revisionary notes on neotropical porcupines (rodentia

Copyright q American Museum of Natural History 2001 ISSN 0003-0082

P U B L I S H E D B Y T H E A M E R I C A N M U S E U M O F N AT U R A L H I S T O RY

CENTRAL PARK WEST AT 79TH STREET, NEW YORK, NY 10024

Number 3351, 36 pp., 15 figures, 5 tables October 31, 2001

Revisionary Notes on Neotropical Porcupines(Rodentia: Erethizontidae). 2. A Review of the

Coendou vestitus Group with Descriptions of TwoNew Species from Amazonia

ROBERT S. VOSS1 AND MARIA N. F. DA SILVA2

CONTENTS

Abstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2Resumen . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2Resumo . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

Materials and Methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 3Species Accounts . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6

Coendou vestitus Thomas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6Coendou pruinosus Thomas . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11Coendou ichillus, new species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17Coendou roosmalenorum, new species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24

Discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 30Acknowledgments . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 32Appendix: Gazetteer . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 34

1 Division of Vertebrate Zoology (Mammalogy), American Museum of Natural History. E-mail: [email protected] Instituto Nacional de Pesquisas da Amazonia, CP 478, Manaus, AM 69083, Brazil. E-mail: [email protected]

2 NO. 3351AMERICAN MUSEUM NOVITATES

ABSTRACT

Neotropical porcupines of the Coendou vestitus group consist of four small-bodied speciesfrom northwestern South America that are hypothesized to form a clade on the basis of theirunique dorsal pelage composition. In addition to soft wool and barbed quills—hair typeswidely shared by other erethizontids—the dorsal pelage of vestitus-group porcupines includeswhat may be called bristle-quills: long, thin, unbarbed quills with flexible tips. The groupincludes Coendou vestitus Thomas, 1899; C. pruinosus Thomas, 1905; and two new species.The latter provide the first documented records of small porcupines from western Amazonia,where only large porcupines (C. prehensilis and C. bicolor) were previously known. Othersmall porcupines probably remain to be discovered in Amazonia, where hydroelectric damprojects offer unique (if unfortunate) faunal-sampling opportunities.

RESUMEN

El grupo de puercoespines neotropicales Coendou vestitus consiste en cuatro especies pe-quenas del noroeste de America del Sur, consideradas parte de un clado con base en la com-posicion particular de su pelaje dorsal. Ademas de lana suave y espinas con puas—tipos depelo compartidos ampliamente con otros eretizontidos—el pelaje dorsal de los puercoespinesdel grupo vestitus tiene lo que podrıa llamarse cerda-espinas: espinas largas, delgadas y ralas,con puntas flexibles. Este grupo incluye a Coendou vestitus Thomas, 1899; C. pruinosusThomas, 1905; y dos nuevas especies. Estas ultimas constituyen los primeros registros docu-mentados de puercoespines pequenos en la Amazonia occidental, donde previamente solo seconocıan puercoespines grandes (C. prehensilis y C. bicolor). Se discute la posibilidad de queotros puercoespines pequenos esten por descubrirse en la Amazonia y se enfatizan las opor-tunidades unicas de muestreo de fauna provistas por la construccion de represas hidroelectricas.

RESUMO

Porcos-espinhos neotropicais do grupo Coendou vestitus sao representados por quatro es-pecies de pequeno porte, que hipotetizamos formar um clado, tendo por base a composicaounica de sua pelagem. Alem da pelagem lanosa e dos espinhos farpados—tipos de pelosamplamente compartilhados por outros eretizontıdeos—a pelagem dorsal dos porcos-espinhosdo grupo vestitus incluem o que podemos chamar de cerdas espinhosas: longos, finos, espinhosnao-farpados de pontas flexıveis. O grupo inclui Coendou vestitus Thomas, 1899; C. pruinosusThomas, 1905; e duas novas especies. Estas fornecem o primeiro registro documentado deporcos-espinhos de pequeno porte no oeste da Amazonia, onde apenas porcos-espinhos degrande porte (C. prehensilis e C. bicolor) eram anteriormente conhecidos. Discutimos a prob-abilidade de que outras especies de porcos-espinhos de pequeno porte sejam descobertas nooeste amazonico, e enfatizamos a importancia unica de amostragens faunısticas criadas pelaconstrucao de barragens hidroeletricas.

INTRODUCTION

Conspicuous among the Neotropical pre-hensile-tailed porcupines currently referredto the genus Coendou (including Sphiggurus;Handley and Pine, 1992) is a group of small-bodied species whose dorsal pelage containsthree intermingled but distinctly differentstructural types: soft fur, quills (defensivespines with hard, sharp, barbed tips), and anunusual structural type that may be called

bristle-quills (fig. 1). The latter are much lon-ger and thinner than conventional quills, buthave the same distinctive root morphology(described and illustrated by Chapman andRoze, 1997), round cross section, stiff cortex,and spongy internal matrix; unlike regularquills, however, the tips of bristle-quills areattenuated, flexible, wire-like filaments, andthey are not provided with microscopicbarbs.

2001 3VOSS AND DA SILVA: NEOTROPICAL PORCUPINES

Fig. 1. Three structural types found in thedorsal pelage of members of the Coendou vestitusspecies group: a, soft wool hairs; b, quills; c, bris-tle-quills. Approximately life size (drawn frompelage of AMNH 126171, holotype of C. ichil-lus).

Although some species of Coendou havebeen incorrectly described as covered byquills only (e.g., C. prehensilis, C. bicolor;Ellerman, 1940), a sparse underfur of shortwool hairs is always present among the quillbases of such forms. Other species of Coen-dou are superficially covered by long fur, butthese taxa always possess an underlying de-fensive coat of sharp quills. In fact, all Coen-dou species possess both soft fur and quills;the distinctive feature of the species treatedin this report consists in the additional pres-ence of thin bristle-quills. Because the latterare not known to occur among any other er-ethizontids, the simplest explanation for theirtaxonomic distribution is that they evolveduniquely in the common ancestor of thegroup of species described below.

The oldest name for any Neotropical por-cupine with thin bristle-quills is Coendou ves-

titus Thomas (1899), originally described onthe basis of a single specimen from Colombiapurchased by the British Museum in 1854.Thereafter, the species remained unknownfrom additional material until the early 1920s,when Hermano Niceforo Marıa obtained sev-en specimens in the western foothills of theeastern Andes near Bogota. Unfortunately, noother records of this obviously uncommon (orelusive) porcupine have been reported in thelast 75 years, and it is not known whether ornot the species is still extant.

A second taxon, closely resembling Coen-dou vestitus but diagnosably different in col-oration and craniodental characters, was de-scribed as C. pruinosus by Thomas (1905).Thomas’s type series consisted of five spec-imens obtained between 2500 and 2600 melevation in the Venezuelan Andes near Me-rida, and most subsequently collected mate-rial assignable to pruinosus is likewise frommontane or foothill habitats in Venezuela andColombia. Although some authors (e.g., Ca-brera, 1961; Honacki et al., 1982; Concep-cion and Molinari, 1991; Woods, 1993; So-riano and Ochoa, 1997) have regarded prui-nosus as a subspecies or junior synonym ofvestitus, others (e.g., Handley, 1976; Handleyand Pine, 1992; Linares, 1998; Alberico etal., 1999) have treated these taxa as validspecies. However, no substantive discussionof character data has been provided to sup-port either usage.

Herein we redescribe Coendou vestitusand C. pruinosus based on our examinationof types and other specimens in North Amer-ican and European museums. Clarifying thediagnostic characteristics of vestitus andpruinosus is necessary in order to diagnosetwo new species that significantly extend theecogeographic distribution of vestitus-likeporcupines into the lowland rainforests ofAmazonia. Unanswered questions concern-ing erethizontid distributions in Amazoniaare briefly considered in a concluding dis-cussion. This paper is the second in a seriesof preliminary reports (initiated by Voss andAngermann, 1997) with the object of resolv-ing key taxonomic problems in advance of acomprehensive revision of Neotropical por-cupines currently in preparation.


MATERIALS AND METHODS

SPECIMENS: Specimens that we examinedand measured for this study—and others re-ferred to in the text—are in the AmericanMuseum of Natural History, New York(AMNH); the Natural History Museum, Lon-don (BMNH); the Coleccion de Vertebradosde la Universidad de los Andes, Merida(CVULA); the Museo de la Estacion Biolo-gica de Rancho Grande, Maracay (EBRG);the Departamento de Biologıa de la EscuelaPolitecnica Nacional, Quito (EPN); the FieldMuseum of Natural History, Chicago(FMNH); the Instituto Nacional de Pesquisasda Amazonia, Manaus (INPA); the Museo deBiologıa de la Universidad Central de Ven-ezuela, Caracas (MBUCV); the Museum ofComparative Zoology of Harvard University,Cambridge (MCZ); the Museo de HistoriaNatural La Salle, Caracas (MHNLS); theMuseum National d’Histoire Naturelle, Paris(MNHN); the National Museum of NaturalHistory, Washington, D.C. (USNM); and theMuseum fur Naturkunde der Humboldt-Universitat zu Berlin (ZMB).

