new species of vampyressa

7/23/2019 New species of Vampyressa

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Copyright American Museum of Natural History 2014 ISSN 0003-0082

AMERICAN MUSEUM NOVITATES

Number 3813, 27 pp. October 2, 2014

Systematics o VampyressamelissaTomas, 1926(Chiroptera: Phyllostomidae), with descriptions o two

new species o Vampyressa

VALRIA DA C. AVARES,1ALFRED L. GARDNER,2 HCOR E. RAMREZ

CHAVES,3AND PAL M. VELAZCO4

ABSRAC

Vampyressa melissa is a poorly known phyllostomid bat listed as vulnerable by the Inter-national Union or Conservation o Nature (IUCN). Since its description in 1926, ewer than40 V. melissa have been reported in the literature, and less than hal o these may have beencorrectly identied. During revisionary studies o Vampyressa, we uncovered two previouslyunrecognized species related to V. melissa, all associated with higher elevation habitats (>1400m), one rom the Andes o Colombia (Vampyressa sinchi, new species) and the other romwestern Panama (Vampyressa elisabethae,new species) revealing that V. melissa, as traditionallydened, is a composite o at least three species. In this paper, we provide a restricted diagnosisor the genus Vampyressa, an emended diagnosis o V. melissa, and descriptions o the two new

species. Te separation o these rugivorous bats, previously identied as V. melissa, into threeisolated upper-elevation species, each having restricted distributions urther highlights theirragile conservation status.

1 Instituto Nacional de Pesquisas da Amaznia (INPA), Brazil; Programa de Ps Graduao em Gentica,Conservao e Biologia Evolutiva, Brazil; Programa de Colees e Acervos Cientcos, PCAC, Brazil; Divi-sion o Vertebrate Zoology (Mammalogy), American Museum o Natural History; the Graduate Center, theCity University o New York, New York.

2USGS Patuxent Wildlie Research Center, National Museum o Natural History MRC111, SmithsonianInstitution, Washington, DC.

3Weisbecker Lab, School o Biological Sciences, University o Queensland, Brisbane, Australia.

4Department o Vertebrate Paleontology, American Museum o Natural History, New York.


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2 AMERICAN MUSEUM NOVIAES NO. 3813

INRODUCION

Commonly known as yellow-eared bats, species o Vampyressaand their allies have been

regarded as primarily or exclusively rugivorous (Gardner, 1977a); nevertheless, little is knownabout their eeding habits and ecology. Te genus Vampyressa(sensu lato Simmons, 2005) hasreceived much attention beginning with the recognition o its paraphyly as a result o the com-prehensive morphological phylogeny developed by Wetterer et al. (2000). Gardner (1977b)illustrated the major chromosomal differences in Vampyressa(sensu lato) and Peterson (1968)recognized three distinct evolutionary lineages or Vampyressa(s.l.) as the subgenera Vampy-ressa, Metavampyressa, and Vampyriscus, a revision acknowledged in Koopmans (1994) clas-sication. Davis (1975), however, recognized two subgenera: Vampyriscus (withMetavampyressaa synonym) to include V. bidens, V. brocki, and V. nymphaea; and Vampyressa to include V.

pusillaand V. melissa. More recent work has provided molecular support or treating Vampy-ressaand Vampyriscusas separate genera (Baker et al., 2003; Hooer and Baker, 2006; Hooeret al., 2008). Tis arrangement was recognized by Arroyo-Cabrales (2008a, 2008b) and is ol-lowed here. A summary o historical taxonomy and contents o Vampyressaand Vampyriscussince the three subgenera proposal o Peterson (1968) is provided in the table 1.

Revisionary work at the species level is scarce; a recent exception is that o Lim et al. (2003)who recognized V. pusillaand V. thyoneas separate species.Vampyressa melissa, as restrictedhere, is among the rarer Neotropical stenodermatines. Te species is restricted to Andeanslopes (between 1180 and 2600 m) with ewer than 40 literature records attributed to this taxon,and even ewer specimens (approximately 20) actually deposited in museum collections. Old-eld Tomas (1926) described the species based on material collected in the northern regiono Chachapoyas, Peru, by R.W. Hendee during the Godman-Tomas Expedition. Tomas(1926: 157) remarked that V. melissa, among the ve species o Vampyressarecognized at thattime, was a very strongly marked species. Goodwin (1963) noted the larger size o V. melissaas compared to other Vampyressa, along with other distinctive characters such as the hairy andringed uropatagium, and the peculiar shape and relative size o its lower molars. Although V.melissa is indeed a distinctive species, easily recognizable upon close examination, variousinterpretations o the characters mentioned in the ew published reports (e.g., Peterson, 1968),together with the lack o taxonomic keys and comparisons with closely related taxa, have con-

tributed to many misidentications.Ongoing morphological and quantitative studies o Vampyressa have revealed that specimens

previously identied as V. melissaactually represent three species, one rom Central America andtwo (including V. melissa) rom South America. Te Central American orm is represented by asmall series o our specimens in the National Museum o Natural History, Washington, DC,collected in the early 1960s during a countrywide survey o the ectoparasites o Panamanianmammals. Another specimen rom Panama was acquired by the National Museum in 1976.

A single specimen collected in the eastern Cordillera o Colombia in the 1970s by Kjell vonSneidern and labeled as Vampyressa melissa (FMNH 114028) also differs rom V. melissa

(BMNH 26.5.3.4) and represents an undescribed South American orm. We later discoveredtwo additional specimens matching the characteristics o FMNH 114028.


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2014 AVARES E AL.: SYSEMAICS OF VAMPYRESSA 3

Herein, we report on our systematic study o these specimens previously identied as V. melissa.In addition, we provide a restricted diagnosis o the genus Vampyressa, a diagnosis and reviseddescription o V. melissa, and descriptions o two previously undescribed species o Vampyressa.

MAERIAL AND MEHODS

External and craniodental characters are based mainly on, but not restricted to, thosedened by avares (2008) and Wetterer et al. (2000). Other anatomical eatures are derivedrom Giannini et al. (2006) and Giannini and Simmons (2007), or are described as ollows:

Accessory cusp near apex o upper canine. A small cusp at the apex o the mediolateralridge orming the lingual border o the longitudinal sulcus on posterior crown o upper canines.In most stenodermatines, the mediolateral ridge is complete to the tip o the canine. In Vampy-ressa(andMesophylla) the mediolateral ridge is incomplete and terminates as a small subapicalstep on the posterointernal crown o the canine. Ectophyllahas a similar accessory subapicalcusp, which differs in that it is thicker and is at the apex o both the mediolateral and a second-ary ridge along the lingual surace o the upper canine (avares, 2008: chars. 191 and 192).

Accessory medial oramen. A small oramen located either in the septum (medial ange othe palatine process o the premaxilla) separating incisive oramina, or located in the midline

between premaxillae anterior to the incisive oramina. In all Vampyressaexamined, this ora-men is in the septum (illustrated herein).

ABLE 1. Historical taxonomy and contents o Vampyressa and Vampyriscusdeparting rom the split intothree subgenera proposed by Peterson (1968).

