A New Species of Spiny Pocket Mouse (Heteromyidae ...

Copyright q American Museum of Natural History 2002 ISSN 0003-0082

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Number 3382, 26 pp., 5 figures, 2 tables August 16, 2002

A New Species of Spiny Pocket Mouse(Heteromyidae: Heteromys) Endemic to

Western Ecuador

ROBERT P. ANDERSON1,2 AND PABLO JARRIN-V.3,4

ABSTRACT

Whereas previous treatments have considered Heteromys australis the only spiny pocketmouse present in Ecuador, morphological and morphometric analyses of specimens from Ec-uador and southwestern Colombia reveal the presence of two species of the genus. Heteromysaustralis is distributed in evergreen forests from eastern Panama and western Venezuelathrough Colombia to extreme northwestern Ecuador, where it inhabits wet, unseasonal areasof the Choco and adjacent western slopes of the Andes. We here describe a new species,Heteromys teleus, found only in evergreen forests of central-western Ecuador, in areas lessmesic and more seasonal than those characteristic of H. australis. Both species possess darkgray dorsal pelage, but H. teleus differs by larger (nonoverlapping) measurements of the hindfoot and distinctive cranial proportions. Most notably, the rostrum of the new species is strik-ingly wide and massive, and the interparietal is narrow and rounded (in contrast to the wide,diamond-shaped interparietal of H. australis). The ranges of the two species together conformto the previously recognized Chocoan evergreen-forest fauna of western Colombia and north-western Ecuador. However, the restriction of H. teleus to evergreen but seasonal forests of thesouthern Choco (transitional between the relatively unseasonal evergreen forests of the centralChoco to the north and highly seasonal xeric regions to the south) is unique within currently

1 Roosevelt Postdoctoral Research Fellow, Division of Vertebrate Zoology (Mammalogy), American Museum ofNatural History. e-mail: [email protected]

2 Natural History Museum and Department of Ecology & Evolutionary Biology, University of Kansas, Lawrence,KS 66045.

3 Museo de Zoologıa, Centro de Biodiversidad y Ambiente, Departamento de Ciencias Biologicas, Pontificia Univ-ersidad Catolica del Ecuador, Avenida 12 de Octubre y Roca, Apartado 17–01–2184, Quito, Pichincha, Ecuador.

4 Present address: Graduate Student, Center for Ecology and Conservation Biology, Department of Biology, BostonUniversity, Boston, MA 02215. e-mail: [email protected]

2 NO. 3382AMERICAN MUSEUM NOVITATES

recognized species of mammals. Biogeographic overviews hint at similar patterns in othergroups, but more alpha-taxonomic research is necessary to evaluate mammalian distributionalpatterns in the region properly. Most suitable habitat for H. teleus has been converted toagricultural uses, and its current distribution is likely restricted to a handful of small-to-me-dium-sized forest patches.

RESUMEN

Mientras que estudios previos han considerado a Heteromys australis como el unico ratonde abazones (5 raton bolsero; raton bolson; raton mochilero) presente en Ecuador, analisismorfologicos y morfometricos de ejemplares provenientes de este paıs y del suroccidente deColombia revelan la presencia de dos especies del genero. Heteromys australis esta distribuidoen bosques siempreverdes, desde el este de Panama y occidente de Venezuela, a traves deColombia, hasta el extremo noroccidental de Ecuador, donde habita areas humedas y sin es-tacionalidad del Choco y estribaciones occidentales de los Andes. En el presente trabajo sedescribe una nueva especie, Heteromys teleus, hallada solamente en bosques siempreverdesde la region centro-occidental del Ecuador, en areas menos humedas y mas estacionales queaquellas caracterısticas para H. australis. Las dos especies poseen un pelaje dorsal de color-acion grisacea oscura, pero H. teleus se caracteriza por tener mayores medidas (sin sobrela-pamiento) de la longitud de la pata trasera y proporciones craneales distintivas. Mas notoria-mente, el rostro de la nueva especie es particularmente mas ancho y robusto, y el interparietalangosto y redondeado (contrastando con el interparietal ancho y en forma de diamante de H.australis). Las distribuciones de las dos especies juntas se ajustan a la previamente aceptadafauna de los bosques siempreverdes del Choco en el occidente de Colombia y el noroccidentedel Ecuador. Sin embargo, la distribucion de H. teleus es unica entre las especies conocidasde mamıferos, estando restringida a bosques siempreverdes, pero con marcada estacionalidad,en el sur del Choco (transicionales entre los bosques siempreverdes de poca estacionalidaddel Choco central al norte y las regiones aridas, fuertemente estacionales hacia el sur). Revi-siones biogeograficas sugieren patrones similares para otros grupos, pero mas investigacionde la taxonomıa alfa es necesaria para evaluar adecuadamente los patrones de distribucion delos mamıferos en la region. La mayor parte del habitat adecuado para H. teleus ha sidomodificado para la agricultura, y su distribucion actual probablemente esta restringida a unospocos parches de vegetacion original de extensions relativamente pequenas o medianas.

INTRODUCTION

The lowland faunas of the mesic Chocoand the xeric Tumbes regions come into con-tact west of the Andes in Ecuador (Chapman,1926; Albuja-V. et al., 1980; Cracraft, 1985;Best and Kessler, 1995). This faunal turnovercorresponds to a rapid and relatively uniformdecrease in precipitation from north to southdue primarily to the influence of the coldHumboldt Current in the south (Chapman,1926; Lynch and Duellman, 1997; Ceron etal., 1999). Evergreen forests of western Ec-uador belong to the Choco, whereas decid-uous (‘‘dry’’) forests and xeric, nonforestedareas to the south comprise the Tumbes re-gion (Cracraft, 1985). The distributions ofmany mammalian groups typical of lowlandNeotropical rainforests terminate west of theAndes in this region (Emmons, 1997; Tirira-

S., 1999). Furthermore, several clades pre-sent in rainforests west of the Andes (trans-Andean) in Central America and western Co-lombia encounter their southernmost distri-butions in western Ecuador and are not foundin Amazonian forests to the east (e.g., Diplo-mys, Heteromys, Hoplomys, and Tylomys;Hershkovitz, 1972; Emmons, 1997). We hereconsider Heteromys, which shows a uniquebiogeographic history and has been used inseveral recent geographic studies (Anderson,1999; Anderson and Soriano, 1999; Ander-son et al., 2002a, 2002b).

The family Heteromyidae originated inNorth America, and three extant subfamiliesare recognized: Dipodomyinae (kangaroorats and kangaroo mice), Perognathinae(silky pocket mice), and Heteromyinae(spiny pocket mice). While the Dipodomyi-nae and Perognathinae have long fossil re-

2002 3ANDERSON AND JARRIN-V.: HETEROMYS FROM ECUADOR

cords in North America (Wood, 1935), mem-bers of the subfamily Heteromyinae showonly Recent and Pleistocene records (Wah-lert, 1993, removed the extinct genera Proh-eteromys, Diprionomys, and Peridiomysfrom the Heteromyinae). Despite this lack offossil sequence, heteromyines comprise awell-defined monophyletic group distinctfrom either of the two other living subfami-lies (Hafner, 1981; Hafner and Hafner, 1983;Wahlert, 1991).

Two genera exist in the subfamily Heter-omyinae (Heteromys and Liomys), and theirconstituent species range latitudinally fromextreme southern Texas (United States) toEcuador. Within Heteromys, six (Williams etal., 1993) or seven (Patton, 1993) species arecurrently recognized, although more proba-bly exist (Handley, 1976; Rogers, 1990; An-derson, 1999). Members of the genus Het-eromys constitute the only heteromyids tohave dispersed to South America (Hershkov-itz, 1972; Simpson, 1980), and the timingand number of colonization events remaincontroversial (Anderson and Soriano, 1999;Anderson et al., 2002a).

In South America, Heteromys is restrictedto the northwestern reaches of the continent,where two species are known to be widelydistributed and one has a marginal range.Heteromys anomalus is found in deciduousand moderately mesic evergreen tropical for-ests across the Caribbean coast and northernfoothills of Colombia and Venezuela, as wellas on the islands of Margarita, Trinidad, andTobago (Handley, 1976; Williams et al.,1993; Anderson, 1999) and from relativelyxeric regions of the upper Magdalena Valleyin Colombia (Hernandez-Camacho, 1956;Anderson, 1999). In contrast, H. australis in-habits very mesic rainforests in the Pacific(Chocoan) lowlands of Ecuador, Colombia,and eastern Panama as well as mesic mon-tane forests eastward in the Colombian An-des. It ranges from sea level to circa 2500 min sufficiently wet areas (Anderson, 1999)and has also been reported from the Cordil-lera de Merida in western Venezuela (An-derson and Soriano, 1999). In addition tothese two widely distributed species, a mem-ber of the H. desmarestianus complex inhab-its high elevations of the Serranıa del Darienin extreme northwestern Colombia (provi-

sionally referred to as H. d. crassirostris byAnderson, 1999).

