterior

face of the main cone of p4, 0

⇒

Node 12—ACCTRAN: (19; 0.286) postorbital

process, absent or poorly developed, 1

⇒

0; (20;

0.333) paraoccipital process, poorly developed, 1

⇒

0. DELTRAN: (19; 0.286) postorbital process,

absent or poorly developed, 1

⇒

0; (20; 0.333)

paraoccipital process, poorly developed, 1

⇒

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Phylogeographic accounts of mammals across fragmented landscapes show high levels of genetic, morphological and ecological variation. The big brown bat (Eptesicus fuscus) widely spans mainland landmasses from Canada to Ecuador and Colombia, and the insular Caribbean through The Bahamas and Greater and Lesser Antilles. Given the distribution of E. fuscus, we hypothesized that insular lineages could represent a different species aided by isolation. We assessed species limits by capitalizing on available mitochondrial and genomic data. Novel morphological and spatial datasets were produced to examine limits phenotypically and whether ecological niches could be associated with differences between groups. Phylogenetics strongly supported the Caribbean as unique compared to the mainland. Genomic data indicated high levels of genetic structure within the Caribbean and no detectable admixture of the Caribbean with continental lineages. Similarly, the Caribbean group shows high phenotypic disparity, and niche models revealed differences in habitat suitability between groups, concordant with the phylogenetic results. This study uncovered signals of divergence supporting the Caribbean clade of E. fuscus as unique through an integrative framework. We endorse re-evaluating the taxonomic status of Caribbean big brown bats as Eptesicus dutertreus. This recognition can help promote local conservation plans for insular lineages of big brown bats.

Catálogo ilustrado de flora y fauna en el municipio de Guarne

Book

Dec 2018

El centro de estudios territoriales- CET; de la Universidad Católica de Oriente, desde su línea de investigación en estudios territoriales, tiene dentro de sus propósitos brindar aportes al desarrollo territorial y promover la generación del nuevo conocimiento. Con su clara misión de acompañar los procesos del territorio, el CET, abre un espacio para realizar el Diagnóstico para el municipio de Guarne, el cual se constituye un aporte significativo desde el campo de la planificación que conduce a fortalecer los procesos de desarrollo en la región. En esta publicación, se logra catalogar las distintas especies de flora, mamíferos, peces y aves que hacen parte del territorio guarneño que, al ser valoradas y preservadas en su habitat natural, conversan la cadena biológica que protege los recursos naturales y proyectan el desarrollo.

List of bats register at high altitudes in Río Abiseo National Park

Article

Full-text available

Dec 2023

We present a list of bats collected above 2000 meters in Río Abiseo National Park. Sixteen bat species belonging to the Phyllostomidae and Vespertilionidae families were collected during four expeditions between 1987 and 1990. Twelve species were found within expected altitudinal ranges, but four exceeded their previously reported maximum altitudinal ranges. A comparison with other bat lists from other localities at similar altitudes is presented.

Taxonomic revision and additional comments of some bats (Mammalia, Chiroptera) reported from Bolivia, with an updated checklist based on voucher material with verified identities

Article

Full-text available

Jun 2023

We review the taxonomic identities of museum specimens of three bat species previously reported from Bolivia. We comment on some erroneously reported taxa, or taxa either not represented by voucher materials or based on insufficient data (including acoustic detections) to verify their taxonomic affinities. As result of this review, the list of bat species known to occur in Bolivia is updated to eight families and 133 species, unlike the nine families and 146 species of previous lists. Some recommendations for future research and a brief historical revision of bat inventories in the country are included.

Taxonomic revision and additional comments of some bats (Mammalia, Chiroptera) reported from Bolivia, with an updated checklist based on voucher material with verified identities