Peterson (1968)

a

Davis (1975)

a

Lim et al. (2003)

e

Arroyo-Cabrales (2008a, b)

Vampyressabmelissa Vampyressabmelissa Vampyressabmelissa Vampyressa melissa

Vampyressabpusilla Vampyressabpusilla Vampyressabpusilla Vampyressa pusilla

pusillac

Synonym: Synonym: Synonym:

natterid nattereri nattereri

thyonec Vampyressabthyone Vampyressa thyone

Synonyms: Synonyms: Synonyms:

minuta minuta minuta

venilla venilla venilla

Metavampyressabbrocki Vampyriscusbbrocki Vampyriscusbbrocki Vampyriscus brocki

Metavampyressabnymphaea Vampyriscusbnymphaea Vampyriscusbnymphaea Vampyriscus nymphaea

Vampyriscusbbidens Vampyriscusbbidens Vampyriscusbbidens Vampyriscus bidens

aClassication based on morphology.bSubgenus o Vampyressa.cSubspecies o V. pusilla.dIncorrect spelling o Vampyressa nattereriGoodwin (1963: 16).eClassication based on morphology and mtDNA data.Te classication ollowed by this paper; supported by the molecular studies o Baker et al. (2003), Hooer and Baker(2006), and Hooer et al. (2008).


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Extension o caudal border o pterygoid.Te caudal border o the pterygoid process extendsposterolaterally across the basisphenoid toward the auditory bullae as a low ridge. Tis ridge

passes anterior to the boney excrescences and orms the anterior margin o the eustachialgroove. In Vampyressathis ridge is low and well separated rom the oramen ovale; in somestenodermatines (e.g., Vampyriscus), the ridge may be more highly developed and betterdescribed as a ange; in others, this ridge may lie along the posterior border o the oramenovale (e.g., Platyrrhinus) or on a bridge separating anterior and posterior segments o a largeoramen ovale (e.g., Chiroderma).

Mesethmoid plate. Tere are differences among species o Vampyressain the morphology othe mesethmoid plate visible in rontal view within the external nares (illustrated herein: g. 5).

Posttympanic process. Te posttympanic process is comprised o a ventral extension o thesquamosal and lateral process o the exoccipital that covers the posterolateral rim o the otic cap-sule. In Vampyressa(andMesophylla) these two bones are unused, whereas in other vampyressinestenodermatids these bones are used at least at the tips. Te unused condition as well as thesubapical cusp on the canine is illustrated in gure 26A (Mesophylla) in Wetterer et al. (2000).

Sphenorbital oramen or ssure. In Vampyressa, this opening, posterior to the optic ora-men, is either a discrete oramen or an elongate ssure (e.g., Giannini et al., 2006: g. 11).

Specimens examined or this study are housed in the ollowing collections:

AMNH American Museum o Natural History, New York

BMNH British Museum (Natural History) London, UKCMUFLA Coleo de Mameros da Universidade de Lavras, Lavras, Minas Gerais,

BrazilDZSJRP Coleo de Chiroptera do Departamento de Zoologia de So Jos do Rio

Preto, So Paulo, BrazilFMNH Field Museum o Natural History, Chicago, IllinoisIAvH Instituto de Investigacin de Recurson Biolgicos Alexander von Humboldt,

Villa de Leyva, Boyac, ColombiaMEPN Museo Escuela Politcnica Nacional, Quito, Ecuador

MHNUC Museo de Historia Natural, Universidad del Cauca, Popayn, ColombiaMUSM Museo de Historia Natural de la Universidad Nacional Mayor de San Marcos,Lima, Peru

MZUSP Museu de Zoologia da Universidade de So Paulo, So Paulo, BrazilRMNH National Museum o Natural History, Leiden, NetherlandsUMMZ Museum o Zoology, University o Michigan, Ann Arbor, MichiganUSNM National Museum o Natural History, Smithsonian Institution, Washington,

D.C.

For the morphometric analyses we measured 56 specimens o the ve species o Vampy-ressa we recognize here(appendix). Our observations are based on adults, with the exceptiono a single individual (MUSM 8915) demonstrated as young by the retention o an upper lef


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deciduous premolar. Te dental homology or premolars ollows Miller (1907): rst upperpremolar (P3), second upper premolar (P4), rst lower premolar (p2), and second lower pre-

molar (p4). We recorded external and craniodental measurements in millimeters (mm) usingdigital calipers accurate to 0.01 mm; body mass is in grams (g). Standard external measure-ments (L, total length; HF, hind oot length; E, ear length; and body mass) were taken romskin labels or database records. Te abbreviation s.u. ollowing the measurement o the oot othe holotype o V. melissameans sine unguis(without claw) and was the standard practice inEurope or recording oot measurements. Measurements are dened as ollows:

otal length (L): Distance rom the tip o the snout to end o the body.Hind oot length(HF): Distance rom the base o calcar to tip o the claw o longest toe.Ear length (E): Perpendicular height rom the ear notch to tip o the pinna without

stretching.ragus length(rL): Perpendicular height o tragus, measured rom the notch to tip.Forearm length(FA): Distance rom the end o olecranon process to wrist (including carpals)

with the wing olded.Tird metacarpal length(MEIII): Distance rom the wrist (including carpals) o the olded

wing to distal end o third metacarpal.Fourth metacarpal length (MEIV): Distance rom the wrist (including carpals) o the olded

wing to distal end o ourth metacarpal.Fifh metacarpal length (MEV): Distance rom the wrist (including carpals) o the olded

wing to distal end o fh metacarpal.ibia length (iL): Distance rom articulation with emur to articulation with oot (including

tarsals).Greatest length o the skull(GLS): Distance rom the posteriormost point o occipital to the

anteriormost surace o premaxillae (not including incisors).Condyloincisive length(CIL): Distance rom a line connecting the posteriormost margins o

occipital condyles to the anterior ace o upper incisor(s).Condylocanine length(CCL): Distance rom the posterior margin o the occipital condyle to

the anterior ace o the canine.

Postorbital breadth (PB): Least breadth behind orbits, always posterior to postorbital bulge ipresent.Interorbital breadth (IB): Least breadth measured across orbitals, always anterior to postor-

bital bulge i present.Braincase breadth (BB): Greatest breadth o braincase.

Mastoid breadth(MB): Greatest breadth across mastoid region.Zygomatic breadth(ZB): Greatest breadth across zygomatic arches.

Maxillary toothrow length(MRL): Distance rom anterior ace o upper canine to posterior-most margin o last upper molar.

Mandibular toothrow length(MANDL): Measured rom anterior ace o lower canine to pos-terior margin o last lower molar. Includes m3 in V. melissaand V. sinchi, sp. nov.; m3missing in V. elisabethae, sp. nov.


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We tested or normality o the several samples o the univariate data using Shapiro-Wilkstests (Shapiro and Wilk, 1965). Seventeen ANOVA and ukeys tests or unequal sample sizeswere used to compare measurements between groups within the Vampyressa complex o spe-cies (V. melissa, V. sinchi, sp. nov., and V. elisabethae, sp. nov.). We then included the selectedexternal and skull measurements useul to discriminate between these groups to build correla-

tion matrices, together with samples o all Vampyressaspecies currently recognized, and per-ormed a principal-component analyses (PCA) or data reduction and visualization. Analyseswere perormed using Systat or Windows, version 11.0, and PAS (Hammer et al. 2001).

RESULS

All external metric characters (FA, MEIII, MEIV, MEV, iL) and six skull metric char-acters (GLS, CIL, CCL, MB, ZB, and MRL) were useul to discriminate among groups amongVampyressa pusilla, V. thyone,and the three orms o the Vampyressa melissacomplex (p


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rst and second components extracted rom the external measurements showed a clear separa-tion in size between the small (V. pusillaand V. thyone) and the large species o Vampyressa(V. melissa,V. sinchi, sp. nov., and V. elisabethae, sp. nov.). Te samples o the V. melissaand V.elisabethae, sp. nov., showed some size overlap but V. sinchi, sp. nov., was clearly segregatedrom all the others (g. 1). Along PC 1, all external metric characters varied equally and posi-tively with size, but PC 2 loadings indicated that iL was not proportional to FA, MEIII,MEIV, and MEV (table 2).