Thus, recent works have considered Het-eromys australis the only spiny pocketmouse present in Ecuador (Albuja-V., 1991;Patton, 1993; Williams et al., 1993; Emmons,1997; Anderson, 1999; Eisenberg and Red-ford, 1999; Nowak, 1999; Tirira-S., 1999).Thomas (1901) described H. australis fromSan Javier, in the coastal lowlands of Es-meraldas province in northwestern Ecuadornear the Colombian border. In the next 100years, the only specimens of the genus re-ported from the country derived from thetype locality and two additional sites in thenorthwestern part of the country (Bulim andRıo Palenque Science Center; Anderson,1999), and one locality in the central-westernregion (Jauneche; Albuja-V., 1992). Here weexamine a much larger number of specimensfrom Ecuador and southwestern Colombia inorder to reevaluate the morphological varia-tion, distributional extent, and taxonomic sta-tus of the spiny pocket mice present in thisregion.

MATERIALS AND METHODS

MUSEUM SPECIMENS

We examined 58 specimens of Heteromysfrom Ecuador, including most known fromthat country (and all known to us in muse-ums in Canada, Ecuador, and the UnitedStates). We compared the Ecuadorian speci-mens with available material of H. australisfrom southwestern Colombia and with rep-resentative samples of other recognized spe-cies in the genus, including the majority ofextant holotypes. Locality information notprovided by the collector appears in brackets.Where original elevation was reported infeet, we provide that datum as well as themetric equivalent to the nearest whole num-ber. Localities in Ecuador and southwesternColombia are detailed in the Gazetteer (ap-pendix 1). Specimens examined (see appen-dix 2) are housed in the following museumcollections (* denotes museums holdingspecimens from Ecuador): Academy of Nat-ural Sciences, Philadelphia (ANSP); Ameri-can Museum of Natural History, New York(AMNH*); Departamento de Ciencias Biol-ogicas, Escuela Politecnica Nacional, Quito


(EPN*); Field Museum, Chicago (FMNH*);Instituto de Ciencias Naturales, UniversidadNacional de Colombia, Bogota (ICN); Mu-seo de Zoologıa, Pontificia Universidad Ca-tolica del Ecuador, Quito (QCAZ*); MuseoEcuatoriano de Ciencias Naturales, Quito(MECN*); Museum of Comparative Zoolo-gy, Harvard University, Cambridge (MCZ);Royal Ontario Museum, Toronto (ROM*);United States National Museum of NaturalHistory, Washington, D.C. (USNM*); Univ-ersidad del Valle, Cali (UV); University ofKansas Natural History Museum, Lawrence(KU*); University of Michigan Museum ofZoology, Ann Arbor (UMMZ); and Univer-sity of Wisconsin Zoological Museum, Mad-ison (UWZM*).

We also searched for specimens of the ge-nus from Ecuador in the following museums,but did not find any records: CaliforniaAcademy of Sciences; Carnegie Museum ofNatural History; Florida Museum of NaturalHistory, University of Florida; LouisianaState University Museum of Natural Science;Michigan State University Museum; Muse-um of Texas Tech University; Museum ofVertebrate Zoology, University of California,Berkeley; Natural History Museum of LosAngeles County; Texas Cooperative WildlifeCollection, Texas A&M University; and Uni-versity of New Mexico Museum of South-western Biology.

We examined external and cranial mor-phological characters, making comparisonsamong specimens of approximately the sameage. Specimens were assigned to age classesbased on patterns of toothwear and molt, fol-lowing Rogers and Schmidly (1982). Mor-phological nomenclature follows Wahlert(1985) and Anderson (1999). To assess thehabitat requirements of the species in Ecua-dor, we used available published reports andattempted to contact all living collectors.

MEASUREMENTS

We here define and illustrate 11 standardcranial measurements for Heteromys (fig. 1).These linear distances generally follow Gen-oways (1973), but we add parietal breadth,modify nasal length, substitute squamosalbreadth for mastoid breadth, and follow Rog-ers and Schmidly (1982) in replacing depth

of braincase with skull depth. Also, we re-name greatest length of skull and alter thewording of Genoways (1973) to unify a tele-graphic style. This current suite of measure-ments better reflects the morphology of thegenus Heteromys relative to its sister genusLiomys, for which the measurements of Gen-oways (1973) were originally designed. Thenasal length and mastoid breadth of Ander-son (1999) and Anderson and Soriano (1999)represent the nasal length and squamosalbreadth presented here, not those of Geno-ways (1973).

We took these measurements on all avail-able intact skulls to the nearest 0.01 mm withdigital calipers (except specimens from theICN and UV collections, which were mea-sured to the nearest 0.1 mm). Only adultspecimens (age classes 4–6) were used forstatistical analyses (see below). Externalmeasurements and mass were copied fromspecimen tags, but measurements taken onlyto the nearest centimeter or half centimeteron two series collected from Narino, Colom-bia, in 1912 were excluded. Because somespecimens lacked hind foot measurementsand others were taken by a variety of collec-tors (some of which did not include theclaws), we verified hind foot measurementswith digital calipers for Ecuadorian speci-mens with complete pedal elements in studyskins or fluid-preserved specimens.

Occipitonasal length (ONL): greatest distancefrom anteriormost projection of nasal bones toposteriormost portion of occipital bone (5greatest skull length of Anderson [1999] andAnderson and Soriano [1999] and greatestlength of skull of Genoways [1973]).

Zygomatic breadth (ZB): greatest width across zy-gomatic arches at right angle to longitudinalaxis of cranium.

Rostral length (RL): greatest distance from notchlateral to lacrimal bone to anteriormost projec-tion of nasal bone on same side of cranium.

Nasal length (NL): greatest distance from anter-iormost projection of one nasal bone to its pos-teriormost projection (not necessarily at medialsuture between nasals).

Least interorbital constriction (IOC): least widthacross interorbital constriction at right angle tolongitudinal axis of cranium.

Squamosal breadth (SB): width across squamosalanterior to external auditory meatus at right an-gle to longitudinal axis of cranium.


Fig. 1. Dorsal, ventral, and lateral views of a cranium of Heteromys showing method of takingmeasurements. Abbreviations and measurements are defined in the text.

Maxillary toothrow length (MTR): distance fromanterior lip of alveolus of premolar to posteriorlip of alveolus of third molar.

Interparietal width (IW): greatest transverse widthmeasured from lateralmost projections of inter-parietal bone at right angle to longitudinal axisof cranium.

Interparietal length (IL): greatest distance fromanteriormost projection of interparietal bone toposteriormost border of interparietal bone, al-ways taken along medial line of cranium evenwhen notch present in posterior border.

Parietal breadth (PB): greatest width across pa-rietal crests at right angle to longitudinal axisof cranium.

Skull depth (SD): greatest distance from dorsal-

most point of braincase to horizontal planepassing through ventral borders of maxillarycheek teeth and ventral borders of occipitalcondyles (taken by placing skull on glass mi-croscope slide with upper incisors rested overedge of slide, and then subtracting width ofslide).

STATISTICS

Specimens from Ecuador and southwest-ern Colombia were used for quantitativecomparisons. Using species assignmentsbased on our qualitative morphological ex-aminations, we tested for sexual dimorphismin cranial measurements within a species in


a series of univariate two-tailed t-tests ofadult specimens in age class 4; based onthose results, the sexes were pooled in allfurther analyses. For each species, we cal-culated descriptive statistics of adult speci-mens in age class 4 for: external measure-ments, mass, cranial measurements, and twoderived ratios (tail length/head-and-bodylength and interparietal width/zygomaticbreadth). All localities for each species werepooled because sample sizes were too smallat individual sites. Additionally, we conduct-ed two-tailed t-tests comparing means of thetwo species for each measurement or ratio.

We also conducted a principal componentsanalysis (PCA) based on the variance-co-variance matrix of loge-transformed cranialmeasurements of adult specimens in ageclasses 4–6. We interpreted the multivariateaxes by examination of both loadings and co-efficients (elements) of the unit eigenvectorcorresponding to each component. Loadingsare Pearson product-moment correlations be-tween specimen scores on an axis, and therespective loge-transformed measurements.We tested for differences between the speciesfor scores on the first three principal com-ponents, by two-tailed t-tests. All statisticalanalyses were performed in Minitab (1998,version 12.1), and probabilities were com-pared to an a of 0.05 for hypothesis testing.

SYSTEMATICS

Our analyses indicate that two species ofHeteromys are present in Ecuador. One spe-cies includes topotypes of H. australis andcorresponds to the morphological diagnosisof that species (Anderson, 1999). The otherspecies displays proportions and measure-ments distinct from those of H. australis aswell as differences from all other recognizedspecies of the genus. As no available nameexists for it, we describe this taxon as:

Heteromys teleus, new speciesFigure 2

HOLOTYPE: AMNH 64694, adult male;skin and skull in excellent condition; left andright rami of mandible separated. Collectedon 2 June 1923 from Ecuador: ProvinciaGuayas: Cerro Manglar Alto, western slopeby G.H.H. Tate; original number 1536. Tate’s

field camp was located at 1500 ft [457 m],but his notes and field catalog indicate thatthe holotype was taken at 2000 ft [610 m].