MEASUREMENTS: All reported measure-ments are in millimeters (mm) and all re-ported weights are in grams (g). Becausehead-and-body length (HBL) and length oftail (LT) were seldom recorded by collectorsfor the specimens treated in this report, weobtained approximate values (prefixed by‘‘ca.’’) for these dimensions by measuringdried skins to the nearest 5 mm with a flex-ible rule. Length of the hindfoot (HF), re-corded to the nearest millimeter, includes theclaws (c.u.) and was either measured by col-lectors in the field and/or remeasured by uson dried skins (only minimally distorteddried hindfeet containing the intact pedalskeleton were remeasured). Measurements ofthe skull and dentition, taken with dial ordigital calipers and recorded to the nearest0.1 mm, are abbreviated and defined as fol-lows (see fig. 2):

CIL, Condylo-incisive Length: Measuredfrom the articular surface of one occipitalcondyle to the greater curvature of the ipsi-lateral upper incisor.

LD, Length of Diastema: Measured fromthe lesser curvature of an upper incisor at the

alveolar margin to the crown of the ipsilat-eral P4 [dP4 of subadults].

LIF, Length of Incisive Foramen: Insidelength of one incisive foramen (or both, ifthe foramina are recessed in a common fos-sa).

BIF, Breadth of Incisive Foramina: Insidebreadth across both incisive foramina.

MTR, Maxillary Tooth Row: Greatestcrown length from P4 [dP4 of subadults] toM3.

LM, Length of Molars: Greatest crownlength of the upper molar series (M1–M3).

BP4, Breadth of P4: Greatest crownbreadth of the permanent upper premolar.

BM1, Breadth of M1: Greatest crownbreadth of the first upper molar.

APB, Anterior Palatal Breadth: Measuredbetween the crowns of the first upper molars.

PPB, Posterior Palatal Breadth: Measuredbetween the crowns of the third upper mo-lars.

PZB, Posterior Zygomatic Breadth: Great-est breadth across the zygomatic arches be-hind the orbits.

HIF, Height of the Infraorbital Foramen:Measured as the greatest inside diameter,usually at an angle of about 30–408 from themidsagittal plane.

ZL, Zygomatic Length: Measured fromthe posterior margin of the infraorbital fora-men to the posterolateral corner of the zy-gomatic arch.

LN, Length of Nasals: Greatest length ofone nasal bone (the longest if right and leftelements are unequal).

BNA, Breadth of Nasal Aperture: Greatesttransverse dimension of the nasal orifice, al-ways at or near the nasal/premaxillary su-tures.

BB, Breadth of Braincase: Transverse di-mension of the braincase, measured by plac-ing the caliper jaws just above the squamosalzygomatic root on each side.

DI, Depth of Incisor: Distance between thegreater and lesser curvatures of an uppertooth.

BIT, Breadth of the Incisor Tips: Measuredacross the enameled tips of both upper teeth.

AGE CLASSIFICATION: We used maxillarytooth eruption, cranial suture closure, andpelage maturation to define a heuristic age


Fig. 2. Anatomical limits of 18 craniodental measurements defined in the text.


classification as follows (after Voss and An-germann, 1997).

Juveniles—Maxillary toothrow incom-plete (three or fewer teeth erupted); all cra-nial sutures open; pelage often conspicuouslyimmature, including long fur even in speciesthat lack visible fur as adults.

Subadults—Immature maxillary dentition(dP4–M3) completely erupted, or dP4 shedand P4 incompletely erupted; all cranial su-tures still visible; pelage always appears ma-ture.

Adults—Permanent maxillary dentition(P4–M3) fully erupted, with light to moder-ate wear (teeth usually not worn below wid-est part of crown and almost always with atleast some occlusal detail remaining); somecranial sutures usually obliterated.

Old adults—Cheekteeth worn below wid-est part of crown (and therefore not measur-able), with little or no occlusal detail remain-ing on M1 and M2; only nasal sutures (ifany) usually visible.

SPECIES ACCOUNTS

The specimens examined for this reportcan be sorted into four distinct groups on thebasis of external and osteological characters.Whereas two of these groups have availablenames based on extant holotypes, the othersare unnamed. The two named forms are re-described below in chronological order oftheir publication, followed by accounts of thenew species. Specimen records are mappedand documented by geographic coordinatesin the appendix.

Coendou vestitus ThomasFigures 3, 4, 5A, 6A

Coendou vestitus Thomas, 1899: 284 (original de-scription).

Coendou (Sphiggurus) vestitus: Tate, 1935: 307(new name combination).

Coendou (Sphiggurus) vestitus vestitus: Cabrera,1961: 603 (new name combination).

Sphiggurus vestitus: Honacki et al., 1982: 572(new name combination).

TYPE MATERIAL: The holotype only,BMNH 54.6.26.1, consisting of the skin,skull, and mandibles of a subadult animal ofunknown sex. The skin, much faded withage, is understuffed and lacks the left fore-

foot; the left hindfoot is detached but intact.The skull lacks the occiput, a common con-sequence of early to mid-19th century spec-imen preparation methods. The original labelgives the place of origin only as ‘‘N[ouvel]leGrenade’’ (5 Colombia). According toThomas (1899), the specimen was purchasedin 1854.

GEOGRAPHIC DISTRIBUTION: Coendou ves-titus is only known from two definite local-ities, both of which are in the western foot-hills of the eastern Andean cordillera of Co-lombia. The first, where Hermano NiceforoMarıa collected six specimens from 1923 to1925, is San Juan de Rıo Seco, a small vil-lage located about 60 km WNW of Bogotaat ca. 1300 m elevation. The second, wherethe same collector took a single specimen in1925, is Quipile, a village only about 15 kmSSE of San Juan and at approximately thesame altitude.

DESCRIPTION: External—The dorsal fur,soft in texture and dull in appearance, is uni-formly blackish brown and averages about50–60 mm middorsally; the fur is shorter andsparser along the flanks, but it is long enoughto conceal most of the quills except on theface. The quills (ranging in length fromabout 25 to 35 mm middorsally) are bicol-ored, pale yellow or ivory-white basally anddark-brown or blackish distally; usually onlythe distalmost 1⁄ or less of each quill is dark,so the pale quill bases are exposed when thefur is parted or ruffled. None of the quillsanywhere on the body is pale-tipped. Scat-tered abundantly but inconspicuouslythroughout most of the dorsal pelage are long(to 70–80 mm) polished bristle-quills thatemerge from the fur like fine wires; these areyellowish basally, but the emergent tips aredark. No bristle-quills occur over the rump,however, which is clothed only by short,sharp quills and fur. The ventral surface ofthe body is densely covered with soft anduniformly dark-brown fur from chin to anus.Two classes of hairs can be distinguished inthe ventral fur (fine, wavy wool hairs andcoarser, straighter guard hairs), but there areno conspicuously thickened spinous hairsgrouped in triads or other clusters.

The tail is short, apparently averagingabout half the length of the combined head-


Fig. 3. Dorsal view of the skin of Coendou vestitus (BMNH 24.2.21.2). Approximately 30.4.


Fig. 4. Dorsal, ventral, and lateral cranial views of Coendou vestitus (MNHN 1929–632). All viewsapproximately 31.5.


Fig. 5. Ventral cranial views of Coendou vestitus (A, MNHN 1929–632) and C. pruinosus (B, MCZ18738) showing species differences in palatal morphology. In Coendou vestitus, the bony palate betweenthe toothrows is marked by a high median keel that is flanked by deep lateral grooves or gutters (lg);in addition, the mesopterygoid fossa (mpf) extends anteriorly between the second molars, and the roofof the fossa is perforated by large sphenopalatine vacuities (spv). In C. pruinosus, the palate is sometimesweakly keeled but never has deep lateral gutters; also, the mesopterygoid fossa extends anteriorly onlybetween the third molars, and the roof of the fossa is not perforated by distinct sphenopalatine vacuities.

and-body (see measurements in table 1). Theproximal half of the tail is clothed dorsallywith quills and woolly fur like the rump, andthe tip has a naked, calloused dorsal prehen-sile surface, but the rest of the tail is coveredabove and below with blackish bristles; thebristles under the base of the tail are muchstiffer and denser than those on the lateraland dorsal surfaces. The dorsal surface of thehands and feet are densely covered withcoarse brownish hairs.

The long mystacial vibrissae are uniformlyblackish and extend behind the pinnae whenlaid back alongside the head. Supraorbital(superciliary), genal, submental, and postcra-nial vibrissae are also present. The postcra-nial vibrissae occur on the forelimb betweenthe elbow and wrist, on the hindlimb be-tween the knee and ankle, and along the ven-tral surface of the body between the forelimband hindlimb.