Te second PCA employing cranial dimensions or the same 55 specimens analyzed orexternal characters also resulted in the variation mostly explained by the two rst compo-nents (g. 2). Bivariate plots o the rst and second components extracted or skull measure-ments (g. 2) also showed clear separations in size between the small species o Vampyressa

(V. pusillaand V. thyone) and the large Vampyressa, and within species o the V. melissa-likespecies (g. 2). All skull metric characters varied equally and positively with size along PCI, but or skull characters, zygomatic breadth (ZB) was not proportional to the charactersdescribing skull length (e.g., MRL) and to the mastoid breadth (M) as demonstrated bythe variation along PC 2 (table 2).

Based on the morphometric results, and our study o several discrete characters that werevariable within the larger V.melissa complex and between these larger species and V. pusillaand V. thyone, we recognized two new orms that are described and named below. o clariythe diagnostic characters dening Vampyressa(sensu stricto) and to distinguish V. melissarom

other taxa diagnosed in this study, we provide emended diagnoses o both the genus Vampy-ressaand V. melissa.

ABLE 2. Factor loadings or the components extracted rom correlation matrices resulting rom the princi-pal-components analysis (PCA) o ve external and six skull characters comparing all the species oVampyressa.

External Characters

Axis

I II III IV V VI

FA 0.42 0.31 -0.16 -0.84 0.01

MEIII 0.43 0.10 0.75 0.12 0.48

MEIV 0.42 0.33 0.17 0.30 -0.77

MEV 0.41 0.32 -0.61 0.45 0.40

iL 0.54 -0.82 -0.14 0.00 -0.09

Skull Characters

GLS 0.39 -0.12 -0.01 -0.37 0.08 0.83CIL 0.46 -0.10 0.28 -0.33 0.63 -0.44

CCL 0.46 -0.07 0.29 -0.22 -0.77 -0.25

MB 0.32 -0.02 0.49 0.78 0.09 0.19

ZB 0.39 0.86 -0.33 0.08 0.02 -0.03

MRL 0.41 -0.49 -0.70 0.29 -0.01 -0.14


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SYSEMAICS

CLASS MAMMALIA

ORDER CHIROPERA

FAMILY PHYLLOSOMIDAE GRAY, 1825

SUBFAMILY SENODERMAINAE GERVAIS, 1856Genus VampyressaTomas, 1900

PhyllostomaWagner, 1843: 366; part; not PhyllostomaCuvier, 1800.Arctibeus: omes, 1860: 260; incorrect subsequent spelling, notArtibeusLeach, 1821.VampyressaTomas, 1900: 270; type species Phyllostoma pusillumWagner, 1843, by original designa-

tion; described as a subgenus o VampyropsPeters, 1865.VampiressaFesta, 1906: 7; incorrect subsequent spelling oVampyressaTomas, 1900.VamqyressaCabrera, 1958: 83; incorrect subsequent spelling o VampyressaTomas, 1900.VampiressaSilva, 1975: 52; incorrect subsequent spelling o VampyressaTomas, 1900.

VampuressaCoimbra, Borges, Guerra, and Mello, 1982: 34; incorrect subsequent spelling o Vampy-ressaTomas, 1900.

-4 -2 0 2 4

PC1 (96.5%)

-1.8

-1.2

-0.6

0

0.6

PC2(3.4

%)

thyone

pusilla

melissasinchi sp.nov.

elisabethae sp.nov.

FIG. 2. Plot o components 1 and 2 extracted rom the PCA analysis o skull measurements rom specimenso Vampyressa; the amount o the variation explained by each principal component is in parenthesis.


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VampiressaMontenegro and Romero-Ruiz, 2000: 646; incorrect subsequent spelling o VampyressaTomas, 1900.

VapyressaContreras-Vega and Cadena, 2000: 287; incorrect subsequent spelling o VampyressaTomas, 1900.

R D V: Te ollowing combination o characters denesthe genus Vampyressa as currently constituted (ve species recognized herein): wo pairs owhite acial stripes; a dorsal stripe is lacking; interemoral membrane usually conspicuouslyurred (hairs may be restricted to medial area) and ofen ringed; a tail is lacking; inner upperincisors relatively long, bid or weakly trid, and usually converging distally; accessory subapi-cal cusp on upper canines; caniniorm p2; p4 lacks a prominent anterior cingulum; secondlower molar (m2) has a well-developed, high, and bladelike entoconid; septum (medial angeo palatine processes o premaxillae) between incisive oramina perorated by large accessorymedial oramen; irregularly enestrated palate, oramen ovale relatively small and located wellanterior to the caudal extension o the pterygoid (the ridge orming the anterolateral border othe eustachial groove); portions o squamosal and exoccipital bones comprising posttympanicprocess unused; dental ormula 2/2, 1/1, 2/2, 2/23 2 = 2830.

C V: Vampyriscus, which we recognize as a separate genus,includes three species (Vampyriscus bidens, V. brocki, and V. nymphaea). Vampyriscuscan bedistinguished rom Vampyressaby the presence o a middorsal stripe (weakly developed insome V. nymphaeaand absent in some V. brocki); lack o extensive ur and absence o a ringeo hair on interemoral membrane; upper inner incisors long, pointed, and parallel in V.

brocki, or unevenly bid and convergent at the tips in V. bidens and V. nymphaea; normalcrowns o upper canines (lack subapical cuspule); p2 not caniniorm; p2 and p4 with well-developed anterior cingulae; entoconid o m2 not prominent as an elevated bladelike cusp;septum between incisive oramina narrow with a small medial accessory oramen in V.bidens, or very narrow and lacking the medial accessory oramen in V. brockiand V. nym-

phaea; palate comparatively at with scattered small, porelike perorations (not obviouslyenestrate as in Vampyressa); oramen ovale relatively large, adjacent to and partially occludedby high ridge o caudal extension o the pterygoid (in posterior view); bones comprisingposttympanic process used.

Vampyressa melissaTomas, 1926

Melissas Yellow-eared Bat

Figures 48

Vampyressa melissaTomas, 1926: 157, type locality Puca ambo, Peru, altitude 7100, San Martn,Peru.

Vampyressa (Vampyressa) melissa: Peterson, 1968: 14; name combination.

M: Te holotype BMNH 26.5.3.4 consists o a relatively well-preserved skinand skull o an adult emale rom Puca ambo (69S, 7716W), Moyobamba, department o


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San Martn, Peru, collected at 1480 m by R.H. Hendee (no. 408) on January 15, 1926, duringthe Godman-Tomas Expedition to Peru.

M H: Te ollowing measurements (mm) o the holotypeare rom the skin label: L 54, HF 10.5 s.u., E 16. Additional measurements (mm) taken byone o us (V.C..) rom the holotype are: FA 36.8, iL 14.9, GLS 21.2, CIL 20.2, CCL 19.5, PB5, IB 6.2, BB 8.9, BH 8.7, MB 10.4, ZB 12.5, MRL 6.9, and MANDL 7.4.

D: As dened here, Vampyressa melissais currently known rom upper-eleva-

tion localities (11802763 m) in Andean Colombia, Ecuador, and Peru (g. 3).E D: A large Vampyressa(FA 34.537.7, n = 9; GLS 21.222.6, n= 9; table

3) with a broad skull (ZB 12.513.5, n= 8; table 3); anteriorly tapering rostrum; discrete orbi-

FIG. 3. Map showing collecting localities o Vampyressa elisabethae,sp. nov. (circle), V.melissa(triangles), andV. sinchi,sp. nov. (squares). Numbers reer to entries in the specimens examined list (appendix).