PARATYPES: We designate the followingspecimens (adult skins and skulls in goodcondition) from a variety of museum collec-tions as paratypes. See also appendix 2 forother specimens not included in the type se-ries (ICZN, 1999: article 72.4.6; recommen-dation 72B). ECUADOR (10): Cotopaxi: SanFrancisco de las Pampas, Reserva BosqueNublado Otonga [5 Bosque Integral Oton-ga], 1900–2000 m, QCAZ 1788. Guayas:Cerro Manglar Alto, eastern slope, Camp I,1200 ft [366 m] (camp), specimen from 1400ft [427 m], AMNH 64676; Cerro ManglarAlto, western slope, Camp II, 1500 ft [457m] (camp), specimens from 2000 ft [610 m],AMNH 64681, 64682, 64686, 64689. LosRıos: Rıo Palenque Biological Station [5 RıoPalenque Science Center], 220 m, KU149133, 149135; Rıo Palenque Science Cen-ter, Santo Domingo, 47 km S of (by road),USNM 528573; Canton Vinces, ParroquiaPalenque, Jauneche, Estacion Biologica Pe-dro Franco Davila, 50 m, EPN 91.3038.

ETYMOLOGY: Latinized from the Greek te-leos, meaning ‘‘having reached its end, fin-ished, complete’’ (Brown 1956: 785) in ref-erence to the species’ distribution at thesouthern end of the Choco, representing thesouthernmost known distribution of the ge-nus as well as of the superfamily Geomy-oidea.

DISTRIBUTION: Known only from central-western Ecuador (fig. 3) from near the RıoEsmeraldas and Rıo Guayllabamba (circa 08209N) south to the Cordillera de Chongon-Colonche (southern extension of the Cordil-lera de la Costa; circa 18479S). Present fromnear sea level on the coastal plain to circa2000 m on the western slopes of the Andes.Specimens from the Cordillera de Chongon-Colonche may be isolated from other popu-lations, as this low cordillera lies in a matrixof drier vegetation (see Chapman, 1926; Fos-ter, 1992b; Ceron et al., 1999; Sierra et al.,1999b). Note that some localities are docu-mented only by specimens provisionally re-ferred to the species (without examined cra-nial material) as listed in appendices 1 and 2.

DIAGNOSIS: A species of spiny pocketmouse with adults showing the following


Fig. 2. Dorsal, ventral, and lateral views of the holotype of Heteromys teleus (AMNH 64694) anda topotype of H. australis (USNM 113305), both males in age class 4. Note the overall larger size androbustness of H. teleus, as well as its extremely wide rostrum and narrow interparietal.

combination of characters (fig. 2): mesopter-ygoid fossa V-shaped, with long, thin ham-ular processes of pterygoids; optic foramensmall, with exterior margin formed by strongbar of bone; p4 with 3 lophs; skull large(ONL circa 35–38 mm in age class 4), ro-bust, and wide (table 1); rostrum extremelywide, especially basal half; interparietal nar-row and laterally rounded; molars distinctlywide; plantar surface of hind feet naked; dor-sal pelage dark gray or blackish; no lateralochraceous band present; pelage alwaysharsh and spiny; ears small to medium in

size; body size large, with hind foot 35 mmor greater in adults; tail (occasionally) ap-proximately equal to or (more often) slightlylonger than head-and-body length.

DESCRIPTION: Dorsal pelage spiny and darkgray or blackish, sharply contrasting withsoft, pure-white pelage of venter (see illus-tration in Jarrın-V., 2001); ears blackish andsmall to medium in size (table 1); tail usuallylonger than head-and-body length (table 1)and moderately bicolored, with some darkhairs present on ventral surface, especiallydistally; hairs on dorsal surface of hind feet


Fig. 3. Map showing collection localities for Heteromys in Ecuador and southwestern Colombia.Triangles represent localities of Heteromys australis, and circles denote localities of H. teleus. Localitiesare numbered to correspond to data given in appendix 1. Regions above 300 m are shown in grayshading, and areas above 2000 m appear in black. The dashed line in the northwestern part of thecountry denotes the division between coastal regions with a dry season more than 5 months long(southwest of the line) and areas with a shorter dry season (northeast of the line), adapted from Sierraet al. (1999b).


TABLE 1Descriptive Statistics and Results of t-Tests Comparing Adults in Age Class 4 of Heteromys

australis and H. teleus in Western Ecuador and Southwestern Colombiaa


white, or white with moderate infusion ofblack hairs; hairs at base of claws of hindfeet white; inner side of forearms white; hindfeet large (table 1), with naked plantar sur-face; skull large (table 1) and robust (fig. 2);posterior half of premaxilla extremely wide,forming a distinctive shelf along dorsolateralborder of rostrum; anterior roots of zygo-matic arches robust; interorbital constrictionmoderately wide; braincase moderately widewith strong parietal crests, but not inflated;interparietal narrow (table 1) and roundedlaterally; anterodorsal lobe of periotic cap-sule well developed, causing distinct undu-lation in posterior margin of squamosal (seeschematic drawing in Anderson, 1999: 619);incisive foramina wide and bowed; molarslarge and wide; permanent upper premolar(P4) with short fold in anterior border of pos-terior loph; mesopterygoid fossa V-shaped,with hamular processes of pterygoids longand thin; optic foramen small, with exteriormargin formed by strong bar of bone; jugularforamen (anterolateral to occipital condyle)large; p4 with 3 lophs.

DENTAL FORMULA: (incisors 1/1, canines 0/0, premolars 1/1; molars 3/3) 3 2 5 total 20.

MEASUREMENTS OF THE HOLOTYPE (mm):Total length 283, tail length 154, hind foot37, ONL 36.44, ZB 18.68, RL 15.87, NL15.30, IOC 8.59, SB 15.95, MTR 6.05, IW8.01, IL 5.23, PB 12.61, SD 11.33.

COMPARISONS: The dark slaty-gray pelageof Heteromys teleus differentiates it fromseveral species of the genus that have dorsalpelage strongly infused with brown or cin-namon (ochraceous) hairs, causing a grizzledeffect. Among these, H. gaumeri and mem-bers of the H. desmarestianus complex (in-cluding H. goldmani; see Rogers, 1990, andWilliams et al., 1993) inhabit Central Amer-ica far from the known distribution of H. te-leus. Specimens of H. gaumeri and manypopulations of the H. desmarestianus com-plex possess a lateral ochraceous band that isabsent in H. teleus. Furthermore, H. gaumeriis much smaller than the new species (seeEngstrom et al., 1987). Members of mostpopulations of H. anomalus (from Colombia,Venezuela, and Trinidad and Tobago) havepale, especially grizzled pelage, in stark con-trast to H. teleus. Heteromys teleus can beseparated from all populations of H. anom-

alus by the narrower rostrum, relatively lon-ger (usually sharply bicolored) tail, and larg-er, rounded ears of H. anomalus, as well asby differences in cranial proportions (rela-tively wider skull in H. teleus and narrowerinterorbit in H. anomalus).

When compared with other species of Het-eromys that have dark pelage, H. teleus iseasily distinguished by its extremely widerostrum. Heteromys oresterus (endemic tothe Cordillera de Talamanca in Costa Rica)has a characteristically long, thin rostrum.Heteromys nelsoni (distributed in southernMexico and western Guatemala) is even larg-er than the new species (Williams et al.,1993), with a more elongated skull (includ-ing rostrum), larger ears, dusky inner fore-arms, and soft dorsal pelage. The only otherspecies of Heteromys with a similarly widerostrum is a member of the H. desmaresti-anus complex present in the Darien region ofeastern Panama and northwestern Colombia(provisionally referred to as H. d. crassiros-tris in Anderson, 1999). That taxon, how-ever, is one of the smallest of the genus(ONL 32.2–35.1 mm and hind foot length29–34 mm in age classes 4–5; Anderson,1999), has barely perceptible temporal fossae(erroneously named masseter-temporal fos-sae by Anderson, 1999), straight posteriormargin of the squamosal, tapered incisive fo-ramina, chocolate-brown dorsal pelage, anda relatively shorter tail.

Heteromys australis, which has a geo-graphic range that probably comes into con-tact with that of H. teleus in northwesternEcuador (or did historically; see appendices1 and 2), merits closer comparisons. The twospecies are virtually indistinguishable exter-nally (see below), but differ in cranial sizeand proportions. No sexual dimorphism wasdetected for any cranial measurement withineither species (P 5 0.18–0.98). In contrast,all raw measurements were significantly dif-ferent between the two species (P rangesfrom , 0.0001 to 0.026 for significant com-parisons; table 1) except for parietal breadth(P 5 0.71). Generally, H. teleus is larger, butthe interparietal width averages greater in H.australis. No overlap occurs between speci-mens (of any age class) of the two species inobserved hind foot measurements (range 535–40 mm for H. teleus, 29–34 mm for H.


Fig. 4. Plot of specimen scores on PC I and PC II from a principal components analysis of loge-transformed cranial measurements of adult specimens (age classes 4–6) of Heteromys from Ecuadorand southwestern Colombia examined in this study. Triangles represent specimens of H. australis, andcircles denote those of H. teleus. Axes are scaled relative to their eigenvalues (proportion of the variationexplained).