Skull—The frontal and nasal sinuses areuninflated, resulting in a flat dorsal profilefrom the nasal tips to the midparietal regionin all the specimens examined. The rostrumis short and tapering in younger animals, butold adults have proportionately longer ros-trums with prominent lateral excavations forthe origin of the infraorbital muscle. The na-sal bones are more-or-less parallel-sided, nei-ther increasing nor decreasing posteriorly inbreadth, with rounded posterior margins thatextend well behind the premaxillae. Viewedfrom above, the zygomatic arches convergeanteriorly from their widest point at the levelof the squamosal roots with only a slight lat-eral deflection at the level of the orbits. Thejugals are slender elements, lacking any con-spicuous postorbital expansion. The dorso-lateral contours of the braincase are stronglysculpted by the origin of the temporalis mus-cle, but the left and right temporal scars are


Fig. 6. Ventrolateral cranial views of Coendou vestitus (A, AMNH 71360) and C. pruinosus (B,MCZ 18738) showing species differences in alisphenoid morphology. In Coendou vestitus, the alisphe-noid is incompletely ossified, resulting in an open sphenopterygoid canal and a buccinator-masticatoryforamen (bmf) that is confluent with the foramen ovale (fo). By contrast, the alisphenoid is fully ossifiedin C. pruinosus, whose sphenopterygoid canal (arrow) is laterally enclosed and whose buccinator-mas-ticatory and oval foramina are separate.

widely separated and do not join middorsallyto form a sagittal crest.

The small incisive foramina are complete-ly contained in the premaxillae (e.g., BMNH24.2.21.2) or shallowly contact the maxillae(e.g., AMNH 71359), but do not deeply pen-etrate between the latter bones; the left andright foramina are incompletely separatedand are recessed in a common fossa in somespecimens, but in others they are completelyseparated and are not recessed together. Theposterior diastema is marked by shallow andwidely separated lateral sulci. In all of thespecimens examined, the palatal bridge (be-tween the toothrows) has a well developedcentral keel and deep lateral gutters (fig. 5A).The mesopterygoid fossa penetrates anteri-orly to or between the second molars, andthe bony roof of the fossa is perforated bywell-formed sphenopalatine vacuities (.1mm in diameter) in all of the specimens athand. The alisphenoid is incompletely ossi-fied, with the result that the sphenopterygoidcanal is open laterally and the buccinator-masticatory foramen is confluent with the fo-ramen ovale (fig. 6A). The auditory bullae

are small (ca. 14–15 mm) rounded capsulesthat are well separated from the base of theparoccipital process on each side. The dorsalroof of the external auditory meatus has anindistinct bony ridge that is less well devel-oped than that seen in some congeneric taxa(e.g., Coendou melanurus; Voss et al., 2001:fig. 70A).

The mandible is distinctive in the absenceof a well-defined coronoid process, which isrepresented only as a rounded convexity atthe base of the ascending ramus in all spec-imens examined.

Dentition—The upper incisors have paleyellow-orange enamel bands and are mod-erately procumbent. The cheekteeth essen-tially resemble those of other erethizontids(except Chaetomys) in occlusal morphology,and the toothrows are subparallel to weaklyconvergent. The permanent fourth upper pre-molar is only slightly larger than the first mo-lar in some specimens (e.g., AMNH 70529),but P4 is conspicuously larger than M1 inothers (e.g., USNM 240035).

COMPARISONS: See the accounts for Coen-


TABLE 1Measurements (mm) of All Known Specimens

of Coendou vestitus

dou pruinosus and the new species describedbelow.

NATURAL HISTORY AND CONSERVATION

STATUS: Nothing has apparently been record-ed about the habitats or behavior of this spe-cies, but the natural vegetation at the two lo-calities where specimens are definitelyknown to have been collected was probablylower montane moist forest (bosque humedosubtropical; Espinal and Montenegro, 1963).Unfortunately, habitat destruction is virtuallycomplete in this part of Colombia (IAVH,1998), and it is possible that the species hasbeen extirpated. Apparently, no specimens orsightings have been recorded since 1925.

SPECIMENS EXAMINED: Colombia—Cundi-namarca, Quipile (AMNH 70529), San Juan

de Rıo Seco (AMNH 70596, 71359; BMNH24.2.21.2; MNHN 1929.631, 1929.632;USNM 240035); ‘‘Colombia’’ (no other lo-cality data, AMNH 71360); ‘‘N[ouvel]le.Grenade’’ (BMNH 54.6.26.1 [holotype]).

Coendou pruinosus ThomasFigures 5B, 6B, 7, 8

Coendou pruinosus Thomas, 1905: 310.Coendou (Sphiggurus) pruinosus: Tate, 1935: 307

(new name combination).Coendou (Sphiggurus) vestitus pruinosus: Ca-

brera, 1961: 602 (new name combination).Sphiggurus vestitus: Honacki et al., 1982: 572

(part; pruinosus treated as a junior synonym).

TYPE MATERIAL: The holotype, BMNH5.7.5.9, consisting of the skin, skull, andmandibles of an adult male collected by S.Briceno Gabaldon e hijos at 2500 m eleva-tion in the Montanas de la Pedregosa nearMerida, Venezuela, on 14 January 1905. Inaddition, Thomas (1905) examined four par-atypes, of which three retain original labelswith Briceno’s commercial imprint: two ofthese are from ‘‘Montana del Valle 2600 m’’,and were collected on 16 November 1903(BMNH 5.7.5.8, 5.7.5.11); the other is from‘‘Montanas del Tabay 2600 m’’, and was col-lected on 28 January 1904 (BMNH 5.7.5.12).The fourth paratype (BMNH 5.7.5.10) has amuseum label in Thomas’s hand with the lo-cality given as ‘‘Merida, Venezuela’’, and thecollection date as 14 June 1904. Details ofskin preparation suggest that the entire typeseries came from Briceno e hijos, well-known commercial collectors and taxider-mists active at Merida around the turn of thelast century.

GEOGRAPHIC DISTRIBUTION: Referred spec-imens are from the foothills and crest of theCordillera Oriental of Colombia, from theadjacent but disjunct Serranıa de la Macar-ena, and from western and northern Vene-zuela. The known range of this species inVenezuela includes the foothills of the Ser-ranıa de Perija, the lowlands of the westernMaracaibo Basin, the Cordillera de Merida,and the Cordillera de la Costa. Recorded el-evations on specimen tags range from 54 to2600 m above sea level.

DESCRIPTION: External—The pelage ofCoendou pruinosus is morphologically sim-


Fig. 7. Dorsal view of the skin of Coendou pruinosus (BMNH 10.12.3.5). Approximately 30.4.


Fig. 8. Dorsal, ventral, and lateral cranial views of Coendou pruinosus (MCZ 18738). All viewsapproximately 31.5.


ilar to that of C. vestitus but differs in col-oration. Although the dorsal fur is predomi-nantly blackish (Thomas, 1905) or darkbrown, the hair tips are pale (grayish or sil-very), producing a frosted effect that is morepronounced in some individuals (e.g.,BMNH 10.12.3.5) than in others (e.g.,BMNH 26.11.4.11). The fur is long (50–70mm midorsally) and dense enough to concealmost of the underlying quills except on thehead. All of the body quills (30–40 mm longmiddorsally) are bicolored (yellowish or ivo-ry-white basally with the extreme tips darkbrown), but some of the quills on the crownof the head and on the cheeks are tricolored(with white bases and tips separated by abroad dark middle band). Scattered through-out the dorsal pelage—except over therump—are many long (to 80–100 mm) wire-like bristle-quills, the pale (usually ivory-white or yellowish) tips of which are con-spicuous against the dark background of thefur; most of the bristle-quills are actually tri-colored (with pale bases and tips separatedby a single dark middle band), but some areentirely pale. The ventral body pelage con-sists of soft frosted-brown fur from chin toanus. Longer, straighter, coarser guard hairscan be distinguished from finer, shorter, wavywool hairs in the ventral pelage, but there areno conspicuously thickened spinous hairsrooted in obvious clusters.

The tail is short, averaging slightly morethan half the combined length of the head-and-body in specimens from montane habi-tats (e.g., the Alturas de Pamplona and Me-rida series in table 2), but specimens fromthe Maracaibo lowlands (e.g., MHNLS 7692)and Caracas (BMNH 26.11.4.11) appear tohave relatively and absolutely longer tails.Whereas the dorsal surface of the proximalhalf of the tail is covered by a mixture ofquills and soft fur like that on the rump, pale-tipped bristles occur along the sides of thetail and converge posteriorly onto the caudaldorsum forming a conspicuous pale (yellow-ish or whitish) chevron. The prehensile tailtip is naked and calloused dorsally, but therest of the tail is covered by blackish bristles;the bristles under the base of the tail aremuch stiffer and denser than those on the lat-eral and dorsal surfaces. The dorsal surface

of the hands and feet are covered with coarsefrosted-brownish hairs.

The cranial and postcranial vibrissae havethe same morphology, coloration, and ana-tomical distribution as previously describedfor Coendou vestitus.

Skull—The frontal and nasal sinuses areuninflated in most specimens, resulting in adorsal profile that is nearly flat from the nasaltips to the midparietal region; a single spec-imen (MHNLS 7692), however, has slightlyinflated dorsal sinuses that produce a smallbut distinct swelling over the maxillary zy-gomatic roots. The rostrum is short and ta-pering in subadults and young adults, butlengthens in older animals, some of whichdevelop shallow lateral excavations for theinfraorbital muscle. The nasal bones are ap-proximately parallel-sided in some speci-mens and taper slightly posteriorly in others,but always have rounded posterior marginsthat extend well behind the premaxillae.Viewed from above, the zygomatic archesconverge anteriorly from the squamosal rootswithout a well-developed secondary widen-ing at the orbits; some specimens (e.g.,FMNH 140260), however, have more round-ed (less angular) zygomatic outlines thanothers (e.g., MCZ 18738; fig. 8). The jugalsare moderately deep (dorsoventrally expand-ed), with some specimens (e.g., BMNH10.12.3.5, USNM 172985) exhibiting a dis-tinct postorbital swelling. The bony crestsand scars associated with the origin of thetemporalis muscle are strongly developed insome specimens, but a sagittal crest is notdeveloped in any of the material examined.