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tosphenoid oramina, well-developed postglenoid processes that project ventrally to the levelo pterygoid processes; mesopterygoid ossa about as long as wide; large basioccipital-basisphe-noid pits, and lacks a well-developed posterolabial cuspulid on p4. V. melissais larger than V.

pusillaand V. thyone, but averages smaller in most dimensions than either o the new species

described below.R: Vampyressa melissahas sparse, long guard hairs conspicuous in the dorsal

pelage; two pairs o pale acial stripes with the medial stripes extending above eyes rom betweenears to upper border o nasal horseshoe, and the lateral stripes extending below eyes rom baseo ears to upper corners o mouth. Te pelage o the dorsum is uniormly brownish, a dorsalstripe is lacking. Te noselea (lancet) is triangular, elongate, and with deep grooves separatingthe lateral aps rom the midrib; bicolored with most o lateral margins o the lancet and outerborder o the horseshoe conspicuously pale yellow or cream color. Te ear pinna is hairy romits base to halway to the tip. Te proximal two thirds o the orearm is densely urred.

All light membranes are dark brown; the antebrachial membrane (propatagium) isnarrow and the posterior plagiopatagium inserts on the hind limb at the base o the toe.he uropatagium is relatively narrow, its greatest breadth approximately hal the length o

ABLE 3. External and craniodental measurements (mm), sample statistics,a and dental ormula (DF) oVampyressa melissa, V. pusilla, and V. thyone.

V. melissa V. pusilla V. thyoneSex 6 , 2 10 , 7 10 , 10

L 61.3 3.4 (5866) 5 50.3 (4853) 3 47.3 4.7 (4365) 20

HF 9.9 0.9 (8.911.0) 7 8.8 1 (69) 10 9 0.9 (710) 20

E 16.1 3.2 (1322) 6 12.9 1.7 (11.215.0) 9 14 1 (1216) 20

FA 36.7 1.1 (34.537.7) 9 33.4 0.6 (32.334.3) 16 30.6 0.7 (29.332.3) 19

MEIII 34.8 2.0 (33.737.3) 9 32.4 1.2 (29.934.2) 15 29 0.7 (2730) 19

MEIV 34.8 0.6 (34.137.0) 9 31.6 0.7 (29.832.5) 15 28.9 1 (27.132.3) 19

MEV 36.6 0.9 (35.138.2) 9 32.5 0.9 (30.633.9) 15 30.2 0.9 (28.732.3) 19

iL 13.7 0.6 (12.214.9) 9 12.4 1 (10.613.7) 5 10.4 0.5 (9.411.2) 19

GLS 21.8 0.5 (21.222.6) 8 19.8 0.4 (19.020.6) 15 18.2 0.6 (17.119.2) 19

CIL 20.3 0.5 (19.521.0) 8 18.1 0.5 (17.619.4) 14 16.4 0.6 (15.517.5) 19

CCL 19.6 0.6 (18.720.5) 8 17.5 0.5 (16.819.1) 15 15.9 0.6 (14.917.0) 19

PB 5.2 0.3 (4.65.7) 8 5.2 0.2 (4.785.6) 15 4.7 0.2 (4.45.0) 20

IB 6.2 0.4 (5.46.6) 7 5.5 0.2 (5.25.7) 10 5 0.2 (4.55.2) 20

BB 9.3 0.4 (8.810.0) 8 8.6 0.23 (8.39.1) 14 8.2 0.2 (7.78.6) 20

MB 10.6 0.3 (10.311.0) 8 9.6 0.3 (8.810.0) 15 9.1 0.2 (8.69.4) 19

ZB 13 0.3 (12.513.5) 8 11.5 0.3 (1112.1) 13 10.6 0.4 (9.511.2) 20

MRL 7.1 0.4 (6.77.9) 8 6.6 0.3 (6.17.3) 16 5.9 0.3 (5.56.5) 20

MANDL 7.7 0.3 (7.18.1) 8 6.9 0.2 (6.57.3) 16 6.2 0.3 (5.96.9) 20

DF i 2/2, c 1/1, p 2/2, m 2/3 i 2/2, c 1/1, p 2/2, m 2/2 i 2/2, c 1/1, p 2/2, m 2/2

aTe sample mean plus or minus the standard error o the mean, the observed range (in parentheses), and the samplesize are provided or each species.


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the tibia; its posterior border has an inverted U-shape. Dorsal and ventral suraces o theuropatagium are conspicuously haired and the posterior margin bears a paler ringe o hair

A

B

C

D

E

F

FIG. 4. Dorsal (A) and ventral (B) views o the skull o Vampyressa elisabethae(USNM 319283 [holotype];male) rom Bocas del oro, Panama. Dorsal (C) and ventral (D) views o the skull o V. melissa (FMNH174909; emale) rom Cuzco, Peru. Dorsal (E) and ventral (F) views o the skull o V. sinchi(FMNH 114028[holotype]; emale) rom Nario, Colombia. Scale bar = 5 mm.


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that is longest toward the midline; lower legs, adjacent uropatagium, and dorsal surace oeet densely covered with reddish-brown hair.

Te skull is robust; the rostrum is elevated anteriorly rom a horizontal plane connectingventral suraces o occipital condyles and crown apices o last upper molars; premaxillae inclinedposteriorly at approximately 60 rom the horizontal plane (g. 6B). Te angle o the oreheadwith the rostrum is not abrupt (~150) and the orehead slopes smoothly to the rostrum in lateraloutline. Te ventral margin o the external nares is V-shaped and bears a small medial pit (g.

5A, D). Te external nares are emarginate dorsally (nasals comparatively short; g. 5D). Terostrum tapers anteriorly and the postorbital constriction is narrower than interorbital breadth(g. 4C). Te sagittal crest is low, but well developed; the posterior border o the skull (inteparietalregion) is not conspicuously inated (g. 6B). Te dorsoventrally thin septum separating thetriangular-shaped incisive oramina is perorated by a large accessory medial oramen (g. 5D).Te lightly and irregularly enestrate hard palate is conspicuously domed behind the incisiveoramina. Te shape o the emargination o the posterior border o palate is variable, yet themesopterygoid ossa is approximately as long as wide (gs. 4D, 8A). A discrete orbitosphenoidoramen is present. Te hamular process o the pterygoid and the oramen ovale are approxi-mately even with (on same lateral plane as) the robust postglenoid process. Te paired basioc-cipital-basisphenoid pits are long and extend anterior to anterior margin o bullae. Te caudal

p

mp

p amf ps

A B C

FED

FIG. 5. Frontal view and dorsal view o external nares o the skulls o V. melissa, USNM 548310 (A, D); V.sinchi, sp. nov., FMNH 114028 [holotype] (B, E); V. elisabethae, sp. nov., USNM 319285 (C) and USNM

319284 (F). Abbreviations: amf,accessory medial oramen; mp,mesethmoid plate; p,median premaxillarypit; s, septum between incisive oramina. Scale bar = 5 mm.


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extension (ridge) o each ptery-goid is directed posterolaterally

and well separated rommedium-sized bony excres-cences (g. 8A). Te oramenmagnum is rounded in outline.