TABLE 2Results of Principal Components Analysis of loge-Transformed Cranial Measurements for AdultSpecimens (Age Classes 4–6) of Heteromys from Ecuador and Southwestern Colombia Examined

in This Study(Elements of the unit eigenvectors, loadings, eigenvalues, and proportion of variance explained are

given for each of the first three principal components.)

australis), or for the derived ratio of inter-parietal width/zygomatic breadth (range 50.423–0.520 for H. teleus, 0.555–0.689 forH. australis).

Multivariate analyses support the morpho-logical distinctiveness of Heteromys teleusfrom H. australis. The two species separateclearly along the first principal component ofthe PCA undertaken on adult specimens (fig.4), with no overlap between the two. The co-

efficients and loadings indicate that PC I rep-resents a contrast between overall size andinterparietal width (table 2). All measure-ments but interparietal width load negatively;thus, specimens with low scores on PC I (leftportion of fig. 4) have relatively large valuesfor all measurements except interparietalwidth. In contrast, specimens with highscores (right portion of fig. 4) hold small val-ues for most measurements, but large ones


Fig. 5. Plot of interparietal width versus zygomatic breadth for all available specimens (age classes2–6) of Heteromys from Ecuador and southwestern Colombia examined in this study. Triangles representindividuals of H. australis, and circles denote those of H. teleus. Although the two species overlap oneither of the variables taken individually when considering specimens of various age classes (as here),the bivariate plot shows that specimens of H. teleus consistently have a narrower interparietal relativeto the zygomatic breadth of the skull. Considering all age classes together, the two species do not overlapin the interparietal width/zygomatic breadth ratio.

for interparietal width. Species means werehighly significantly different on PC I (P ,0.0001), but not on PC II or PC III (P 50.24–0.97).

In sum, adults of Heteromys teleus showoverall larger body size than H. australis anddiffer in cranial proportions. The hind foot islonger (nonoverlapping measurements re-gardless of age class) and the zygomaticbreadth wider (nonoverlapping measure-ments for adults in age class 4; table 1). Therostrum is strikingly wider in H. teleus, bothrelatively and absolutely (although not mea-sured), and the anterior roots of the zygo-matic arches are more robust (fig. 2). In ad-dition, the jugular foramina anterolateral tothe occipital condyles are larger in H. teleus,and the molars are wider. Finally, regardlessof age class, the interparietal is laterallyrounded and relatively narrower in the newspecies, in contrast to the interparietal of H.australis, which is wide and flattened-dia-mond shaped (at least in this geographic re-gion; figs. 2 and 5).

Both species possess dark gray dorsal pel-age, but whereas the dorsal pelage of Het-eromys teleus is always harsh and spiny, thatof H. australis varies with altitude (spiny in

lowlands, softer in highlands; see Anderson,1999). The length of the hind foot representsthe only reliable external character we knowof for distinguishing the two species. Eventhe single available specimen of H. teleus inage class 1 has a larger hind foot (35 mm)than does any specimen of H. australis fromEcuador of any age class (the largest knownspecimens have a hind foot measuring 34mm). Nevertheless, definitive identificationsrequire collection of museum voucher spec-imens, given the difficulty of taking accuratehind foot measurements on live animals andthe paucity of available specimens of the ge-nus from Ecuadorian localities in museumcollections.

NATURAL HISTORY: In contrast to somepopulations of Heteromys currently referredto H. desmarestianus in Central America(e.g., Fleming, 1970, 1974a, 1974b, 1977a,1977b; Fleming and Brown, 1975; Vander-meer, 1979; Coates-Estrada and Estrada,1988; Quintero and Sanchez-Cordero, 1989;Martınez-Gallardo and Sanchez-Cordero,1993; Sanchez-Cordero, 1993; Sanchez-Cor-dero and Fleming, 1993; Brewer, 2001) or toH. anomalus in Venezuela (Pirlot, 1963;Rood, 1963; Rood and Test, 1968; Valdez et


al., 1985), very little ecological informationis available for the two species of Heteromyspresent in Ecuador (but see Gonzalez-M. andAlberico [1993] for information on H. aus-tralis in western Colombia and Anderson[1999] for natural history information on H.australis gleaned from specimen tags andfield notes). G.H.H. Tate offered in his fieldnotes of 17 May 1923 at Cerro Manglar Altothis tantalizing report on the species that wenow recognize as H. teleus: the Heteromys‘‘use distinct runways, these are well wornand frequently placed about a foot below thetop of the steep bank of the quebrada [5stream]. Every specimen but one [at eastern-slope camp] taken in quebrada, the otherabout 50 yards above in a cornfield. Onespecimen had a large cockroach in onecheek-pouch. Nocturnal.’’ On 21 May 1923at the western camp, he added that the spe-cies ‘‘is probably distributed in the small wa-tercourses throughout the region. It may bea water rat, but it is certainly a bank-rat, be-ing almost always taken along the banks ofthe quebradas.’’ Another specimen had smallseeds in its pouches.

DISCUSSION

ECOGEOGRAPHY

Within Ecuador, Heteromys australis in-habits areas wetter than the known localitiesof H. teleus. Whereas H. australis is restrict-ed to the extreme northwestern portion of thecountry in very wet forests of the lowlandChoco and on the western versant of the An-des, H. teleus occupies slightly drier (but stillevergreen) forests in the southernmost exten-sion of the Choco, also on the Pacific coastalplain and western slopes of the Andes (fig.3; see also below and appendix 1). Both spe-cies inhabit a wide range of altitudes, fromnear sea level to approximately 2000 m (H.teleus) or to 1400 m (H. australis) in Ecua-dor, although the latter species is known torange up to circa 2500 m in parts of Colom-bia (Anderson, 1999). Lowland localities ofH. australis in Ecuador are restricted to areasreceiving more than 3000 mm of annual pre-cipitation, but the two piedmont localities ofthat species lie in regions of between 2000and 3000 mm of precipitation (IGM, 1995b).Such a shift in tolerances corresponds to low-

er potential evapotranspiration at higher (andthus cooler) elevations (Holdridge et al.,1971) and agrees with patterns for H. aus-tralis throughout the rest of its range (An-derson et al., 2002b). In contrast, northernlocalities of H. teleus hug the 3000-mm iso-pleth of annual precipitation, and southernones enter regions receiving 1300–2000 mmof rainfall or even less (800–1300 mm forCerro Manglar Alto; IGM, 1995b). However,the vegetation on the summits and high west-ern slopes of this last site is nourished by‘‘horizontal’’ precipitation in the form ofmist and fog coming off the Pacific, makingsuch areas functionally much more mesicthan suggested by their direct precipitation(G.H.H. Tate field notes, 1923; see also Fos-ter, 1992b, and Best and Kessler, 1995, fordescriptions of similar sites).

This difference in absolute precipitationbetween collection localities of the two spe-cies correlates with seasonality of rainfall.Regions of Ecuador with over approximately3000 mm in rainfall receive that precipitationrelatively uniformly throughout the year (#4 dry months; Sierra et al., 1999b) and arecategorized as ‘‘humedo tropical’’ in a broadscheme of climates and potential vegetation(IGM, 1995a). All localities of Heteromysaustralis in Ecuador correspond to this cat-egory. In contrast, areas receiving less rain-fall in western Ecuador experience a strongdry season (IGM, 1995a, 1995b; Lynch andDuellman, 1997; see also Sierra et al., 1999a,1999b). All but one locality of H. teleus fallinto or very near areas of the ‘‘tropical mon-zon’’ category, with a pronounced dry seasonfrom June to November (IGM, 1995a). Theone exception is Cerro Manglar Alto, whichhas a more stable climate due to coastal mistsand fog. Hence, the macrogeographic distri-butions of these species in western Ecuadorappear to correspond to climatic factors.

Current faunal and floral classifications ofthe region do not distinguish between areascorresponding to the respective distributionsof Heteromys australis and H. teleus. To-gether, their distributions generally corre-spond to the ‘‘tropical noroccidental’’ andwestern ‘‘subtropical’’ zoogeographic re-gions of Albuja-V. et al. (1980; see also Al-buja-V., 1999), the Colombian-Pacific Faunaof Chapman (1926), and the ‘‘humid tropi-


cal,’’ ‘‘humid subtropical,’’ and ‘‘subhumidtropical’’ regimes of Lynch and Duellman(1997). However, the individual distributionsof the two species do not coincide with de-marcations within the former systems, orwith categories in a recent classification ofvegetation types in the country (Sierra et al.,1999b; see also supporting documentation inSierra et al., 1999a). While Sierra et al.(1999b) recognized 46 vegetational forma-tions, they did not make a distinction be-tween the very wet, relatively unseasonal ar-eas of evergreen forest where H. australis isfound and the slightly drier, often highly sea-sonal (but still evergreen) regions character-istic of H. teleus. Both species have been col-lected in (or within 3 km of, given potentialerror in our geographic coordinates) regionsclassified as ‘‘bosque siempreverde de tierrasbajas,’’ ‘‘bosque siempreverde piemontano,’’and ‘‘bosque simpreverde montano bajo.’’ Inaddition, H. teleus also is documented fromareas of ‘‘bosque de neblina montano’’(Bosque Integral Otonga, the highest recordfor the species in the country) and ‘‘bosquede neblina montano bajo’’ (Cerro ManglarAlto, which is aberrantly mesic due to fogand mist off the ocean). This lack of predic-tive information regarding the individualspecies’ distributions in these phytogeo-graphic, zoogeographic, or bioclimatic clas-sification systems highlights the value of anindependent modeling approach for eachspecies in faunal mapping projects (Petersonet al., 1999, 2002) rather than a vegetation-surrogate method.