Article

Full-text available

Jun 2023

On the cusp of adaptive change: the hierarchical radiation of phyllostomid bats

Preprint

Full-text available

May 2023

Adaptive radiations are bursts in biodiversity that lead to the origin of new evolutionary lineages and phenotypes. However, adaptive radiations typically occur over millions of years and it is unclear how the macroevolutionary dynamics that underpin them vary through time and among groups of organisms. Phyllostomid bats radiated extensively for diverse diets –from insects to vertebrates, fruit, nectar, and blood– and we use their molars as a model system to examine the dynamics of adaptive radiations. Three-dimensional shape analyses of lower molars of Noctilionoidea (Phyllostomidae and close relatives) indicate that different diet groups exhibit distinct morphotypes. Comparative analyses further reveal that phyllostomids are a striking example of a hierarchical radiation; their initial, higher-level diversification involved an ‘early burst’ in molar morphological disparity as lineages invaded new diet-affiliated adaptive zones, followed by subsequent lower-level diversifications within adaptive zones involving less dramatic morphological changes. We posit that strong selective pressures related to initial shifts to derived diets may have freed molars from morpho-functional constraints associated with the ancestral molar morphotype. Then, lineages with derived diets (frugivores and nectarivores) diversified considerably within broad adaptive zones, likely reflecting finer-scale niche partitioning. The observed early burst pattern is only evident when examining molar traits that are strongly linked to diet, highlighting the importance of ecomorphological traits in comparative studies. Our results support the hypothesis that adaptive radiations are commonly hierarchical and involve different tempos and modes at different phylogenetic scales, with early bursts being more common at broader scales. SIGNIFICANCE STATEMENT Many groups of organisms are exceptionally diverse in their ecology, morphology, and number of species. But there is debate as to whether these groups commonly achieved this diversity through ‘bursts’ in diversification early in their history. Phyllostomid bats are one of the most ecologically diverse mammalian families and a classic example of an adaptive radiation. We use their molar shapes, which correlate with diet, as a model for examining macroevolutionary patterns during diversifications. We find that phyllostomids experienced a two-step process of diversification; the first step involved a rapid burst, whereas the second involved finer-scale changes as lineages filled ecological niches. We posit that this is a common, yet underappreciated, pattern during the early histories of many diverse clades.

A new species of Vampyrops (Chiroptera: Phyllostomatidae) from South American

Book

Jan 1972

A reappraisal of interfamilial relationships of bats;

Article

Jan 1998

N.B. Simmons

Distribution and zoogeography of Peruvian bats, with comments on natural history

Article

Oct 1970

M.D. Tuttle

Biogeography of the bats of South America

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Jan 1982

K.F. Koopman

Mammals of the Smithsonian Venezuelan Project

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Jan 1976

Charles O Handley

A new species of Micronycteris (Chiroptera: Phyllostomidae) from northeastern Brazil, with comments on phylogenetic relationships

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Jan 1996

N.B. Simmons

XLI.—Three small mammals from S. America

Article

Oldfield Thomas

Phylogenetic relationships of mormoopid bats (Chiroptera: Mormoopidae) based on morphological data

Article

Jan 2001

Mormoopidae is a small family of Neotropical microchiropteran bats that includes two genera (Mormoops and Pteronotus) and ten species, two of which are known only from fossils. Mormoopidae is typically classified as a member of Noctilionoidea, a group that minimally includes two other Neotropical families (Phyllostomidae and Noctilionidae) and may also include Mystacinidae, a taxon endemic to New Zealand and Australia. Phylogenetic relationships of extant mormoopid species and one extinct taxon, Pteronotus pristinus, were investigated in a series of parsimony analyses of 209 morphological characters including features of the skull, dentition, vomeronasal organ complex and brain, trachea and hyoid apparatus, tongue, face, ears, pelage, patagia, postcranial skeleton, postcranial myology, reproductive tract, and digestive tract. Three extant phyllostomid species, two noctilionids, two mystacinids, and one emballonurid species were included as outgroups to test monophyly of Mormoopidae and to provide a context for determining the sister group of the family. Results of parsimony analyses under a variety of different assumption sets indicate that Mormoopidae is monophyletic, and that Pteronotus and Mormoops are monophyletic sister taxa. Within the genus Pteronotus, several clades were repeatedly recovered: (1) P. davyi + P. gymnonotus (= subgenus Pteronotus); (2) P. macleayi + P. quadridens; (3) P. personatus + P. macleayi + P. quadridens (= subgenus Chilonycteris); (4) P. parnellii + P. pristinus + P. personatus + P. macleayi + P. quadridens; and (5) P. parnellii + P. pristinus (= subgenus Phyllodia). These results support monophyly of all subgenera of Pteronotus previously recognized, and additionally indicate that the subgenera Pteronotus and Chilonycteris are sister taxa. Comprehensive diagnoses for each species and clade of Mormoopidae are provided based on character optimizations and ancillary morphometric data from the literature. Results of our parsimony analyses also have implications for understanding higher-level phylogeny of noctilionoid bats. Monophyly of each of the traditionally recognized families (i.e., Noctilionidae, Mystacinidae, Mormoopidae, and Phyllostomidae) was strongly supported. In congruence with recent analyses of mitochondrial gene-sequence data and DNA hybridization experiments, we found strong support for inclusion of Mystacinidae in Noctilionoidea. Using an emballonurid species to root the tree, we found the following interfamilial relationships of noctilionoids: (Noctilionidae (Mystacinidae (Phyllostomidae, Mormoopidae))). Lists of morphological synapomorphies of each of these groups are provided based on character optimizations.

Description of New Species of Diatoms

Article

H.L. Smith

A systematic review of the large species of Artibeus Leach, 1821 (Mammalia: Chiroptera), with some phylogenetic inferences

Article

Jan 1994

S.A. Marques-Aguiar

Morphological Phylogeny of the Bat Genus Platyrrhinus Saussure, 1860 (Chiroptera: Phyllostomidae) with the Description of Four New Species

Abstract and Figures

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