Crowns o upper innerincisors (I1s) are large, approxi-mately evenly bilobate, con-

verge apically, and havewell-separated roots; outerupper incisors (I2s) also arebilobate, but much smaller,extending to or slightly beyondcingulae o upper canines (g.5A), and anterior portions are

visible in dorsal view o theskull. Tere are two subequaland weakly bid lower incisorson each side o the mandible.

Te upper canine (C) is

about twice the length o P4,and has a conspicuous smallcusp at the apex o the medio-lateral ridge orming the lin-gual border o the longitudinalsulcus on its posterior crown.P3 is about a hal or less thesize o P4, which is nearly tri-angular, has a well-developed

labial cingulum and a smallposterior cuspule paired with asmaller cuspule on the postero-labial cingulum. he lower

A

C

B

FIG. 6. Lateral views o the skull andlower jaw o (A) Vampyressa elisabe-thae (USNM 319283 [holotype];male). Lateral views o the skull andlower jaw o (B)V. melissa (FMNH174909; emale). Lateral views o theskull and lower jaw o (C) V. sinchi

(FMNH 114028 [holotype]; emale).Scale bar = 5 mm.


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canine (c) has a well-developed posterior cingulum; p2 strongly resembles the lower canine,although less than hal its size; p4 has a ridge along the posterior crown o the major cusp thatextends across the heel o the tooth to the posterior cingulum, but lacks a discrete posterolabialcuspulid (gs. 6B, 7A).

Te rst upper molar (M1) has a relatively well-developed protocone, paracone, andsmaller metacone. Te second upper molar (M2) has a rudimentary protocone, a much-reduced metacone, and a small protoconule behind the large paracone. Te rst lower molar

(m1) is quadrate in occlusal outline, with a large protoconid and a small blunt hypoconid. Tesecond lower molar (m2) has a long metaconid that is twice the size o the protoconid and alarge, L-shaped (bladelike) entoconid approximately the same size as the metaconid. Tomas(1926: 1) commented on the lower molars o V. melissa: Its peculiarly shaped m1, and theunusual development o the characteristic cusps on m2 all readily distinguish it rom the otherspecies o the genus [sensu lato]. A small peglike m3 is present.

C: Previously, V. melissawas compared with presumed congeners Vampyriscusbidensand V. nymphaea. Goodwin (1963: 23), or example, briey redescribed the holotype oV. melissa and commented that the skull was about like that o Vampyriscus nymphaeabut

more robust. However, these two taxa along with Vampyriscus brockiare not species o Vampy-ressaand need not be compared herein (see diagnosis or Vampyressa).

ABLE 4. External and craniodental measurements o the type series o Vampyressa sinchi,sp. nov. Measure-ments are in millimeters, except weight, in grams.

Character HolotypeFMNH114028

ParatypeMHNUC1514

ParatypeIAvH2282

Weight 16 g

L 70 70 68

HF 13 11 11.6

E 17 18

FA 40.5 41.5 39.1

MEIII 40.0 38.0 39.0

MEIV 38.2 38.6 37.9

MEV 38.9 40.3 38.8iL 15.0 15.9 15.1

GLS 23.7 23.6 23.3

CIL 22.7 22.3 21.9

CCL 22.0 21.6 21.3

PB 5.3 5.3 5.5

IB 6.3 5.6 6.0

MB 11.2 11.4 10.8

ZB 14.2 14.8 13.9

MRL 7.6 7.5 7.7MANDL 8.8 8.7 8.7


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Vampyressa melissa is considerably larger than either o the other two known species oVampyressa(contrast measurements o V. melissawith those o V. pusillaand V. thyonein table

3). V. melissadoes merit comparison with each o the two new taxa described below (see thosesections or comparisons).

During research on her doctoral dissertation, the lead author realized that Colombianspecimens identied as V. melissa rom upper elevations in the Andes were larger and morerobust than V. melissa, and appeared to be undescribed. Here we describe this species as:

Vampyressa sinchi, new species

Quechuan Yellow-eared Bat

Figures 46, 810

Vampyressa melissa:Lemke et al., 1982: 231.Vampyressa melissa: Alberico et al., 2000: 55 (part).

H: An adult emale (FMNH 114028; gs. 4EF, 5B, 5E, 6C, 8B, 9A) collected atLlorente (04640N, 772150W; 1700 m), municipio de Crdoba, departamento de Nario,Colombia, by Kjell von Sneidern (original number: 1736) on May 10 1971. Te holotype ispreserved as a skin and skull (table 4).

P: An adult emale (MHNUC 1514, g. 10) caught at San Juan de Villalobos

(13318N, 761819W; 1620 m), Vereda La Esmeralda, Santa Rosa, departamento de Cauca,Colombia, in the Caquet River basin, eastern Colombian slopes o the eastern Andes by H.E.Ramrez-Chaves on 5 May 2005; and an adult male (IAvH 2282) collected in Parque NacionalNatural Cueva de los Gucharos (138 N, 7558W, 1900 m), Acevedo, departamento deHuila, Colombia, by .O. Lemke (Lemke et al. 1982) see table 4.

D: Know only rom the type series rom the eastern slope o the Central Cor-dillera (Llorente) and eastern slopes o the Eastern Cordillera (San Juan de Villalobos andParque Nacional Cueva de los Gucharos) o the Andes in Colombia (g. 3).

E: A Quechuan word, sinchiconveys the meaning robust and strong. Te namehonors the Quechuan people o Colombia (the Ingas), and indicates the robustness o this spe-cies, the largest Vampyressaknown. Te name is to be treated as a noun in apposition.

FIG. 7. Rami o V. melissa, USNM 548310 (A) and V. elisabethae,sp. nov., USNM 319284 (B). Note spiculelikem3 in V. melissa (lacking in V. elisabethae) and posterolabial cusp on p4 in V. elisabethae (lacking in V.melissa). Scale bar = 5 mm.

A B


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D: Vampyressa sinchiis similar to V. melissa, but larger in most dimensions withew overlapping measurements (tables 34, gs. 1, 2, 4E, 4F, 6C). Te skull, including dentition,is larger and more robust than any other known species o Vampyressa. Te strongly developedpostglenoid processes extend below the level o pterygoid processes; the mesopterygoid ossa

is longer than wide; the ventral surace o the postdental extension o the palate is at, its mar-gins comparatively sharp, and bears small winglike projections on each side at approximatelythe level o the orbitosphenoid oramen.

D:Vampyressa sinchihas pale brown to dark-beige dorsal pelage; the darker dorsumcontrasts with the paler venter; short, sparse guard hairs on the head and dorsum; orearm denselyhaired proximally or about two thirds o its length; the uropatagium is narrow (about hal thelength o the tibia in dried skins), urred on both suraces, and has a sparse ringe o hairs along itsentire posterior margin but longest toward the midline; tibia, adjacent uropatagium, and oot con-spicuously covered with reddish-brown hairs similar to the condition in V. melissa. Te third and

fh metacarpals are similar in length and the ourth metacarpal averages shorter than the thirdand fh. Otherwise, V. sinchiis very similar externally to V. melissaas described in that account.Te skull o Vampyressa sinchi (gs. 4E, 4F, 6C) also is similar to V. melissa, but more

robust, and larger in most dimensions with ew overlapping measurements (tables 34). Temedial, incisor-bearing anterior premaxillae o V. sinchiare inclined at approximately 60 romthe horizontal plane. Te external nares are emarginate dorsally; the ventral margin is V-shapedbetween the roots o the upper inner incisors and bears a medial pit (gs. 5B, 5E). Te nasalsare relatively short resulting in a comparatively longer narial opening (deeper dorsal emargina-tion, g. 5E). Te mesethmoid plate is not swollen and its anterior edge is thin (g. 5B).