CONSERVATION STATUS

Data provided by collectors indicate thatboth Heteromys teleus and H. australis havebeen captured in disturbed as well as primaryforests (appendix 1). The disturbed forestsvary in the forms of low-intensity humanuse, but they include systems that containsmall cultivated plots, that are occasionallyentered by free-ranging livestock, or fromwhich wood has been selectively extractedby local inhabitants. No Ecuadorian collec-tion locality of either species derives from atruly secondary site (recovering from beingclearcut recently), deciduous forest, or open,nonforested area.

Six of the seven localities where Hetero-mys teleus has been collected now corre-spond to landscapes where most of the nat-ural vegetation has been removed, and theother one (Bosque Integral Otonga) lies with-in 3 km of nonnatural vegetation (Sierra,1999; our geographic coordinates have ac-curacy of approximately 3 km). Most of theoriginal vegetation on the coastal plain ofcentral-western Ecuador has been convertedto agriculture, especially plantations of ba-nanas and oil palms (Parker and Carr, 1992;Best and Kessler, 1995). Recent collectionsof H. teleus are from very small reservescontaining relictual forest patches within theagricultural matrix (see appendices 1 and 2).As is typical elsewhere in the Neotropics, in-ventories of small mammals in Ecuador inpast few decades have focused on areas ofremaining natural or seminatural vegetation,and sampling effort for H. teleus in agricul-tural areas is inadequate. However, species ofthe genus Heteromys characteristically in-habit closed-canopy forests (Sanchez-Cor-dero and Fleming, 1993; Anderson, 1999),although H. anomalus also occurs in low-in-tensity or ‘‘subsistence’’ agricultural areasbut not in high-intensity, large-scale indus-trial croplands (Anderson, 1999; Utrera et al.,2000). Hence, while the presence of H. teleusin areas of agricultural plantations in westernEcuador is unlikely, further collecting effortsare needed to establish the status of popula-tions of native mammals there (emulatingCartaya and Aguilera, 1985; Martino andAguilera, 1993; Utrera et al., 2000).

The long-term survival of Heteromys te-leus in small, isolated forest remnants is un-likely (Harrison, 1991). Some larger tracts offorest remain along the western slope of theAndes (Sierra, 1999), but only one localityof the new species is from such high eleva-tions (Bosque Integral Otonga). Among hab-itats that we consider typical for the species,the largest areas of intact forest probably liein the Reserva Ecologica Mache-Chindul,which holds forest from near sea level to cir-ca 700 m in the northern portion of the coast-al cordillera (Sierra, 1999), but H. teleus hasnot yet been collected there. The other largeprotected area within the species’ likelyrange is the Parque Nacional Machalilla, inthe Cordillera de Chongon-Colonche (south-


ern portion of the Cordillera de la Costa).However, much of this national park is de-ciduous forest not suitable for the species,and the park has been heavily logged andgrazed by goats and cattle (L. Emmons, inlitt. January 2002). In addition, all nationalparks and other governmentally protected ar-eas in Ecuador face encroachment and vari-ous human uses, and the future of currentlyintact habitat at Mache-Chindul cannot betaken for granted. Like most rodents, no spe-cies of Heteromys was evaluated for the Li-bro rojo de los mamıferos del Ecuador (Ti-rira-S., 2001), but the limited range suggest-ed by known localities indicates that H. te-leus should be evaluated for potential listing.

In contrast, relatively large expanses ofunseasonal wet forests remain in extremenorthwestern Ecuador, in the range of Het-eromys australis (Sierra, 1999). Because ofextreme rainfall, these regions are not as suit-able for agriculture, but timber is currentlybeing exploited for large-scale industrial use(Parker and Carr, 1992; see also Best andKessler, 1995). Seven of the 12 Ecuadorianlocalities of H. australis currently hold nat-ural vegetation (Sierra, 1999), although theyare within 3 km of areas converted to agri-culture. The others lie in areas now defor-ested (Sierra, 1999), but which held forestsat the dates of collection no more than 15years ago (1987–1996; see appendix 1). TheReserva Cotacachi-Cayapas protects exten-sive premontane forests in northwestern Ec-uador, but no large reserve exists in westernEcuador in unseasonal wet lowland rainfo-rest. While expanses of suitable habitat re-main in the Pacific lowlands of western Co-lombia (IAvH and Etter-R., 1998), little isprotected in national parks (Garces-Guerreroand De la Zerda-Lerner, 1994). Overall, theAndean distribution of H. australis has be-come highly fragmented by deforestation inthe past 50 years, but its lowland distributionappears more intact (Anderson, 1999).

BIOGEOGRAPHY OF HETEROMYS IN SOUTH

AMERICA

The occurrence of Heteromys only west ofthe Andes apparently results from the north-ern origin of the group and contingent, his-torical factors related to its subsequent col-

onization of South America (Hershkovitz,1972; Anderson, 1999). The existence of 19collection localities west of the Andes in Ec-uador, with none known from the easternslopes or Amazonian lowlands, makes thepresence of any species of Heteromys east ofthe Andes in Ecuador unlikely. Albuja-V.(1999) presented a relatively uniform distri-bution of collection localities for bats eastand west of the Andes in the country. Thus,because patterns of fieldwork for small ro-dents in the northern Neotropics are gener-ally similar to those for bats, the genus’ ab-sence east of the Andes is not likely an ar-tifact of biased sampling. Predictive modelsof the potential geographic distribution of H.australis indicated suitable environmentalconditions for the species along both easternand western flanks of the Andes in Ecuador(Anderson et al., 2002a). Heteromys austral-is is not known from the eastern slopes ofthe Andes in Colombia either, although thespecies has been recorded for the headwatersof the Rıo Magdalena near the crest of theCordillera Oriental in southern Colombia. Itsevident failure to cross this low pass, inseemingly suitable habitat, to colonize theeastern versant of the Andes of Colombiaand Ecuador remains mysterious (Chapman,1926: 44; Anderson, 1999; Anderson et al.,2002a).

Heteromys classically has been consideredone of the latest North American groups todisperse to South America, with only a min-imal level of differentiation on that continent(Hershkovitz, 1972; Simpson, 1980; Mar-shall et al., 1982; Webb, 1997). While pocketgophers (Geomyidae), which represent thesister group to heteromyids, have dispersedonly to extreme northwestern South Americawest of the Rıo Atrato/Rıo San Juan low-lands or Bolıvar Geosyncline (Alberico,1990), heteromyids are distributed muchmore extensively (Handley, 1976; Anderson,1999). Although the timing of their entryinto South America is controversial, recentalpha-systematic (species-level) and geo-graphic-modeling research has shown thatHeteromys distributions in South Americaare more complex than previously recog-nized (Anderson, 1999; Anderson and Sori-ano, 1999; Anderson et al., 2002a). The pres-ent discovery of a new species endemic to


western Ecuador adds a new level of com-plexity to the emerging understanding of thegroup’s differentiation in South America.Evolutionary relationships among species ofthe genus remain unresolved (Rogers, 1990),but a phylogenetic study of the subfamilyHeteromyinae using morphological charac-ters is underway (Anderson, unpubl.). Fur-thermore, other undescribed species of Het-eromys exist in South America (Handley,1976; Anderson, 1999). Thus, an overall ac-count of heteromyid diversity and evolutionin South America is not yet possible.

FAUNAL RELATIONSHIPS AND ENDEMISM

Under various names, the faunal turnoverin western Ecuador between Chocoan rain-forest elements and Tumbesan species char-acteristic of southern xeric areas has beenlong established (Chapman, 1926; Albuja-V.et al., 1980; Cracraft, 1985). Many mam-malian species characteristic of trans-Andeanrainforests reach their southernmost geo-graphic distribution in Ecuador west of theAndes (e.g., primates Alouatta palliata andCebus capuchinus; rodents Microsciurus mi-mulus and Oryzomys bolivaris; and the batRhinophylla alethina; Emmons, 1997; Tirira-S., 1999). Most such trans-Andean rainforestspecies are closely related to Amazonian spe-cies in the same genus, and—at least follow-ing current taxonomy—many other rainforestspecies are found on both sides of the Andes(Emmons, 1997; Tirira-S., 1999). In contrast,Heteromys and Tylomys (naked-tailed climb-ing rats) represent the only mammalianclades of North American origin to reachtheir southern termination in western Ecua-dor (Hershkovitz, 1972; Tirira-S., 1999; notethat Reithrodontomys, harvest mice, andCryptotis, shrews, also represent diverseNorth American clades with marginal rangesin South America, but there they have An-dean distributions that terminate in montaneEcuador and Peru, respectively; Hooper,1952; Vivar et al., 1997).