The incisive foramina are moderately longand usually bordered posteriorly by the max-illae, but they do not penetrate deeply be-tween those bones; in most specimens, theleft and right foramina are completely sepa-rated by a bony septum and are not recessedin a common fossa. (A single specimenamong those we examined, USNM 496172,is an exception: the foramina are containedwithin the premaxillae, and are incompletelyseparated and recessed in a common fossa.)The posterior diastema is marked by shallowand widely separated lateral sulci. The pala-tal bridge between the toothrows is smooth(e.g., AMNH 21350) or weakly keeled(USNM 172985), but lacks deep lateral gut-


TABLE 2Measurements (mm) of Adult Specimens of Coendou pruinosus

ters (fig. 5B). The mesopterygoid fossa isshallow in most specimens (penetrating onlybetween the third molars) and the bony roofof the fossa is perforated by tiny nutrient fo-ramina or by small irregular holes that do nothave the aspect of distinct sphenopalatine va-cuities. The alisphenoid is completely ossi-fied in all specimens examined, such that thesphenopterygoid canal is enclosed laterallyand the buccinator-masticatory foramen isnot confluent with the foramen ovale (fig.6B). The auditory bullae are small (ca. 14–15 mm) rounded capsules that are well sep-arated from the paroccipital processes inmost specimens, but three skulls (BMNH26.11.4.11; FMNH 140261; MHNLS 7692)

have unusually large (16–18 mm) bullae.The dorsal roof of the external auditory me-atus has an indistinct bony ridge in mostadult specimens, but two specimens(MHNLS 7692, USNM 496172) have dis-tinct auditory ridges.

The mandible bears a small but distinctcoronoid process in all specimens examined.

Dentition—The dentition of Coendoupruinosus is similar to that of C. vestitus, ap-parently without any distinctive features.

COMPARISONS: Coendou pruinosus and C.vestitus are similar in many points of com-parison, yet each is morphologically distinc-tive. Externally, pruinosus differs from ves-titus by its frosted (versus unicolored-dark)


dorsal and ventral fur, tricolored (versus bi-colored) head quills, pale-tipped (versusdark-tipped) bristle-quills, and chevron ofpale-tipped caudal bristles (versus caudalchevron absent). Cranially, the two speciesare clearly distinguishable by palatal mor-phology (the bony palate between the tooth-rows is more distinctly ridged and groovedin vestitus than in pruinosus), mesopterygoidperforation (distinct sphenopalatine vacuitiesare present in vestitus versus absent in prui-nosus), alisphenoid ossification (the sphen-opterygoid canal is laterally open in vestitusversus closed in pruinosus), and mandibularmorphology (a distinct coronoid process isabsent in vestitus but present in pruinosus).In addition, other modal differences supportthe hypothesis that these are genetically dif-ferentiated taxa: (1) rostral morphology (thesides of the rostrum appear to be more deep-ly excavated for the origin of the infraorbitalmuscle in vestitus than in like-aged pruino-sus); (2) depth of the jugal (more dorsoven-trally expanded below the orbit in pruinosusthan in vestitus); (3) morphology of the in-cisive foramina (short, incompletely separat-ed, and contained entirely by the premaxillaein most vestitus versus longer, completelyseparated, and bordered by the maxillae inmost pruinosus); and (4) depth of the me-sopterygoid fossa (penetrating to or betweenthe second molars in vestitus versus shallow-er in most pruinosus). Although only smallsamples are currently available for taxonom-ic comparisons, these differences permit un-ambiguous identifications and seem morethan sufficient to recognize C. vestitus andC. pruinosus as valid species.

VARIATION: Consistent with its larger ec-ogeographic range, Coendou pruinosus ismore morphologically variable than C. ves-titus, but all of the referred material shares acommon aspect and we provisionally regardit as representing conspecific populations.Nevertheless, a few specimens from geo-graphically peripheral localities are notablyatypical. In particular, AMNH 136312 (fromthe foothills of the Cordillera Oriental in De-partamento Meta, Colombia) has unusuallysparse fur, a deep mesopterygoid fossa (ex-tending between the second molars), andsmall toothrows (MTR, 13.3 mm); its bullaeare relatively longer (approaching the base of

the paroccipital process on each side) andmore laterally compressed than those seen intopotypical material. Likewise, MHNLS7692 (from the foothills of the Serranıa dePerija) has uniquely swollen frontal sinuses,dorsal bristles tipped with pale orange (in-stead of white or pale yellow), and very largebullae. Finally, the single specimen exam-ined from the Cordillera de la Costa (BMNH26.11.4.11, a subadult) has unusually shortincisive foramina, small teeth, and large bul-lae. The possibility that such outliers repre-sent distinct taxa merits testing when morematerial becomes available from these areas.

REMARKS: A specimen collected ca. 75 kmSW of the type locality was karyotyped byConcepcion and Molinari (1991), who report-ed a diploid number of 42 and a fundamentalnumber of 76. Their morphological descrip-tion of the karyotyped animal is consistentwith that given above for Coendou pruinosus,with only minor differences. The reported ex-ternal dimensions and weight of this specimen(CVULA I-3030, an adult male) are the onlysuch data obtained to date from positivelyidentified fresh material near the type locality:300 mm (HBL) 3 220 mm (Tail) 3 55 mm(HF, c.u.) 3 22 (Ear) 5 1010 g.

Specimens currently labeled as ’’Sphig-gurus vestitus’’ in Venezuelan museums arepresumably referable to Coendou pruinosusas herein recognized, but the following ma-terial should be examined for diagnostic attri-butes to confirm this identification: EBRG126 (Aragua, Rancho Grande), EBRG 4350(Merida, 2 km E Jajı), EBRG 10281 (Miran-da, Parque Nacional El Avila), EBRG 20243(Aragua, Colonia Tovar), MBUCV 1472(Distrito Federal, Carayaca), MBUCV 4155(Miranda, Estacion Experimental de Rıo Ne-gro), MHNLS 58 (Miranda, Turgua),MHNLS 3879 (Distrito Federal, Caracas),MHNLS 8480 (Zulia, Hacienda La Ceiba).

NATURAL HISTORY: All known specimensof Coendou pruinosus have been collected inregions where the original (non-anthropogen-ic) vegetation is (or was) humid forest. Atelevations below 1000 m, the species inhab-its lowland rainforest, but at higher eleva-tions the typical habitat is probably cloudforest (Lower Montane or Upper MontaneRainforest; Grubb, 1977). To the best of ourknowledge, no specimens have been taken in


unforested (e.g., savanna or paramo) land-scapes.

Published natural history information canbe associated with only three positively iden-tifiable records of Coendou pruinosus. Han-dley (1976: 55) reported that one individual(USNM 496172, an adult male) was ‘‘foundin a tree in an upland area in mature ever-green forest’’ near El Rosario (54 m eleva-tion; Estado Zulia, Venezuela), where the lo-cal vegetation was further characterized as‘‘[m]ature evergreen forest 18–30 m high,with many palms and vines’’ (op. cit.: 69).Concepcion and Molinari (1991: 238) sub-sequently reported another specimen (CVU-LA 1–3030, also an adult male) that was cap-tured ‘‘in an area dominated by cloud forest’’at an elevation of 1500 m near the town ofZea (Estado Merida, Venezuela).

On the night of 6 July 1986, Voss shot an-other adult male specimen (MHNLS 7692)that was perched about 20 m above theground on the trunk of a large buttressed treefestooned with lianas and epiphytes near Mi-sion Tukuko (ca. 300 m elevation; Estado Zu-lia, Venezuela). The tree was part of the orig-inal rainforest canopy at this site, but most ofthe surrounding undergrowth had beencleared for cacao cultivation (for a detaileddescription of local habitats, see Voss, 1991:68–70). The freshly dissected stomach wasfull of brownish paste, possibly consisting ofchewed bark; no insect parts, seeds, or otheridentifiable food fragments were observed.

SPECIMENS EXAMINED: Colombia—Meta,Villavicencio (AMNH 136312), Pico Rengifo(FMNH 87896); Norte de Santander, Alturasde Pamplona (FMNH 140260, 140261). Ven-ezuela—Distrito Federal, Caracas (BMNH26.11.4.11); Merida, Merida (AMNH 21350,BMNH 5.7.5.10, 10.12.3.5), Montanas de laPedregosa (BMNH 5.7.5.9), ‘‘Montana de laSierra’’ (MCZ 18738, ZMB 33377), Montan-as de Tabay (BMNH 5.7.5.12), ‘‘Montana delValle’’ (BMNH 5.7.5.8, 5.7.5.11); Zulia, ElRosario (USNM 496172), Mision Tukuko(MHNLS 7692). ’’South America’’ (USNM172985).