Te rostrum is elevated above the horizontal plane connecting occipital condyles and thetips o the last upper molars. Te septum between the incisive oramina is dorsoventrally thin

FIG. 8. Partial views o the basicranial region o V. melissa, USNM 548310 (A); V. sinchi, sp. nov., FMNH114028 [holotype] (B); V. elisabethae,sp. nov., USNM 319282 (C). Abbreviations: be,bony excrescence; cb,extension o caudal border o pterygoid; fo,oramen ovale; pg,postglenoid process; sf,orbitosphenoid ssure.Scale bar = 5 mm.

CA B C

s

cbe

cb fo

pg sf

cb fo


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and perorated by a large accessory medial oramen (g. 5E). Te lightly and irregularly enes-trated palate is conspicuously domed behind the incisive oramina. Te sagittal and lambdoidalcrests are low, but well developed; the interparietal-supraoccipital region is not conspicuouslyinated. Te mesopterygoid ossa is longer than wide; basioccipital-basisphenoid pits aresharply dened, deep, and extend well anterior to the anterior level o the auditory bullae.

Te upper inner incisors (I1s) are subequally bilobed and convergent at the tips. Te dentitionis similar to that o V. melissa, but more massive throughout with stronger cingular development.C:Vampyressa sinchiis larger than all other Vampyressaknown (gs. 12, tables

34; avares et al., 2008) andtends to have longer ur and a darker dorsum. Among species oVampyressa, V. sinchiis most similar to V. melissarom which it differs primarily in larger size;heavier, more robust dentition; unevenly bilobed upper inner incisors, deeper dorsal emargina-tion o anterior nares, longer than wide mesopterygoid ossa, and a distinctly larger and longerpostglenoid process that extends below the level o the hamular process o the pterygoid.

Like V. melissa, V. sinchihas sloping premaxillae (approximately 60 rom horizontal plane)that bear a pit located in the midline between roots o upper inner incisors; narrow anterior

margin o mesethmoid plate (g. 5E); anteriorly tapering rostrum; dorsoventrally thin septumseparating wider than long incisive oramina and perorated by a large accessory medial oramen;

ABLE 5. External and craniodental measurements o the type series o Vampyressa elisabethaesp. nov. (inmm)

Character HolotypeUSNM319283

ParatypeUSNM319282

ParatypeUSNM319284

ParatypeUSNM319285

ParatypeUSNM520851

L 581 532

HF 9.31 10.91 9.71 10.51 10.52

E 162

FA 37.8 36.8 36.2 37.8 37.2

MEIII 35.5 35.6 35.4 35.2 34.7

MEIV 35.7 35.4 35.5 35.3 35.2

MEV 37.0 37.2 36.1 36.5 36.1

iL 12.3 12.7 12.4 12.4 12.9GLS 22.9 22.9 22.9 23.0 22.9

CIL 21.4 21.3 21.6 21.6 21.5

CCL 20.7 20.7 20.8 20.8 20.8

PB 5.2 5.2 5.4 5.3 5.3

IB 6.3 5.9 6.0 6.1 5.7

BB 9.3 9.0 9.1 9.2 9.4

MB 10.9 10.6 10.7 10.9 11.0

ZB 12.6 12.0 12.0 12.1 12.4

MRL 7.8 7.7 7.7 7.9 7.7MANDL 8.6 8.3 8.5 8.1 8.4

1Measurements rom dried skin.2Measurements rom uid-preserved specimen.


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conspicuously domed hard palate behind incisive oramina; orbitosphenoid oramina (g. 9A);oramen ovale and ventral projection o hamular process o pterygoid on same lateral plane asthe base o the postglenoid process (g. 8B); large basisphenoid-basioccipital pits that extend wellanterior to anterior suraces o auditory bullae; caudal ridge rom pterygoid well separated romcomparatively small boney excrescences (located on basisphenoid anteromedial to auditory bul-lae) (g. 8B); rounded oramen magnum; robust mandibular rami and angular process. Like V.melissa, in V. sinchithe outer upper incisors are visible rom above; p4 has a posterior ridge thatextends down rom the crown o the major cuspid and across heel to the posterior cingulum, andp4 also lacks a posterolabial cuspulid. V. sinchiand V. melissaare the only species o Vampyressathat retain the spiculelike last lower molar; the dental ormula is 2/2, 1/1, 2/2, 2/3 2 = 30.

Te late Charles O. Handley, Jr., acquired our specimens o a large Vampyressarom theBocas del oro region in the 1960s during eldwork in Panama related to a countrywide survey

o the ectoparasites o vertebrates, primarily mammals. Although cataloged as Vampyressamelissa, Handley (1966: 767) reported these specimens as Vampyressaspecies, but never gotaround to describing them. An additional specimen rom the same region was acquired byRobert K. Ender in 1976. Tese specimens are described here as:

Vampyressaelisabethae, new species

Elisabeths Yellow-eared Bat

Figures 49Vampyressa species Handley, 1966: 767.

FIG. 9. Oblique views o the basicranial region o V. sinchi,sp. nov., FMNH 114028 [holotype] (A); V. elisa-bethae,sp. nov., USNM 319282 (B). Abbreviations: be,bony excrescence; cb,extension o caudal border opterygoid; fo,oramen ovale; of,orbitosphenoid oramen; sf,orbitosphenoid ssure. Scale bar = 5 mm.

of fo

cbbe

sf

fo

cb be

A B


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H: An adult male (USNM 319283; gs. 4AB, 6A) caught in evergreen orests oRancho Mojica, upper Changena River, 20 miles [32.2 km] SSW o Changuinola (96 N,8234W; 1463 m), Provincia Bocas del oro, Panama, by Vernon J. ipton (original number8006) on 13 September 1961. Te holotype is preserved as a skin and skull (table 5).

P: Te skins and skulls o one adult emale (USNM 319282; gs. 8C, 9B) and twoadult males (USNM 319284 [gs. 5F, 7B], 319285 [g. 5C]) caught at Rancho Mojica, upperChangena River, 20 miles SSW o Changuinola (096 N, 08234 W, 1463 m), ProvinciaBocas del oro, Panama (original numbers V.J. ipton 8006, 8120, 8122). A uid-preservedspecimen o a male (USNM 520851), with skull removed, collected by Robert K. Enders atFish Camp (0858N, 08240W, 1494 m), Bocas del oro, Panama on 26 March 1976.

D: Know only rom provincia de Bocas del oro, Panama (g. 3).E: Te name elisabethaehonors the legacy o the late Elisabeth Klara Victoria Kalko

in the study o the natural history, ecology, conservation, vocal recognition, and behavior o bats.D: A large Vampyressa(FA 36.237.8, n= 5; GLS 22.923.0, n= 5; table 5) having

a relatively long, narrow skull (ZB 12.012.6, n= 5; table 5) with nearly parallel sides o the ros-trum. Externally and dentally similar in appearance and size to V. melissaand V. sinchi, V. elisa-

FIG. 10. Photograph o an adult emale Vampyressa sinchi(MHNUC 1514) captured at San Juan de Villalobos,Caquet River basin, eastern Colombian slopes o the Eastern Cordillera, collected at 1620 m. Photograph byOelia Meja-Egas.