Further, Heteromys teleus holds a uniqueposition among mammals endemic to Ecua-dor. The overwhelming majority of Ecuador-ian endemics are rodents, and H. teleus is theonly one of them found on the western coast-al plain. Among the 27 terrestrial mammals

listed as endemic to the country by Tirira-S.(1999), most are montane (13; found onlyover 1000 m) or from the Galapagos islands(8). Two others (Proechimys gularis andOryzomys tatei; rodents) are Amazonian, oneis of unknown locality within the country(Bolomys punctulatus, a rodent; see Voss,1991), and the remainder (3) are from westof the Andes. Of the three western species(all bats), one remains undescribed (Sturnirasp. A of Tirira-S., 1999), and one has sincebeen reported for southwestern Colombia(Balantiopteryx infusca; Alberico et al.,2000; see also McCarthy et al., 2000). Thelast (Cabreramops [5 Molossops] aequato-rianus) is a rare, high-flying bat only knownfrom two localities in central-western Ecua-dor (Ibanez, 1980; Tirira-S., 1999: 138). Thatspecies’ distribution and evolutionary historyremain enigmatic, but it appears to inhabitareas drier than those typical of H. teleus,including deciduous forests where H. teleusis not known. Albuja-V. (1999) also men-tioned undescribed species of the chiropterangenera Lonchophylla and Sturnira from ex-treme northwestern regions of the country,areas wetter than those inhabited by H. te-leus. Hence, H. teleus may represent the onlymammalian species restricted to seasonal ev-ergreen forests of the southern Choco. How-ever, Emmons and Albuja (1992) mentionedthe possible species-level status of a brocketdeer (currently Mazama americana fuscata)and a white-fronted capuchin (currently Ce-bus albifrons aequatorialis) in central-west-ern Ecuador. The taxonomic status of thoseforms has not yet been resolved, but they doshow distributional patterns similar to that ofH. teleus.

We know of no formalized faunal divisionwithin Ecuador corresponding to the distri-butions of Heteromys australis and H. teleus,but several studies hint that may be the case.Several authors have divided the Choco intonorthern and southern faunal regions formammals and herpetofauna, with the divi-sion present at approximately Buenaventurain central-western Colombia (e.g., Lynch,1979; Hernandez-Camacho et al., 1992).This division is much less marked than thatrecognized for birds between the Choco andthe Nechı region north of the Andes in Co-lombia (Cracraft, 1985). In contrast to these


northern divisions for faunal groups, Gentry(1992: 57) was the first to recognize westernEcuador south of the town of Esmeraldas asa ‘‘unique and distinctive floristic region forwet and moist forest . . . rather than as thetail-end of the Choco flora.’’ Although notformalized as a faunal division, distributionalmaps for rainforest lizards and snakes sug-gest a similar pattern (Dixon, 1979). Fur-thermore, Almendariz and Carr (1992) statethat 42% of the species of herpetofauna thatthey collected in western Ecuador were en-demic to that region, including some speciescharacteristic of moist forests.

Nevertheless, as mentioned by Emmonsand Albuja (1992), Hernandez-Camacho etal. (1992), Voss and Emmons (1996), andCadena et al. (1998), vast regions of the Cho-co remain unsampled or inadequately sam-pled, and taxonomic understanding of smallmammals (and most other taxonomic groups)of this and other adjacent regions remainswoefully incomplete. Thus, fine faunal divi-sions and analyses of endemism remain pre-mature and dependent upon scattered and in-complete collection localities, as well as in-adequate alpha-taxonomic work. Future in-ventories and comprehensive taxonomicrevisions will undoubtedly bring to light cur-rently unrecognized species and refine ourunderstanding of the true distributional pat-terns of small mammals in the region, whichin most cases can now only be imperfectlyapproximated.

ACKNOWLEDGMENTS

We thank the following curators and col-lection managers for access to specimens un-der their care: Ted Daeschler (ANSP); RobertS. Voss and Nancy B. Simmons (AMNH);Luis Albuja-V. (EPN); Bruce D. Patterson,Larry R. Heaney, and William Stanley(FMNH); Alberto Cadena (ICN); MariaRutzmoser (MCZ); Juan Francisco Rivade-neira (MECN); Robert M. Timm and ThorHolmes (KU); Luis A. Coloma (QCAZ);Mark D. Engstrom and Burton K. Lim(ROM); Philip Myers and Priscilla Tucker(UMMZ); Michael D. Carleton, the lateCharles O. Handley, Jr., Linda K. Gordon,Robert D. Fisher, and Richard W. Thoring-ton, Jr. (USNM); Michael S. Alberico (UV);

and Holly McEntee and E. Elizabeth Pillaert(UWZM). The following collectors kindlyprovided crucial unpublished data on thehabitats in which they encountered Hetero-mys in Ecuador: Luis Albuja-V., Alvaro Bar-ragan-Yanez, Felipe Campos-Y., Jaime Cev-allos, Burton K. Lim, Lincoln Nolivos-D.,Andrew T. Smith, Francisco Sornoza-Molina,Diego Tirira-S., and Don E. Wilson. Holly I.McEntee, Fredy Trujillo, and Neal Woodmansupplied information on specimens at theirinstitutions. Rodrigo Sierra made his GISmaps available to us in digital format. LuisAlbuja-V. and Santiago Ron facilitated An-derson’s research in Ecuador. Rene Fonsecaand David Lasso participated in fieldworkwith PJV at Otonga. Luis A. Coloma, LouiseH. Emmons, Hugh H. Genoways, MarcelaGomez-Laverde, Santiago Ron, and RobertS. Voss read and made helpful comments onprevious drafts of the manuscript. The Centrode Biodiversidad y Ambiente (UniversidadCatolica del Ecuador) and Fundacion Otongacontributed funding to PJV’s research. An-derson’s work was supported by a NationalScience Foundation Graduate Research Fel-lowship, the Panorama Society and the E.Raymond and Mary Hall Fund (Universityof Kansas Natural History Museum), a Grantin Aid of Research (American Society ofMammalogists), the Thomas J. Dee Fund(Field Museum), and a Collection StudyGrant and Theodore Roosevelt PostdoctoralResearch Fellowship (American Museum ofNatural History).

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APPENDIX 1: Gazetteer

Collection localities for Heteromys in Ecuadorand southwestern Colombia follow. Only abbre-viated locality names are given here; see appendix2 for full provenience and museum catalog num-bers. Secondary political divisions (Departamen-to, Provincia) are italicized, and localities are ar-ranged from N to S within secondary political di-visions.

Heteromys australis

COLOMBIANARINO

1. Barbacoas [18419N, 788099W; 35 m; Payn-ter, 1997]. Collected by W.B. Richardsonin August 1912. ‘‘Thick forest and impen-etrable jungle’’ on Pacific coastal plain(Chapman, 1917: 641).

2. Buenavista [18299N, 788059W; Paynter,1997], 1200 ft [366 m]. Collected by W.B.Richardson in September–October 1912.General area was ‘‘uncultivated and . . .inhabited [by humans] only along theroad’’ Chapman (1917: 50), but conditionof exact collection site unclear.

3. Reserva Natural del Rıo Nambı [18189N,788039W; Cadena et al., 1998], 1300 m.Collected by A. Cadena, P. Rivas-Pava, andR.P. Anderson in March 1995. Very wetprimary premontane rainforest, with muchfog, epiphytes, mosses, and palms (Cadenaet al., 1998).

4. Planada de Maindes (Junın) [18159N,788109W; Orejuela-Gartner et al., 1982],870 m. Collected by M.S. Alberico in Au-

gust 1981. Pluvial forest (transition be-tween tropical and premontane) with mod-erate extraction of wood by locals; mossand orchids very common (Orejuela-Gart-ner et al., 1982).

ECUADORCARCHI

5. El Pailon [18009N, 788149W; Albuja-V.,1989, 1999], 1400 m (includes sublocali-ties). Collected by L. Albuja-V. and P.Mena-V. in November 1987. Disturbed for-est (i.e., with some selective extraction ofwood) in an area with strongly broken to-pography (Albuja-V., 1989; L. Albuja-V.,personal commun. January 2001).

ESMERALDAS

6. Mataje [18229N, 788439W; 50 m; Albuja-V.,1999]. Collected by L. Albuja-V. and P.Mena-V. in October 1991. Disturbed forest(L. Albuja-V., personal commun. January2001).

7. Bulim [5 Pulun; 18059N, 788409W; Payn-ter, 1993], 160 ft [49 m]. Collected by W.Rosenberg in November 1900. Tropicalzone of coastal plain at foot of mountains(Paynter, 1993). State of forest at time ofcollection unknown but likely quite intact.

8. San Javier [5 San Javier de Cachavı;18049N, 788479W; Paynter, 1993], 60–120ft [18–37 m]. Collected by G. Flemming inMay–June 1900. ‘‘Interior northern Esmer-aldas on coastal plain, on railroad’’ (Payn-ter, 1993). State of forest at time of collec-tion unknown, but likely intact away fromrailroad; exact site of collection unknown.