Coendou ichillus, new speciesFigures 9–12

TYPE MATERIAL: The holotype, AMNH126171, consists of the skin, skull, and man-

dibles of a young adult of unknown sex col-lected by the von Baumann-Roosevelt Ex-pedition in July 1936 on the Rıo Pastaza, Ec-uador.3 The stuffed skin (fig. 9) appears to beslightly faded, but is in otherwise perfectcondition with no missing elements. Theskull (fig. 10) lacks part of both nasal bonestogether with most of the right premaxilla,but is otherwise complete. Two paratypes arefrom unknown localities on the Rıo Con-ambo (EPN 806, an adult female) and theRıo Yana Rumi (FMNH 43289, an adultmale), both in Provincia Pastaza, Ecuador.

GEOGRAPHIC DISTRIBUTION: All unambigu-ous records of Coendou ichillus are from theAmazonian lowlands of eastern Ecuador, buta referred juvenile specimen purchased nearIquitos (see Variation, below) suggests thatthe species is more widely distributed.

ETYMOLOGY: From ichilla, meaning‘‘small’’ (Orr, 1978) in the dialect of the low-land Quichua, within whose tribal territorythe new species occurs.

DIAGNOSIS: A member of the Coendou ves-titus group distinguished from other speciesby its long tail, lack of visible fur in the adultpelage, more extensively black-tipped quills,tricolored (pale-tipped) bristle-quills, spinyventral pelage, and a unique combination ofcranial traits.

DESCRIPTION: External—No soft fur is ap-parent in the dorsal pelage except by closeexamination because the blackish wool hairsare sparse and hidden among the quills andbristles; a few dorsal wool hairs pluckedfrom two specimens were about 40 mm inlength. All of the quills (30–40 mm longmiddorsally) are bicolored (yellowish basallywith the terminal ⅓ to ½ dark-brown or

3 The original specimen label gives the locality as‘‘Rio Pastaza’’ only, but the entire von Baumann-Roo-sevelt collection (including 10 other mammals and 25birds variously labeled ‘‘Rio Tigre’’, ‘‘Rio Bobonaza’’,Rio Blanco’’, ‘‘Rio Pastaza’’, and ‘‘Rio Napo’’) wasknown to have been obtained in Ecuador, so the lower(Peruvian) course of this river can be ruled out. Unfor-tunately, the exact route traveled by the von Baumann-Roosevelt Expedition is unknown, and no archival re-cord of the purpose or staffing of this mysterious outfitappears to exist. Filed correspondence with its director,Cyril von Baumann, consists only of an AMNH acces-sions receipt dated 23 January 1937, together with a curtacknowledgment from von Baumann dated 1 February.


Fig. 9. Dorsal view of the skin of Coendou ichillus (AMNH 126171, holotype). Approximately30.4.


Fig. 10. Dorsal, ventral, and lateral cranial views of Coendou ichillus (AMNH 126171, holotype).All views approximately 31.5.


Fig. 11. Dorsal, ventral, and lateral cranial views of Coendou ichillus (FMNH 43289, paratype). Allviews approximately 31.5.


Fig. 12. An adult female Coendou ichillus eating bananas suspended from the rafters of a buildingat La Selva Jungle Lodge, Provincia Sucumbıos, Ecuador, on 9 July 1996 (photo courtesy of P. J.DeVries).

blackish), except on the head, where somequills are tricolored (with ivory-white basesand tips separated by a dark middle band).Densely scattered among the quills are manylong (to ca. 80 mm) thin bristle-quills withyellowish bases and tips separated by a broaddark-brown or blackish middle band; the paletips of the bristle-quills produce a character-

istically streaked effect over the whole dor-sum with the exception of the rump (whichis covered only with short bicolored quillsand a few wool hairs). The ventral body pel-age consists of very coarse bicolored or tri-colored hairs (ca. 0.30–0.40 mm in diameterand 15 mm long) that are usually rooted intriplets but occasionally in groups of two or


TABLE 3Measurements (mm) of All Known

Adult Specimens of Coendou ichillus

four; a few short wool hairs are almost in-visibly scattered among this harsh covering.

The tail is moderately long, averagingrather more than ¾ of head-and-body lengthon the two skins from which approximatemeasurements could be obtained (table 3); inlife, the extended tail of one individual (onlypartly visible in fig. 12) seemed to be almostas long as the head and body, but no accuratemeasurement could be taken from this un-restrained animal (see Natural History, be-low). The dorsal part of the base of the tailis densely covered with short bicolored quillslike those of the rump, but tricolored bristlesextend along the lateral caudal surfaces andconverge dorsally to form an indistinct whit-ish or yellowish chevron near the middle ofthe tail. The prehensile tail-tip is callousedand naked dorsally, but the rest of the tail isdensely covered by blackish bristles; the cau-dal bristles are conspicuously stiffer and

denser under the base of the tail than else-where. The hands and feet are covered dor-sally with coarse blackish hairs.

The cranial and postcranial vibrissae ofthis species have the same morphology, col-oration, and distribution as previously de-scribed for Coendou vestitus.

Skull—The frontal and nasal sinuses areuninflated, resulting in a flat dorsal profilefrom the nasal tips to the midparietal region.In dorsal view, the rostrum is short and mod-erately broad, lacking obvious lateral emar-ginations for the origin of the infraorbitalmuscle. The nasal bones are damaged in twoof the three specimens at hand, but appear totaper gently from front to back; the posteriornasal margins are rounded and extend behindthe premaxillae (much more so in FMNH43289 than in AMNH 126171). Viewed fromabove, the zygomatic arches are rounded(AMNH 126171) or biconcave (EPN 807,FMNH 43289) because they are anteriorlydeflected to accommodate relatively largerorbits than those of either Coendou vestitusor typical C. pruinosus. The jugals have ei-ther a small postorbital expansion (AMNH126171, EPN 807) or taper more-or-lessevenly from front to back (FMNH 43289).The dorsolateral contours of the braincaseshow only slight to moderate sculpting forthe origin of the temporalis muscle; the rightand left temporalis scars are widely separatedin all three specimens examined.

The small incisive foramina are incom-pletely separated and recessed in a commonfossa that is contained entirely by the pre-maxillae (EPN 807) or bordered narrowly bythe maxillae behind. The posterior diastemais marked by shallow and widely separatedlateral sulci. The palatal bridge between thetoothrows does not have a distinct mediankeel or deep lateral gutters in any of the spec-imens examined. The mesopterygoid fossaextends anteriorly just beyond the point ofcontact between M2 and M3, and the bonyroof of the fossa (in AMNH 126171 andEPN 807) is not perforated by distinct sphe-nopalatine vacuities. The alisphenoid is com-pletely ossified such that the sphenoptery-goid canal is enclosed laterally and the buc-cinator-masticatory and oval foramina areseparate. The auditory bullae are large (ca.


16–17 mm) inflated capsules that contact thebase of the paroccipital process on each side.The dorsal roof of the external auditory me-atus lacks a distinct bony keel in both spec-imens at hand (AMNH 126171, FMNH43289).

The mandible is provided with a small butwell-developed coronoid process.

Dentition—The dentition is qualitativelysimilar to that of Coendou vestitus and C.pruinosus, but measurements suggest that theincisors are relatively broader than in thosespecies, and they appear to be more deeplypigmented as well.

COMPARISONS: No other species of theCoendou vestitus group lacks a visible coatof long fur, and this trait together with itsrelatively long tail and spiny ventral pelagemake C. ichillus externally unmistakable.Additionally, the pigmented tips of the de-fensive quills are much longer in ichillusthan in either vestitus or pruinosus; as a re-sult, when the long fur of the latter two spe-cies is shed or parted, the exposed quills andbristles produce the effect of a pale (yellow-ish or whitish) animal streaked and stippledwith dark brown, whereas the permanentlyexposed quills and bristles of ichillus pro-duce the effect of a blackish animal streakedwith white or yellow.

Cranially, Coendou ichillus is distin-guished from C. vestitus and from typical ex-amples of C. pruinosus by its proportionatelysmaller and less laterally excavated rostrum,more rounded (or biconcave) zygomaticarches, and proportionately larger bullae.From vestitus, the new species additionallydiffers by its less strongly muscle-scarredbraincase, less strongly keeled and groovedpalatal bridge, unperforated mesopterygoidfossa, and distinct mandibular coronoid pro-cess. From pruinosus, the new species addi-tionally differs by its smaller incisive foram-ina and more anterior mesopterygoid pene-tration.

External and cranial comparisons with thesecond new species are provided in the fol-lowing account.

VARIATION: Cranial differences among thethree available adult specimens of Coendouichillus do not exceed the range of osteolog-ical variation routinely observed within localpopulations of other Neotropical porcupines.

However, the skin of the holotype appears tobe faded by comparison with the more viv-idly colored paratypes. The latter closely re-semble living examples (fig. 12), whose darkpelage markings are distinctly blackish. Bycontrast, the corresponding pigmentation onAMNH 126171 is brownish (near Smithe’s[1975] Sepia or Raw Umber), a chromaticdifference that might have resulted from in-adequate specimen storage (e.g., exposure tolight or bleaching agents) at some point inthe past.

An immature specimen (FMNH 112565)purchased by Pekka Soini in 1971 near Iqui-tos is mounted for display with the skull in-side and has painted seeds for eyes; obvi-ously intended for the tourist trade, it is un-accompanied by records of sex, measure-ments, or habitat. The dorsal body pelageconsists of bicolored quills, tricolored bristle-quills, and long reddish fur; the ventral pel-age consists mostly of reddish fur, but a fewclusters of soft spines are emerging along themidline. Measurements made with a flexiblerule suggest that the tail in life was more thanthree-quarters of the combined length ofhead and body; the estimated length of thedried hind foot is ca. 48 mm.