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bethaehas a swollen anterior margin o mesethmoid plate (g. 5C); dorsal contour o externalnares not deeply emarginate (g. 5F); an orbitosphenoid ssure (instead o a discrete oramen,

gs. 4B, 9B); caudal border o pterygoid extending posteriorly as a low ridge over basisphenoidto near contact with the large bony excrescence (gs. 8C, 9B); laterally ovoid oramen magnum;unequally bilobate to weakly trilobate upper inner incisors (g. 5C); a small cuspulid on postero-labial crown o p4 (g. 7B); m3 absent; dental ormula 2/2, 1/1, 2/2, 2/2 2 = 28.

D: Vampyressa elisabethaeis closely similar in color pattern and other externaleatures to both V. melissaand V. sinchi, although slightly smaller than V. sinchiand averagingequal to or slightly larger than V. melissa. On average, V. elisabethaehas slightly paler dorsalpelage (pale ochre and yellowish brown to beige) than do either V. melissaor V. sinchi. Teorearm is densely urred or a little more that its proximal hal, and dorsal and ventral suraceso the propatagium and plagiopatagium are hairy along sides o the body. Te dorsal and ven-tral uropatagium, legs, and eet are sparsely urred, except toward the ventral midline o theuropatagium, where the ur is especially dense and long. Te uropatagium lacks a conspicuousringe o hair along the entire posterior margin. Like V. melissaand V. sinchi, the uropatagiumis relatively narrow, its greatest breadth measuring less than hal the length o the tibia. Tethird and ourth metacarpals are subequal, and the fh metacarpal is longer than either thethird or ourth. Te tibia is short (12.312.9, n= 5) relative to the overall size o the bat.

Vampyressa elisabethaeis cranially and dentally similar to V. melissaand V. sinchi, but canbe distinguished by its relatively longer, narrower skull; nearly parallel sides o the rostrum;nearly straight prole o rontals and rostrum in lateral view (g. 6A); less convergent upper

toothrow; and narrower zygomatic breadth (ZB 12.012.6, n= 5). Te anterior plane o pre-maxillae is near vertical (80) rom the horizontal plane o skull; the rostrum is aligned withthe cranium (tips o canines and second upper molars on same horizontal plane as ventralsuraces o occipital condyles); lower margin o anterior external nares U-shaped and lackinga medial pit between roots o upper inner incisors; anterior margin o mesethmoid plate swol-len (g. 5C); dorsal contour o external nares not deeply emarginate (g. 5F); an orbitosphe-noid ssure present (gs. 4B, 9B); incisive oramina comparatively round with medial septumdorsoventrally thick and perorated by a comparatively small accessory medial oramen; palatenot conspicuously domed behind incisive oramina; short, shallow basioccipital-basisphenoid

pits; caudal border o pterygoid extending posteriorly as a low ridge to near contact with largebony excrescence on basisphenoid; oramen magnum laterally ovoid (g. 8C); ventral suraceo postdental palate weakly depressed (indented) along ventral midline and its lateral marginsrounded along at least part o their length; oramen ovale and pterygoid process located behindmedial base o postglenoid process; each dentary is proportionally slender, dorsoventrally nar-row, has a low coronoid process, and a small articular process that is close to the small angularprocess; upper toothrows not strongly convergent anteriorly; unequally bilobate to weakly tri-lobate upper inner incisors; upper outer incisors not easily visible in dorsal view o the skull;a posterolabial cuspulid present on p4, but lacking a medial ridge rom major anterior cuspidto cingulum on heel; m3 absent; the dental ormula is 2/2, 1/1, 2/2, 2/2 2 = 28.

C: Vampyressa elisabethaeneeds no comparison with the much smaller V.pusillaand V. thyone. Although externally similar to V. melissaand V. sinchi, V. elisabethae


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has somewhat paler pelage, less hairy eet and uropatagium (except ventrally near mid-line), and shorter tibiae. he skull is conspicuously narrower and the anterior premaxillae

more vertically oriented (near 80 rom the horizontal plane versus approximately 60 inV. melissa and V. sinchi). he anterior nasal cavity is considerably dierent in that theventral margin in V. elisabethaeis U-shaped and lacks a medial pit (V-shaped with a medialpit between roots o upper inner incisors in the other two taxa), less emarginate upperborder (deeper emargination in V. melissaand the much more deeply emarginate borderin V. sinchi; ig. 5), and swollen anterior margin o the conspicuously thicker mesethmoidplate (plate thinner in V. melissa and V. sinchi; ig. 5). he sphenorbital opening in thecranium is a issure in V. elisabethae in contrast with all other species o Vampyressa inwhich the opening is a oramen. he morphology o the anterior palate diers between V.elisabethae, V. melissa, and V. sinchi (ig. 5) and the palate is domed behind the incisiveoramina in V. melissa and V. sinchi, but not domed in V. elisabethae. he oramen mag-num is transversely ovoid in V. elisabethae, but more rounded in all other Vampyressa.Each dentary is massive, the coronoid process high, and both articular and angular pro-cesses are larger and not particularly close together in V. melissaand V. sinchi(ig. 6BC),whereas in V. elisabethaeeach dentary is proportionally more slender, dorsoventrally nar-row with a low coronoid process, and the smaller articular process is closer to the smallerangular process (ig. 6A).

Vampyressa melissaand V. sinchi obviously are closely related as reected by their closesimilarity. V. elisabethaeis not as closely related to these two species, nor is it closely related to

V. pusillaand V. thyonebased on its clearly distinctive eatures. Nevertheless, all known specieso Vampyressashare the same major dental eatures, particularly the caniniorm p2 and thecharacteristic well-developed, high, and bladelike entoconid on m2.

NAURAL HISORY AND CONSERVAION

Vampyressa melissa was listed as vulnerable by the most recent IUCN Global MammalAssessment because o presumed population decline and shrinkage in distribution inerredrom habitat degradation and conversion (avares et al., 2008). Te recognition that the popu-

lations previously identied as V. melissacomprise three distinct species reinorces the percep-tion o threatened status. Tese three species o rugivorous bats are uncommonly encountered,and occur in ragmented, higher-elevation habitats. One Ecuadorian specimen o V. melissa(USNM 548310) was pregnant when captured on 12 February 1983 with a etus having acrown-to-rump length o 27 mm.

Te rarity o Vampyressa melissaand its allies constrains study o their variation and obvi-ously hinders the possibility o gaining knowledge about their biology. On the other hand, V.melissahas been recorded in three instances rom 1970 to 1990 in habitats described as dis-turbed. Gardner (1976) and Patton et al. (1982) reported collecting Peruvian specimens o V.melissain lower-elevation humid orest o different secondary stages along the eastern slope othe Andes, and one Ecuadorian specimen came rom a disturbed orest habitat (Albuja-V.,


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1991). Te supposed rst specimen o V. melissareported rom Colombia, collected in highlandcloud orest (Lemke et al., 1982), is actually V. sinchi.

Te distribution o each o the three large Vampyressaspecies suggests delity to a particu-lar high elevational range, an unusual pattern among stenodermatines. Graham (1983) reportedelevational limits or V. melissa in Peru as 10001800 m, based on the scanty inormationavailable at that time. Tese species appear to be distributed at elevations above 1200 m andpossibly range as high as or higher than 2600 m. Because these bats are rugivores, the primaryactor limiting their distribution is the year-round availability o consumable ruit. Tat actoralone places these species in jeopardy because o continued habitat conversion and destructionat the elevations where these Vampyressaoccur.

AUHOR CONRIBUIONSTis report was redrafed by V.C.. based on her revisionary work o Vampyressa(s.s.) as

part o her doctoral dissertation. A.L.G. prepared gures 5, 7, 8, and 9; wrote most o thedescription o the new species rom Panama; and contributed to the redraf the manuscript.H.E.R.C. provided measurements and observations on the paratypes o V. sinchihoused in theColombian museums, and contributed with natural history inormation in a previous draf.P.M.V. prepared gures 3, 4, and 6, and contributed to the analyses.