9. Alto Tambo [08569N, 788329W; 750 m; Ti-rira-S., 1999]. Collected by D. Tirira-S. inDecember 1990. Based only on adult fluid-preserved specimen with small hind foot;not confirmed by cranial material; but noteother nearby localities. Caught in a hand-crafted wooden trap used by local inhabi-tants to capture small rodents for food (seealso Albuja-V. and Mena-V., 1987, for il-lustration of and Cadena et al., 1998, formention of similar traps). Relatively pris-tine forest with low-intensity use by nearbyhuman settlements at time of capture; thenatural vegetation in this area was subse-quently destroyed concomitant with theconstruction of the Ibarra-San Lorenzohighway (D. Tirira-S., in litt. August 2001).

10. Estero Zapote [08559N, 788519W for El Za-pote; IGM, 1985]. At 190 m on ‘‘left bankof the Rıo Santiago approximately an


hour’s journey in motorized canoe fromPlaya de Oro/Luis Vargas Torres’’ (J. Cev-allos, in litt. October 2001; translationours). Collected by J. Cevallos in June1993. Based only on adult study skin withsmall hind foot; not confirmed with cranialmaterial; but note nearby localities. Trap-line was situated in the upper parts of theswamp and included both mature forestand areas regenerating after recent agricul-tural uses; habitat type of capture area notavailable (J. Cevallos, in litt. October2001).

11. Alto Tambo, 2 km S of (08549N, 788339W;collection tag), 700 m. Collected by B.K.Lim and F. Sornoza-Molina in March 1996.General area was disturbed humid forest(B.K. Lim, in litt. August 2001), but theHeteromys was captured on a downed login a swampy area of otherwise well-drained primary forest, which was subse-quently clearcut (F. Sornoza-Molina, in litt.October 2001).

12. Reserva La Mayronga [08539N, 798109W;L. Nolivos-D., in litt. February 2001; seealso Lynch and Duellman, 1997]. Collectedby G. Onore and A. Barragan-Yanez in Oc-tober 1993 and by L. Nolivos-D. and I.Castro in September–October 1998. Onoreand Barragan-Yanez captured the mice byhand after turning over fallen palm fronds(Astrocaryum sp.; common name chambi-ra) in primary humid tropical forest (A.Barragan-Yanez, in litt. October 2001).Several individuals were encountered eachtime a frond was disturbed, but only twoHeteromys (both juveniles) were preservedas voucher specimens; thus it is not knownwhether adults were present there as well.Nuts of the same palm were present un-derneath the fronds. L. Nolivos-D. (in litt.February 2001) provided the following in-formation on his and Castro’s later field-work in the reserve: specimens were foundin disturbed tropical forest (ENSOA man-agement treatment ‘‘Area Bajo ExplotacionNormal’’) with irregular topography and apartially open canopy; two individualswere captured near a swampy area, and theother next to a pambil palm (Iriartea del-toidea).

13. Luis Vargas Torres [5 Playa de Oro;08539N, 788489W; 50 m; Albuja-V., 1989,1999, but see Albuja-V. and Mena-V.,1987; not Tirira-S., 1999]. Collected by L.Albuja-V. in October 1994. Specimenswere trapped in both primary forest fartherfrom the town of Luis Vargas Torres and

disturbed forest (which held some smallplots of cultivars) closer to town (L. Al-buja-V., personal commun. January 2001;see also Albuja-V. and Mena-V., 1987).

14. Hacienda La Granada [08529N, 788319W;Albuja-V., 1989, 1999], 670 m. Collectedby L. Albuja-V. and P. Mena-V. in August1994. Disturbed forest (L. Albuja-V., per-sonal commun. January 2001) in an ex-tremely wet area of highly irregular terrain(Albuja-V., 1989).

IMBABURA

15. Los Cedros [08189N, 788479W; IGM, 1973;Young-Owl et al., 1992], 1400–1450 m.Collected by M. Bronsvoort in August1993. Based only on adult fluid-preservedspecimen with small hind foot; not con-firmed with cranial material. State of forestat time of collection unknown, but generalregion holds much intact forest (C. Carri-on, personal commun. January 2001).

PICHINCHA

16. Salcedo Lindo [08139N, 798059W; circa 450m; IGM, 1991]. Collected by R.E. Blei-weiss, F.A. Iwen, and F. Sornoza-Molina inNovember 1991. ‘‘Trapped in forest; densevegetation and downed logs’’ (R.E. Blei-weiss field notes, November 1991); at thebase of a tree in primary forest (F. Sornoza-Molina, in litt. October 2001).

Heteromys teleus

ECUADORCOTOPAXI

17. Bosque Integral Otonga [08259S, 798009W;Freiberg, 1998 and Jarrın-V., 2001; notLynch and Duellman, 1997 or Tirira-S.,1999], 1900–2000 m. Collected by P. Jar-rın-V. in September 1996. Small reserve(1000 ha) of mainly primary premontaneand montane wet tropical forest on thewestern slopes of the Andes, adjacent tolarger protected areas (Jarrın-V., 2001).Canopy not completely closed; with ‘‘adense shrubby and herbaceous ground flo-ra’’ (Freiberg, 1998: 168). The Heteromyswas caught at ground level in primarycloud forest, in an area with highly irreg-ular terrain. See Moret (1991), Schatti andKramer (1991, 1993), Coloma (1995), Ki-zirian (1996), Lynch and Duellman (1997),


and Jarrın-V. (2001) for other faunal re-ports.

ESMERALDAS

18. Quininde [08209N, 798289W; 100 m; Al-buja-V. and Mena-V., 1987; Albuja-V.,1999]. Collected by R. Rageot in October1980. Based only on adult study skin withlarge hind foot; not confirmed by cranialmaterial. Most of the region has been con-verted to cattle pastures and plantations ofoil palms and bananas, but with remnantdisturbed forests still present at least in1971 (Albuja-V. and Mena-V., 1987; Lynchand Duellman, 1997). Habitat conditions atsite of capture not available.

GUAYAS

19. Cerro Manglar Alto, eastern slope, [18469S,808389W; IGM, 1989; G.H.H. Tate fieldnotes, May 1923; not Voss, 1992 or Musseret al., 1998, who followed the coordinatesgiven for the town of Manglar Alto byPaynter and Traylor, 1977 and Paynter,1993, respectively], specimens from 1400ft [427 m]. Collected by G.H.H. Tate inMay 1923. See site 20 for general site in-formation. A large cornfield and severalsmall pastures lay in the immediate vicinityof this camp; the surrounding area was‘‘dense, fairly dry, tagua [Phytelephas ae-quatorialis, a palm] jungle.’’ Specimenstaken in stream bank running through for-est and in nearby cornfield.

20. Cerro Manglar Alto, western slope [18479S,808379W; IGM, 1989; G.H.H. Tate fieldnotes, May 1923; not Voss, 1992 or Musseret al., 1998, who followed the coordinatesgiven for the town of Manglar Alto byPaynter and Traylor, 1977 and Paynter,1993, respectively], specimens from 2000ft [610 m]. Collected by G.H.H. Tate inMay–June 1923. Cerro Manglar Alto is thename used by Tate to refer to the portionof the Cordillera de Chongon-Colonche(southern extension of the Cordillera de laCosta) at the headwaters of the Rıo Mang-lar Alto (5 Rıo Culebra) immediately eastof the coastal town Manglar Alto. The lowpeaks in this region receive various nameson detailed maps (IGM, 1988, 1989), butnone of them is known as Cerro ManglarAlto. Nevertheless, consultation of Tate’sfield notes and a detailed, modern map(IGM, 1989) allows for a fairly precise lo-calization of his collecting sites and tenta-tive assignment of coordinates, which we

judge to be accurate to within a minute.Both of Tate’s camps apparently were lo-cated between the Cerro Las Mesas and theLoma Colonche, and near the crest of thecordillera (IGM, 1989). A general charac-terization of the region follows: ‘‘Dense,wet, tagua [Phytelephas aequatorialis, apalm] forest, which continues up to 1600or 2000 ft [488–610 m] in the most west-erly ridges of the range. Moisture [is]caught by these spurs leaving the rest ofthe hillcrest further east practically dry al-though just about as high’’ (G.H.H. Tatefield notes, May 1923). Specimens fromthis camp on the western slopes were takenalong the Rıo Domene (not found on ourmaps), a small tributary of the Rıo ManglarAlto/Rıo Culebra, in undisturbed forest.There, Heteromys was ‘‘only taken in trapsplaced in runways which passed immedi-ately below the overhang of the bank in itssteepest parts’’ (G.H.H. Tate field notes,May 1923).

LOS RıOS

21. Rıo Palenque Science Center [08349S798209W; Lynch and Duellman, 1997; seealso Best and Kessler, 1995 and Albuja-V.,1999], 220 m. Collected by A.T. Smith inMarch 1977 and by D.E. Wilson in Feb-ruary 1979. Found in primary forest (A.T.Smith, in litt. August 2001); habitat infor-mation regarding the specimen collectedby Wilson unknown. Now-isolated forestreserve containing both primary (87 ha)and secondary (80 ha) habitats (Paynter,1993; see also Dodson and Gentry, 1978).