Long reddish juvenile fur appears to be ataxonomically widespread trait among Neo-tropical porcupines, occurring even in thosespecies that lack visible fur as adults (Rob-erts et al., 1985; Handley and Pine, 1992).We identify FMNH 112565 as Coendou ich-illus because its long tail, boldly black-tippedquills, tricolored bristle-quills, and emergingventral spines suggest a similar external phe-notype at maturity. Although this identifica-tion is biogeographically plausible (othereastern Ecuadorean taxa have ranges that ex-tend southeastward into Peru, e.g., Saguinusnigricollis graellsi; Hershkovitz, 1977: fig.X.21), the exact provenance of FMNH112565 is uncertain because tourist itemssold near Iquitos might have originated farfrom the city.

NATURAL HISTORY: Although no usefulecological association can be inferred fromthe type locality, ‘‘Rıo Pastaza’’, a river thattraverses a considerable range of habitats inits descent from the Andes to the Amazon,both paratypes are from densely rainforestedcatchments below 500 m elevation. The orig-


inal skin tag attached to FMNH 43289 in-cludes the notation ‘‘Col[ectado]: con cer-batana y beneno’’ (collected with blowgunand poisoned dart), suggesting that the ani-mal was taken from a tree by a native hunter,probably belonging to one of the lowlandQuichua groups described by Whitten(1976). No other natural history informationis available from any of the specimens athand.

Fortunately, field observations of Coendouichillus are available from at least one defi-nite locality: La Selva Jungle Lodge, an eco-tourist facility and biological station at ca.300 m elevation on the left (north) bank ofthe Rıo Napo in Provincia Sucumbıos, Ec-uador. Situated between the oxbow lakes ofGarza Cocha and Mandi Cocha, the lodgeand its surrounding trail system occupy thetypical ridge-and-swale landscape created bymeandering white-water rivers, with tall for-est on the high ground and swamps in thevalley bottoms. Balslev et al. (1987) provid-ed a detailed description of the local vege-tation based on their study of flooded andunflooded forest at nearby Anangu (on theopposite bank of the Napo). Several years ofweather records from La Selva suggest thatthis region receives between 3500 and 4000mm of annual rainfall (DeVries et al., 1999).

An adult female porcupine, unambiguous-ly identifiable as Coendou ichillus, was brief-ly observed at La Selva (by J. E. Cadle, P. J.DeVries, L. H. Emmons, and R. S. Voss) onthe night of 9 July 1996 as it ate ripe bananassuspended by a cord from the rafters of abuilding surrounded by gardens and second-ary vegetation (fig. 12). Several days later,another adult C. ichillus was observed atnight (by J. E. Cadle and L. H. Emmons) inadjacent primary floodplain forest, where itwas perched on a liana about 8 m above theground. At least two individuals were said bylodge employees to inhabit holes in hollowpalm trunks that were used as upright sup-ports for the kitchen and other buildings atLa Selva.

SPECIMENS EXAMINED: Ecuador—Pastaza,Rıo Conambo (EPN 807), Rıo Yana Rumi(FMNH 43289); Rıo Pastaza (AMNH126171 [holotype]). Peru—Loreto, Iquitosregion (FMNH 112565).

Coendou roosmalenorum, new speciesFigures 13, 14

TYPE MATERIAL: The holotype, INPA 2586(original number CCM 58), consists of thewell-preserved skin, skull, and mandibles ofan adult female collected by M.G.M. and T.van Roosmalen on 23 November 1996 at thecaboclo settlement of Novo Jerusalem nearthe left bank of the middle Rio Madeira inthe Brazilian state of Amazonas. One para-type (INPA 2587) is from the nearby villageof Santa Maria on the same side of the Ma-deira, but the other (INPA 677) is from 49km E Porto Velho on the opposite bank inthe Brazilian state of Rondonia.

GEOGRAPHIC DISTRIBUTION: Known fromboth banks of the middle Rio Madeira inBrazil between 5 and 98 S latitude.

ETYMOLOGY: For Marc van Roosmalen andhis son Tomas, whose collections from themiddle Madeira included this distinctive por-cupine together with other previously undes-cribed mammalian taxa (Roosmalen et al.,1998, 2000).

DIAGNOSIS: A member of the Coendou ves-titus group distinguished from other memberspecies by its small size, long tail, long adultfur, minutely black-tipped quills, nonspinousventral fur, and unique combination of cran-iodental traits.

DESCRIPTION: External—The dorsal fur ofsoft wool hairs is dull in appearance and ap-parently variable in coloration, pale grayish-brown in one specimen (INPA 2586) butdark brown in the other two (INPA 677,2587); in the specimens with brownish dorsalfur, the hair tips are pale (grayish or silvery)but these do not produce a distinctly frostedmass effect. The dorsal fur is long (50–70mm) and dense enough over the back andrump to partly (INPA 2586, 2587) or com-pletely (INPA 677) conceal the underlyingquills. All of the quills (ranging in lengthfrom about 25 to 35 mm middorsally) arebicolored (yellowish basally with the ex-treme tips dark brown) over the entire dor-sum, including the head. Scattered abun-dantly throughout most of the dorsal pelage(except the rump) are long (40–70 mm) wire-like bristle-quills; almost all of these are tri-colored—with pale (yellowish or ivory-white) bases and tips separated by a single


Fig. 13. Holotype of Coendou roosmalenorum (INPA 2586) in life. Photos courtesy of M.G.M. vanRoosmalen.


Fig. 14. Dorsal, ventral and cranial views of Coendou roosmalenorum (INPA 2586 [original numberCCM 58], holotype). All views approximately 31.5.


TABLE 4Measurements (mm) of All Known Specimens

of Coendou roosmalenorum

dark-brown middle band of variable width—but a few bristles are bicolored (with palebases and dark tips). The ventral surface ofthe body is densely covered with a mixtureof fine, soft, brownish wool and coarser,banded (tricolored or bicolored) hairs fromchin to anus; the banded hairs (0.20–0.25mm in diameter and 15–20 mm long) arerooted in clusters, usually triplets but occa-sionally groups of two or four.

The long tail is almost 90% of head-and-body length in both of the specimens fromwhich measurement data are available (table4). The basal third of the tail is covered dor-sally with bicolored quills and woolly fur,and the prehensile tail-tip is bare and cal-loused dorsally, but the rest of the tail (aboveand below) is uniformly covered with black-ish bristles; the caudal bristles under the baseof the tail are stiffer and denser than thoseoccurring elsewhere. The dorsal surfaces of

the hands and feet are densely covered withcoarse dark-brown or blackish hairs.

The cranial and postcranial vibrissae haveessentially the same morphology, coloration,and anatomical distribution as previously de-scribed for other vestitus-group porcupines.

Skull—The frontal and nasal sinuses areslightly inflated in the younger left-bankspecimens (INPA 2586, 2587), producing aslightly swollen dorsal profile above the or-bits; the sinuses of the older right-bank spec-imen (INPA 677) are uninflated, however, re-sulting in a flat dorsal profile from the nasaltips to the midparietal region. The rostrum isshort and tapering in our younger examplesbut moderately broad in INPA 677; however,there are no well-developed lateral emargi-nations for the origin of the infraorbital mus-cle in any specimen. The nasal bones tapergently from front to back, with rounded pos-terior margins that extend well behind thepremaxillae. Viewed from above, the zygo-matic arches converge anteriorly from thesquamosal roots without a conspicuous wid-ening at the level of the orbits. The jugalsare moderately deep with distinct postorbitalprocesses in all examined material. The dor-solateral contours of the braincase are onlyslightly (INPA 2587) to moderately (INPA2586 and 677) sculpted by the origin of thetemporalis muscles. The left and right tem-poralis scars are widely separated and are notjoined middorsally to form a sagittal crest inany specimen.

The incisive foramina are short and almostcompletely contained in the premaxillae(INPA 2586), or longer and penetrating be-tween the maxillae posteriorly (INPA 2587,677); the left and right foramina are com-pletely separated by a bony septum in INPA2586, but the septum is incomplete and bothforamina are recessed in a common fossa inthe other two specimens. The posterior dia-stema is marked by shallow and widely sep-arated lateral sulci. Although the palatalbridge between the toothrows has a weaklydeveloped central keel and shallow lateralgutters anterior to M1, the posterior palate ismore-or-less smooth. The mesopterygoidfossa extends slightly anterior to the point ofcontact between M2 and M3 in the holotype,but not in either paratype. Small sphenopal-atine vacuities (ca. 1 mm in diameter) per-


forate the bony roof of the mesopterygoidfossa in INPA 677, but not in INPA 2586(the mesopterygoid region of INPA 2587 istoo damaged to score confidently for thistrait). The alisphenoid is completely ossified,such that the sphenopterygoid canal is en-closed laterally and the buccinator-mastica-tory foramen and the foramen ovale are sep-arate. The auditory bullae are large (ca. 17mm) inflated capsules that contact the baseof the paroccipital process on each side. Thedorsal roof of the external auditory meatushas a weakly developed bony ridge in twospecimens (INPA 677 and 2587) but not inINPA 2586.