ACKNOWLEDGMENS

Tis work was part o V.C..s doctoral studies at City University o New York in a joint pro-gram with the American Museum o Natural History; V.C.. is grateul to the committee mem-bers N. Simmons, C. Raxworthy, J. Wahlert, R. Rockwell, and R. Voss; special thanks to N.Simmons or her careul review o an earlier draf o this paper. Te ollowing curators and col-lection staff kindly permitted access to specimens under their care: R. Jenkins and P. Jenkins(BMNH); E.M. Versute (DZSJRP); B.D. Patterson (FMNH); J.E. Castillo (IAvH); H. Lpez Arvaloand Y. Muoz-Saba (ICN); R.M. imm (KU); M. Rivas Pava (MHNUC); L. Albuja and J. Regalado(MPEN); V. Pacheco (MUSM); J.L. Patton (MVZ); M. de Vivo (MZUSP); L. Gordon (USNM);

O.M. Garcia (UV), B. Lim and M. Engstrom (ROM), R.J. Baker (U), and P. Myers (UMMZ).We are grateul to Oelia Meja-Egas or allowing us to use her photograph o a live individual oVampyressa sinchi. Tis material is based upon work supported by the NASA under awardsNAG5-12333 and NAG5-8543, the New York Community rust, S. Donnelley, and the Centeror Biodiversity and Conservation at the American Museum o Natural History to V.C..; V.C..also thanks CAPES and FAPEAM postdoctoral ellowship under the program PRO-DPD/AMPPGSS. o those people and institutions we express our sincere thanks. We also are grateul to. Chesser, USGS-PWRC, Biological Survey Unit, National Museum o Natural History; B.D.Patterson, Field Museum o Natural History; one anonymous reviewer; and to R.S. Voss or theircomments and recommendation. Any mention o trade, product, or rm names is or descriptivepurposes only and does not imply endorsement by the United States government.


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REFERENCES

Alberico, M., A. Cadena, J, Hernndez-Camacho, and Y. Muoz-Saba. 2000. Mameros (Synapsida:

Teria) de Colombia. Biota Colombiana 1 (1): 4375.Albuja-V., L. 1991. Mameros, lista del vertebrados del Ecuador. Politcnica 16: 163204.Arroyo-Cabrales, J. 2008a (2007). Genus VampyressaO. Tomas, 1900. InA.L. Gardner (editor), Mam-

mals o South America, vol 1: marsupials, xenarthrans, shrews, and bats: 346350. Chicago: Uni-versity o Chicago Press.

Arroyo-Cabrales, J. 2008b (2007). Genus VampyriscusO. Tomas, 1900. In A.L. Gardner (editor),Mammals o South America, vol 1: marsupials, xenarthrans, shrews, and bats: 350355. Chicago:University o Chicago Press.

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APPENDIX

S E

Te ollowing list includes all the specimens used in this study, with their respective locali-ties. See Material and Methods or abbreviations.

Vampyressa elisabethae,sp. nov. (5): PANAMA: Bocas del oro: Changuinola, 20 miles SSW,upper Changena River, Rancho Mojica, 906 N, 8234 W (USNM 319282, ; 319283,

[holotype]; 319284, ; 319285, ); Fish Camp, 858N, 8240W (USNM 520851, ).Vampyressa melissa(11): COLOMBIA: Norte de Santander: Parque Natural am, 706N,

7213W (IAvH 6761, ). ECUADOR: Pastaza: Mera, 122S, 7801W (USNM 548310, ).Napo: Cantn Archidona, Parroquia Cotundo, Ro Urcusiqui, 049S, 7747W (MEPN 5392,). Zamora-Chinchipe: km 30 va Loja-Zamora, Estacin Cientca San Francisco, 0353S,7905W (MEPN 7578, ). PERU: Cuzco: Bosque de las Nubes, carretera Paucartambo-Pill-copata, km 150, Puente Union, 1322S, 7161W (MUSM 8914, ); Quitacalzn, carreteraPaucartambo-Pillcopata, Puente at km 163, 1299 S, 7145 W (MUSM 8915, juvenile ;MUSM 8916,); Paucartambo, Consuelo, 15.9 km SW Pilcopata, 1302S, 7149W (FMNH

174909, ). Huanuco: Cerros del Sira, 930S, 7447W (AMNH 233761, ; AMNH 233769,). San Martn: Moyobamba, Puca ambo, 69 S, 7716 W (BMNH 26.5.3.4, [holotype]).

Vampyressa pusilla (17): BRAZIL: Minas Gerais: APA Coqueiral, 2111 S, 4526 W(CMUFLA 165, ; CMUFLA 166, ); Apolo 21, Raposos, 1957S, 4348W (CMUFLA 167,; CMUFLA 569,); Lavras, 2114 S, 4459 W (CMUFLA 484, ; CMUFLA 530, );Minduri, 2139 S, 4436 W (CMUFLA CPM 26, ); Viosa, 2044 S, 4250W (USNM395703, ). So Paulo: Canania, 2501S, 4755 W (MZUSP 22215, sex unknown); IlhaAnchieta, 2331S, 4502(MZUSP 29438, ); Ilha do Cardoso, 2511S 4759 W (MZUSP27729, ; 28132, ); Mirassol, Stio Progresso, 2049S, 4931W (DZSJRP 3920, ); Varjodo Guaratuba, 2345S, 4553W (MZUSP 21081, ); Ipanema, 2327S, 4735 W (RMNH


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17256, ). PARAGUAY: Paraguar: Parque Nacional Ybycu, 2601 S, 5703 W (UMMZ133730, ; 133731, ).

Vampyressa sinchi,sp. nov. (3): COLOMBIA: Cauca: Santa Rosa, Vereda La Esmeralda SanJuan de Villalobos, 13318 N, 761819 W (MHNUC 1514, ). Huila: Acevedo, ParqueNacional Natural Cueva de los Gucharos, 138N, 7558W (IAvH 2282, ). Nario: Cr-doba, Llorente, 04640N, 772150W (FMNH 114028, [holotype]).

Vampyressa thyone (20): COLOMBIA:Antioquia: 24 km S and 22 km W, near La irana,520 m, ca. 718N, 7505W (USNM 499464, ; 499466,; 499467, ). Meta: Finca ElBuque, Villavicencio, 409 N, 7339 W (USNM 507198, ). COSA RICA: Puntarenas:Corcovado N.P., Osa Peninsula, Quebrada Camaronal, Sirena, 833 N 8335 W (USNM565811, ). ECUADOR: El Oro: Puente de Moromoro, 1 km SW, 920 m, 344S, 7944W(USNM 513466, ; 513467 , ; 513469, ). PANAMA: Canal Zone:Barro Colorado Island,0909N, 7951W (USNM 514966, ; 514967, ); Bohio Peninsule, 0911N, 7950 W(USNM 503634 ; 503629, ; 503632, ; 503633, ; 503636, ). PERU: Cuzco: Pagoreni, 465m, 1142S, 7254W (USNM 582870, ). VENEZUELA:Apure: Nulita, Selvas de San Camilo,29 km SSW o Santo Domingo, 24 m, 719N, 7157W (USNM 4407, ). Yaracuy: Minas deAroa, Bolvar R.R., 1025, 6854W (USNM 440660, ; 440733, ; 440734, ; 440736, ).


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