22. Jauneche (Estacion Biologica Pedro FrancoDavila) [18209S 798359W; Albuja-V., 1992,1999], 50 m. Collected by L. Albuja-V. inJuly 1991. Found in disturbed forest (L.Albuja-V., personal commun. January2001). Reserve encompasses 138 ha (Bestand Kessler, 1995); see also Dodson et al.(1985) and Foster (1992a) for botanicalcharacterizations.

PICHINCHA

23. Bosque Protector La Perla [08009N,798239W; Lynch and Duellman, 1997; notTirira-S., 1999], 220 m. Collected by F.Campos-Y. in May 1990. Based only onadult study skin with large hind foot; notconfirmed by cranial material. This reservecharacterized as a ‘‘remnant of humid trop-ical rainforest’’ (Lynch and Duellman,1997) surrounded by agricultural areas.


The Heteromys was captured in a dis-turbed, seasonally flooded forest (250 ha inarea at the time) dominated by various spe-cies of palms (especially Iriartea delto-idea), with many downed trunks present.The forest has been selectively logged; thispractice largely ceased with the creation ofthe reserve, but cattle continued to enterfrom the surrounding pastures in 1990 (F.Campos-Y., in litt. August 2001).

APPENDIX 2: Specimens Examined

Heteromys anomalus (25): COLOMBIA (13):CUNDINAMARCA: Caparrapı, Volcanes, ICN 409,1827–1831, 1832 (holotype of Heteromys anom-alus hershkovitzi), 1833–1835, 1866. MAGDALE-NA: below Minca, 1000 ft [305 m], AMNH 15347(holotype of Heteromys jesupi), 15348. TRINI-DAD AND TOBAGO (6): TRINIDAD: Caura,AMNH 7567/5960, 7575/5967, 7578/5970, 7588/5979, 7589/5980, 7594/5985. VENEZUELA (6):ZULIA: Rıo Aurare, FMNH 18622, 18623 (holo-type of Heteromys anomalus brachialis), 18624–18627. See also 64 other specimens from Vene-zuela reported in Anderson and Soriano (1999)and 201 from Colombia in Anderson (1999).

Heteromys australis (59): COLOMBIA (17):NARINO: Barbacoas, AMNH 34178–34180,34182, 34184–34185; Buenavista, 1200 ft [366m], AMNH 34181, 34183, 34186–34188; Muni-cipio Barbacoas, Corregimiento Altaquer, circa 7km NNE Altaquer, Reserva Natural del Rıo Nam-bı (FELCA), 1300 m, ICN 13659; Junın, Planadade Maindes, 870 m, UV 3067–3069, 3040A. VAL-LE DEL CAUCA: Las Lomitas, 5000 ft [1524 m],AMNH 32240 (holotype of Heteromys lomiten-sis). ECUADOR (27): CARCHI: Parroquia TobarDonoso, El Pailon, 1400 m, EPN 87.1810,87.1811; Parroquia Tobar Donoso, El Pailon, jun-to al rıo Pailon, 1400 m, EPN 87.1746; ParroquiaTobar Donoso, El Pailon, Loma del Tigre, margenizquierda rıo Pailon, 1400 m, EPN 87.1786,87.1787. ESMERALDAS: Canton San Lorenzo, Ma-taje, EPN 91.3174; Bulim [5 Pulun], 160 ft [49m], FMNH 18871; USNM 172940; San Javier [5San Javier de Cachavı], 60–120 ft [18–37 m],USNM 113304–13307; Alto Tambo, vıa Ibarra-Lita-San Lorenzo, Reserva Ecologica Cotacachi-Cayapas, QCAZ 840; Playa de Oro [5 Luis Var-gas Torres], Estero Zapote, MECN uncataloged,ear tag J. Cevallos 688; Alto Tambo, 2 km S of,700 m, ROM 105784; Reserva La Mayronga, 80m, MECN 1611, 1630, 1645; Reserva La May-ronga, Lagarto, 100 m, QCAZ 932–933; Luis Var-gas Torres, 2 km S de, EPN 84.561, 84.579; LuisVargas Torres, 4 km S de, EPN 84.581; Alto Tam-

bo, Hacienda La Granada, Lita, 3 km de, vıa aSan Lorenzo, 670 m, EPN 84.198, 84.199. IM-BABURA: Cotocachi, Manduriacu, Los Cedros,1400–1450 m, EPN 93.3971. PICHINCHA: Coop-erativa Salcedo Lindo, road from Pedro VicenteMaldonado to campamento de CODESA, 4 km Sof CODESA camp, UWZM S.31291. PANAMA(15): DARIEN: Cana, 1800–2000 ft [549–610 m],USNM 178621, 178698, 178699 (holotype ofHeteromys australis consicus), 178700, 179595;Amagal, 1000–2000 ft [305–610 m], ANSP19491–19498, 19499 (holotype of Heteromys aus-tralis pacificus), 19779. Plus see Anderson (1999)for 134 additional specimens of H. australis fromColombia, Panama, and Venezuela. Note that Al-berico et al. (2000) miscited Anderson (1999) byfalsely mentioning the documented presence of H.australis from the Departamento del Cauca insouthwestern Colombia, while failing to note ver-ified specimens from the departments of Antio-quia, Boyaca, Caldas, Cordoba, Cundinamarca,Huila, and Quindıo or to give credit to reportedspecimens of this species in the Museo La Salle(MLS, Bogota) and MCZ collections.

Heteromys desmarestianus (species complex;64): COSTA RICA (11): CARTAGO: Angostura,USNM 12903/38590, 12904/38591 (holotype ofHeteromys desmarestianus subaffinis), 12905/36848; SAN JOSE: Escazu, 3000 ft [914 m],AMNH 131729 (holotype of Heteromys desma-restianus underwoodi); Escazu, Los Higuerones,AMNH 137299–137302; San Geronimo Pirris,USNM 250348 (holotype of Heteromys desma-restianus planifrons), 250349, 256445. MEXICO(10): CHIAPAS: Chicharras, USNM 77516 (holo-type of Heteromys goldmani), 77577, 77579–77582, 77694; VERACRUZ: Motzorongo, USNM63718, 63719 (holotype of Heteromys temporal-is), 63720. PANAMA (43) COLON: Rıo Indio,near Gatun, USNM 170919–170920, 170975,170976 (holotype of Heteromys zonalis); CHIRI-QUı: Boquete, 4000–5800 ft [1219–1768 m],MCZ B10355, B10356 (holotype of Heteromysrepens), B10358, B10360–B10361; Cerro Pando,ANSP 17835 (holotype of Heteromys desmares-tianus chiriquensis), 17836–17837, 18374–18377; DARIEN: Mount Pirri, near head of Rıo Li-mon, 4500–5200 ft [1372–1585 m], USNM178998–179004, 179006–179015, 179016 (holo-type of Heteromys crassirostris), 179018–179020; PANAMA: Cerro Azul, 2500–3000 ft[762–914 m], USNM 171107 (holotype of Het-eromys panamensis), 171108–171111, 171128.

Heteromys gaumeri (2): MEXICO (2): CAM-PECHE: 7 km N, 51 km E Escarcega, KU 93644;103 km SE Escarcega, KU 93647.

Heteromys nelsoni (2): MEXICO (2): CHIA-


PAS: Pinabete, USNM 77920 (holotype of Heter-omys nelsoni), 77578.

Heteromys oresterus (9): COSTA RICA (9):SAN JOSE: El Copey de Dota, 6000–6500 ft[1829–1981 m], UMMZ 64026, 64027 (holotypeof Heteromys oresterus), 64028–64034.

Heteromys teleus (31): ECUADOR (31): CO-TOPAXI: San Francisco de las Pampas, ReservaBosque Nublado Otonga [5 Bosque IntegralOtonga], 1900–2000 m, QCAZ 1788. ESMERAL-DAS: Quininde, EPN .22 (provisional identificationbased on study skin). GUAYAS: Cerro ManglarAlto, eastern slope, Camp I, 1200 ft [366 m](camp), [specimens from 1400 feet, 5 427 m;

G.H.H. Tate field notes, May 1923], AMNH64675–64679; Cerro Manglar Alto, western slope,Camp II, 1500 ft [457 m] (camp), [specimensfrom 2000 feet, 5 610 m; G.H.H. Tate field notes,May 1923], AMNH 64680–64696. LOS RıOS: RıoPalenque Biological Station [5 Rıo Palenque Sci-ence Center], 220 m, KU 149132–149135; RıoPalenque Science Center, Santo Domingo, 47 kmS of (by road), USNM 528573; Canton Vinces,Parroquia Palenque, Jauneche, Estacion BiologicaPedro Franco Davila, 50 m, EPN 91.3038. PI-CHINCHA: La Concordia, 2 km de, Bosque Protec-tor La Perla, 220 m, QCAZ 905 (provisional iden-tification based on study skin).

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a This paper meets the requirements of ANSI/NISO Z39.48-1992 (Permanence of Paper).

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