The coronoid process of the mandible iswell-developed in all examined specimens.

Dentition—The dentition is qualitativelysimilar to that of the remaining species of thevestitus group, but the toothrows of INPA2586 and INPA 677 are remarkably small(table 4). The upper incisors are strongly pro-cumbent and deeply pigmented (orange).

COMPARISONS: Coendou roosmalenorumcan be distinguished unambiguously fromother species of the vestitus group by aunique combination of characters, the mostsalient of which are tabulated for easy ref-erence (table 5). Externally, roosmalenorumresembles vestitus and pruinosus by its long,dense dorsal fur and nonspinous ventral pel-age, but it differs from both of those speciesin details of fur coloration (grayish or brown-ish versus blackish), in body size (muchsmaller, as indicated by hindfoot measure-ments), and by its conspicuously longer tail.Other external characters provide differentpatterns of taxonomic contrasts. Like vesti-tus, roosmalenorum has only bicolored cra-nial quills, whereas at least some cranialquills are tricolored in all examined speci-mens of pruinosus. Like pruinosus, however,roosmalenorum has tricolored bristle-quills,whereas the bristle-quills of vestitus are bi-colored. In craniodental traits, roosmaleno-rum differs from vestitus in palatal sculptur-ing, alisphenoid ossification, bullar size,mandibular morphology, and in many addi-tional (but subtler) details. From pruinosus,roosmalenorum differs craniodentally by itsrelatively much larger bullae, but other os-teological contrasts between these taxa areharder to assess due to the wide range of var-

iation among referred specimens. For exam-ple, the contrast in rostral morphology be-tween some exemplar crania of pruinosus(e.g., MCZ 18738; fig. 8) and roosmaleno-rum (e.g., INPA 2586; fig. 14) is striking, butother specimens that we could have chosento represent each species (e.g., FMNH140260 [pruinosus] and INPA 677 [roos-malenorum]) are much more similar in thischaracter. Average (or modal) character dif-ferences between roosmalenorum and prui-nosus, however, include dental size (roos-malenorum has smaller molars), mesoptery-goid penetration between the toothrows(deeper in roosmalenorum), mesopterygoidfenestration (sphenopalatine vacuities arebetter developed in roosmalenorum), and na-sal/frontal sinus inflation (perhaps more fre-quent in roosmalenorum).

Although Coendou roosmalenorum and C.ichillus are both long-tailed species, they dif-fer markedly in external appearance, mostnotably in dorsal fur length (long versusshort, respectively), density (dense versussparse), and color (gray or brown versusblackish); in addition, whereas the quills ofroosmalenorum are only dark at the extremetips, those of ichillus are much more exten-sively pigmented. Other trenchant externaldifferences include size (roosmalenorum issmaller, as evidenced by hindfoot measure-ments), and ventral pelage composition(coarse banded hairs in ichillus, woollier inroosmalenorum). The sum of such contrastsis sufficiently striking that it is unlikely thatthese species could be confused in the fieldby competent observers. By contrast, cranialdifferences between roosmalenorum and ich-illus are harder to identify given the smallsamples at hand. Although it is possible thatthe morphology of the nasal/frontal sinuses(perhaps more frequently inflated in roos-malenorum), zygomatic arches (perhapsmore rounded in ichillus), and the mesopter-ygoid roof (perhaps more frequently fenes-trated in roosmalenorum) might have somediagnostic value alone or in combination,larger samples are clearly needed to assesssuch modal differences.

Body weight data suggest that Coendouroosmalenorum is one of the smallest livingerethizontids, but the adult holotype (INPA2586, 600 g) may have been emaciated (see


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below), and the other weighed specimen(INPA 2587, 800 g) is a subadult. By com-parison, 42 postjuvenile4 specimens of C.melanurus weighed by Richard-Hansen et al.(1999) ranged from 1500 to 2600 g, and 70specimens of C. prehensilis weighed by thesame authors ranged from 1800 to 5800 g.

VARIATION: With only three specimens inhand, the possible significance of observedcharacter variation in Coendou roosmaleno-rum is hard to evaluate. Although certain dif-ferences observed in our material are plau-sibly attributable to ontogeny, others are not.An example of the former is the rostrum ofINPA 677 (an old adult), which is relativelyand absolutely larger than the rostrums ofINPA 2587 (a subadult) and INPA 2586 (ayoung adult). An example of the latter is thegrayish dorsal fur of INPA 2586 versus thebrownish fur of the other two skins.

Although we examined specimens fromboth banks of the Rio Madeira, a major Am-azonian tributary and one of the largest riversin the world, the similarities uniting our ma-terial as a taxon diagnosable from otherforms of Coendou are more persuasive thanthe few differences that segregate our left-bank examples (INPA 2586, 2587) from ourright-bank singleton (INPA 677). However,the more inflated sinuses of the former spec-imens are perhaps noteworthy in this context,as is the extremely small toothrow of the lat-ter. Future studies based on larger samplesfrom both populations will doubtless contrib-ute to a more informed judgement about theirtaxonomic status.

NATURAL HISTORY: According to infor-mation kindly summarized for us by Marcvan Roosmalen, the holotype was capturedby caboclos at Novo Jerusalem when itemerged from a felled tree. Kept alive as apet, the animal was released every eveningto roam freely in the surrounding terra firmeforest, returning at dawn to sleep throughoutthe day in a box underneath its owners’house. This individual subsequently lived forseveral months in captivity at Manaus, whereit maintained its nocturnal habits (sleeping in

4 Richard-Hansen et al. (1999) judged their weighedspecimens to be adults, but those authors did not employdental criteria to assess maturity. Most subadult porcu-pines are externally indistinguishable from adults.

a tree hole by day), and ate a variety of seedsand fruit. It died of unknown causes whileseeming perfectly healthy.

The paratype from Santa Maria was shotat night by a caboclo boy who mistook itseyeshine for that of a paca (Cuniculus paca)as it was walking on the ground no morethan 100 m from the shore of Lago Matupiri,a blackwater lake. The local vegetation wastall ‘‘seringal’’ forest growing on terra firmewith abundant rubber trees (Hevea brasilien-sis), Brazil nuts (Bertholletia excelsa), ingas(Inga spp.), and other trees known locally asorelha de macaco (Enterolobium schomburg-kii), bacuri (Rheedia macrophylla), and tap-ereba (Spondias mombin).

The paratype from 49 km E Porto Velhowas taken in the course of faunal rescue ef-forts at the Samuel hydrolectric dam site, butthe ecological circumstances of its captureare unknown.

REMARKS: Three specimens (two malesand one female) collected at the Samuel hy-droelectric dam site (49 km E of Porto Velhoon BR 364) were identified by Naiff et al.(1996) as the ‘‘Black dwarf porcupine’’ ofEmmons (1990: 199), a taxon that is tech-nically known as Coendou nycthemera Ol-fers (see Voss and Angermann [1997], whoexplain that C. koopmani Handley and Pineis a junior synonym). One of those speci-mens (the female, INPA 677) is a paratypeof C. roosmalenorum, but the whereaboutsof the two male specimens is unknown. It isdifficult to understand how INPA 677 couldpossibly have been mistaken for Emmons’black dwarf porcupine, which was clearlydescribed by her as having entirely spinyblackish upperparts. One possible explana-tion is that the two missing specimens fromNaiff’s series were, in fact, C. nycthemera(which is known to occur on the right bankof the lower Madeira near Borba), and thatINPA 677 was mistaken for a conspecificlong-furred juvenile. Whether or not thisconjecture is correct, future collectors shouldbe alert to the possibility that two small spe-cies of Coendou may be sympatrically orparapatrically distributed along the right(east) bank of the Madeira.

SPECIMENS EXAMINED: Brazil—Amazonas,Novo Jerusalem (INPA 2586), Santa Maria


Fig. 15. Collection localities and sight records for members of the Coendou vestitus species group.Numbered symbols for C. vestitus (n), C. pruinosus (●), C. ichillus (m), and C. roosmalenorum (.) arekeyed to entries in the accompanying gazetteer (appendix). The geographic configuration of localities9–11 is approximate because specimen labels provide only river names. Western Amazonia (as definedby Voss and Emmons, 1996: 8) is that part of the basin west of the Rio Negro and the Rio Madeira.The Cashinahua Indian village of Balta is the source of an unvouchered record of small porcupinesdiscussed in the text.

(INPA 2587); Rondonia, 49 km E Porto Vel-ho (INPA 677).

DISCUSSION

All published inventories of rainforestmammals from western Amazonia (i.e., westof the Negro and Madeira rivers; see fig. 15)have hitherto reported only one species ofporcupine at any given site (Patton et al.,1982; Janson and Emmons, 1990; Woodmanet al., 1991; Hutterer et al., 1995; Peres,1999; Fleck and Harder, 2000; Patton et al.,2000). Invariably, the single reported speciesis a large (3–5 kg) animal that has been iden-tified as either Coendou prehensilis or C. bi-color. By contrast, small (1–2 kg) porcupineshave long been known to occur sympatricallywith C. prehensilis in eastern Amazonia (Ca-

banis, 1848; Walsh and Gannon, 1967; Pine,1973; Husson, 1978; George et al., 1988;Handley and Pine, 1992; Vie, 1999; Voss etal., 2001).5

This report provides the first definite re-cords of small Coendou from western Ama-zonia, where they are almost certainly sym-patric with one or more large (prehensilis-like) porcupines. Why these diminutive spe-cies, and perhaps others of the same size

5 The small porcupine that occurs throughout north-eastern Amazonia is Coendou melanurus (see Voss etal., 2001), but this species has often been misidentifiedas C. insidiosus (e.g., by Husson, 1978). The small spe-cies of southeastern Amazonia is C. nycthemera, previ-ously known by its junior synonym C. koopmani (seeVoss and Angermann, 1997). The large porcupines cur-rently known as C. prehensilis are unrevised and mayrepresent a complex of closely related species.


class, have remained undetected for so longbegs explanation. Two hypotheses merit dis-cussion.

First, it is possible that small porcupineshave very restricted distributions in westernAmazonia and simply do not occur in re-gions where most faunal inventory projectshave been carried out. By this interpretation,Coendou ichillus and C. roosmalenorummight survive as relicts of a formerly wide-spread ancestral taxon that went extinct else-where for ecological reasons. Perhaps vesti-tus-like porcupines are narrowly adapted tosome particular type of forest that occursonly along the middle Rio Madeira and ineastern Ecuador. Unfortunately, vegetationalmapping in this region is currently too coarseto test such ad hoc conjectures.

Alternatively, the absence of other recordsof small porcupines from western Amazoniacould be an artifact of inadequate collectingefforts in this still largely inaccessible region.Although impressive faunal sampling effortshave been made at a few sites, small porcu-pines appear to be difficult to detect by anyknown inventory method (e.g., those de-scribed by Voss and Emmons, 1996). In-stead, most collected specimens appear tohave been encountered by chance. To thebest of our knowledge, none have ever beentaken in traps, and few have been taken bydeliberately hunting for them at night.

Only annecdotal evidence is available tosupport the hypothesis that small porcupinesare actually widespread in western Amazon-ia. For example, Cashinahua hunters at Balta(in Departamento Ucayali, Peru: fig. 15) toldA. L. Gardner about a morphologically dis-tinctive small porcupine that occurs sympat-rically with Coendou bicolor at that locality,but only the latter was collected (see Vossand Emmons, 1996: appendix 9). BecauseBalta is far from the known range of both C.ichillus and C. roosmalenorum, it is not sur-prising that the small porcupine described bythe Cashinahua is clearly not referable to ei-ther taxon (A. L. Gardner, personal com-mun.).

We predict that more new species of smallCoendou, perhaps including some that be-long to other clades than the vestitus group,will eventually be found in western Amazon-ia, but opportunities to collect specimens of

these elusive animals are so infrequent thatnone should be ignored. In particular, faunalrescue operations following hydroelectricdam construction afford unique opportunitiesfor wildlife researchers to examine largenumbers of porcupines (Walsh and Gannon,1967; Mascarenhas and Puorto, 1988; Vie,1999). Regrettable as such events are fromother perspectives, each is a rare chance togain valuable information about erethizontiddiversity.

ACKNOWLEDGMENTS

Numerous museum curators generously al-lowed us to borrow rare material from theircollections, and tolerantly extended overdueloans for several years to make this reportpossible. In this context, we are especiallygrateful for the assistance of Renate Anger-mann (ZMB), Luis Albuja (EPN), C. O. Han-dley, Jr. (USNM), Paula Jenkins (BMNH),Daniel Lew (MHNLS), Bruce Patterson(FMNH), Maria Rutzmoser (MCZ), andMichel Tranier (MNHN). We thank PeterGoldberg for the cranial photographs in thisreport, Dennis Finnin for the skin photo-graphs, Pat Wynn for the line art, Phil De-Vries for the color slide reproduced in figure12, and Marc van Roosmalen for the colorprints reproduced in figure 13. Rob Andersonkindly tallied information about unexaminedspecimens in Venezuelan museums, and heremeasured other material in Ecuadorean mu-seums at our request; Francisco Bisbal(EBRG), Daniel Lew (MHNLS), Roger Perez(MBUCV), Belkis Rivas (MHNLS), and Ja-vier Sanchez (EBRG) facilitated Rob’s effortson our behalf. Thanks are also due to themanagement of La Selva Jungle Lodge andto Phil DeVries for hosting our brief visit tothe surrounding habitat of Coendou ichillus in1996.

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APPENDIX: Gazetteer

Below are listed all localities at which exam-ined specimens referable to the Coendou vestitusgroup have been collected. Italicized place namesare those of the largest political divisions (state/department/province) within countries; boldfaceidentifies collection localities as they appear in thetext of this report. Unless recorded by the collec-tor, geographic coordinates (degrees and minuteslatitude/longitude) and elevation above sea levelin meters (m) are provided in square brackets witha cited secondary source for these data. The spe-cies name, name(s) of collector(s), and date(s) ofcollection are separated from the locality entry bya colon. Numbered localities are plotted on theaccompanying map (fig. 15).

BRAZIL

1. Amazonas, Novo Jerusalem on Lago Ma-tupirizinho (58339280S, 618079200W, ,100m): Coendou roosmalenorum (coll. M. G.M. van Roosmalen and T. van Roosmalen,23 November 1996).

2. Amazonas, Santa Maria on Lago Matupiri(58339150S, 618159470W, ,100 m): Coen-dou roosmalenorum (coll. M.G.M. vanRoosmalen and T. van Roosmalen, April1997.

3. Rondonia, 49 km E Porto Velho [coordi-nates unknown, ca. 100 m]: Coendou roos-malenorum (coll. ‘‘Equipe J. Arias,’’ 9 July1985).

COLOMBIA

4. Cundinamarca, Quipile [48459N, 748329W;DMA, 1988], W of Bogota: Coendou ves-titus (coll. Niceforo Marıa, February 1925).

5. Cundinamarca, San Juan de Rıo Seco[48519N, 748389W, 1303 m; Paynter andTraylor, 1981]: Coendou vestitus (coll. Ni-ceforo Marıa, 1923–1925).

6. Meta, [Serranıa de] La Macarena, PicoRengifo [ca. 38069N, 738559W; Paynter and

Traylor, 1981], 4500 ft: Coendou pruinosus(coll. K. von Sneidern, 2 April 1957).

7. Meta, Villavicencio [48099N, 738379W;Paynter and Traylor, 1981], 500 m: Coen-dou pruinosus (coll. R. M. Gilmore, 15June 1939).

8. Norte de Santander, Alturas de Pamplona[78239N, 728399W, 2340 m; Paynter andTraylor, 1981]: Coendou pruinosus (coll.Niceforo Marıa, July 1950).

ECUADOR

9. Pastaza, Rıo Conambo [ca. 18529S,768479W, ca. 300 m at village of Conambo;Paynter and Traylor, 1977]: Coendou ich-illus (coll. C. Estrella, 18 February 1966).

10. ? Pastaza, Rıo Pastaza [no other localitydata]: Coendou ichillus (coll. von Bau-mann-Roosevelt Expedition, July 1936).

11. Pastaza, Rıo Yana Rumi [presumably theRıo Yanarumiyacu, with mouth at 18389S,768599W; DMA, 1987]: Coendou ichillus(coll. R. Olalla, 19 October 1934).

12. Sucumbıos, La Selva Jungle Lodge(08309S, 768229W): Coendou ichillus (un-vouchered sightings by J. E. Cadle, P. J.DeVries, L. H. Emmons, and R. S. Voss,July 1996).

PERU

13. Loreto, Iquitos region [ca. 38469S,738159W; Stephens and Traylor, 1983]:Coendou ichillus. (purchased by P. Soini,1971).

VENEZUELA

14. Distrito Federal, Caracas [ca. 108309N,668559W, 917 m; Paynter, 1982]: Coendoupruinosus (collector unknown, ca. 1926).

15. Merida, Finca el Cumbe, 3.5 km SW Zea[88239N, 718479W; Paynter, 1982], eleva-


tion 1500 m: Coendou pruinosus (see Con-cepcion and Molinari, 1991).

16. Merida, Merida (and localities presumablynear this city including ‘‘Montana Sierra2100[?] m’’ and ‘‘Montana del Valle 2600m’’) [ca. 88369N, 718089W]: Coendou prui-nosus (coll. S. Briceno Gabaldon e hijos,ca. 1900–1905).

17. Merida, Montanas de la Pedregosa, ele-vation 2500 m [ca. 88369N, 718129W; Payn-ter, 1982]: Coendou pruinosus (coll. S. Bri-ceno Gabaldon e hijos, 14 January 1905).

18. Merida, Montanas de Tabay, elevation2600 m [ca. 88389N, 718049W; Paynter,1982): Coendou pruinosus (coll. S. BricenoGabaldon e hijos, 28 January 1904).

19. Zulia, El Rosario [ca. 98099N, 728369W;Handley, 1976], 48 km WNW Encontra-dos, 54 m: Coendou pruinosus (coll. A.Tuttle et al., 1 March 1968).

20. Zulia, Mision Tukuko [98509N, 728529W;Voss, 1991], elevation 200 m: Coendoupruinosus (coll. R. S. Voss, 7 July 1986).

revisionary notes on neotropical porcupines (rodentia

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