PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, NY 10024 Number 3373, 42 pp., 19 ®gures, 10 tables June 21, 2002

A New for Aepeomys fuscatus Allen, 1912, and intectus Thomas, 1921: Enigmatic Murid from Andean Cloud Forests

ROBERT S. VOSS,1 MARCELA GOÂ MEZ-LAVERDE,2 AND VICTOR PACHECO3

CONTENTS Abstract ...... 2 Resumen ...... 2 Introduction ...... 2 Materials and Methods ...... 3 Systematics ...... 4 Taxonomic History ...... 4 Handleyomys, new genus ...... 5 Geographic Variation and Species Limits ...... 20 Handleyomys fuscatus (Allen) ...... 23 Handleyomys intectus (Thomas) ...... 24 Natural History and Biogeography ...... 25 Habitat Descriptions ...... 27 Summary ...... 33 Discussion ...... 35 Acknowledgments ...... 37 References ...... 37 Appendix: Gazetteer ...... 40

1 Associate Curator, Division of Vertebrate (Mammalogy), American Museum of Natural History. e-mail: [email protected] 2 FundacioÂn UlamaÂ, Apartado AeÂreo 93674, Santafe de BogotaÂ, Colombia. e-mail: [email protected] 3 Curador de MamõÂferos, Museo de Historia Natural, Universidad Nacional Mayor de San Marcos, Apartado 14- 0434, Lima 14, Peru. e-mail: [email protected]

Copyright ᭧ American Museum of Natural History 2002 ISSN 0003-0082 2 AMERICAN MUSEUM NOVITATES NO. 3373

ABSTRACT Two nominal species of Neotropical murid rodents (subfamily ) that have long been referred to different genera are here placed in a new genus in recognition of their distinctness from other named supraspeci®c taxa. Aepeomys fuscatus Allen and Oryzomys intectus Thomas share a unique combination of external and craniodental character states that diagnose Handleyomys, new genus, with fuscatus as its type species. Morphological compar- isons of Handleyomys with the type species of Aepeomys Thomas and Oryzomys Baird provide a basis for preliminary inferences about phylogenetic relationships. Five shared, derived char- acter states support the hypothesis that Handleyomys is an oryzomyine, but no close relation- ship between the new genus and any particular oryzomyine clade is indicated by the data at hand. All known specimens of Handleyomys are from the western Andes (Cordillera Occi- dental) and the central Andes (Cordillera Central) of Colombia, where they have been collected at 20 localities ranging in elevation from 1500 to 2800 m above sea level. Analyses of mor- phological data suggest that two valid allopatric species are represented, of which H. fuscatus is endemic to the western Andes and H. intectus to the central Andes. Although no other mammalian clade is known to have the same geographic distribution, recent analyses of am- phibian biogeography in Colombia suggest that Handleyomys is part of a nonvolant cloud- forest vertebrate fauna with allopatric sister taxa in the Cordillera Occidental and Cordillera Central. Much revisionary taxonomic research, however, is needed to assess the generality of this pattern of endemism among other cloud-forest .

RESUMEN Dos especies nominales de roedores muÂridos neotropicales (subfamilia Sigmodontinae) que habõÂan sido ubicadas durante mucho tiempo en geÂneros diferentes, se asignan a un nuevo geÂnero reconociendo sus diferencias con otros taxones nominales supraespecõ®cos. Aepeomys fuscatus Allen y Oryzomys intectus Thomas comparten una combinacioÂn uÂnica de estados de caracteres externos y craneodentales, que permiten diagnosticar a Handleyomys como un geÂ- nero nuevo, con fuscatus como su especie tõÂpica. Las comparaciones morfoloÂgicas entre Han- dleyomys y las especies tõÂpicas de los geÂneros Aepeomys Thomas y Oryzomys Baird proveen una base para hacer deducciones preliminares sobre sus relaciones ®logeneÂticas. La hipoÂtesis de que Handleyomys es un orizomino esta apoyada por cinco estados de caracteres compartidos derivados, pero los datos disponibles no indican una relacioÂn estrecha entre el nuevo geÂnero y ninguÂn clado particular de orizomino. Todos los espeÂcimenes conocidos de Handleyomys provienen de de las Cordilleras Occidental y Central de los Andes de Colombia, donde fueron recolectados en 20 localidades en alturas comprendidas entre 1500 y 2800 m sobre el nivel del mar. El anaÂlisis de los datos morfoloÂgicos sugiere la presencia de dos especies alopaÂtricas vaÂlidas, de las cuales H. fuscatus es endeÂmica de la Cordillera Occidental y H. intectus de la Cordillera Central. Aunque no se conoce ninguÂn otro clado de mamõÂferos con una distribucioÂn geogra®ca similar, anaÂlisis biogeogra®cos recientes de an®bios de Colombia sugieren que Handleyomys forma parte de una fauna vertebrada no voladora de los bosques de niebla, con taxones alopaÂtricos hermanos en la Cordillera Occidental y en la Cordillera Central. Para evaluar la generalidad de este patroÂn de endemismo entre otros mamõÂferos de los bosques nublados se requiere de mayores investigaciones taxonoÂmicas.

INTRODUCTION tane rain forests in the southern Andes by treeless alpine deserts and grasslands, the Extending from the SerranõÂa de Perija west cloud forests of the northern and central An- of Lake Maracaibo southward to the yungas des harbor taxonomically distinctive mam- of central Bolivia is the largest region of malian communities that are among the least tropical montane rain forest in the world. Iso- known of all terrestrial vertebrate faunas. Al- lated from similar habitats in Central Amer- though a few large species are relatively well ica by the RõÂo Atrato lowlands of north- known as focal taxa for international conser- western Colombia, and from temperate mon- vation effortsÐwoolly tapirs (Tapirus pin- 2002 VOSS ET AL.: NEW GENUS OF ANDEAN RODENTS 3 chaque), spectacled bears (Tremarctos or- computed head-and-body length (HBL) by natus), pudus (Pudu mephistopheles), and subtracting LT from TL. Length of the hind yellow-tailed woolly monkeys (Lagothrix foot (HF, including claws) and of the external ¯avicauda)Ðthe overwhelming majority of ear (Ear, from notch to distalmost margin of Andean cloud-forest mammals are small and the pinna) were also transcribed from speci- virtually unstudied. Indeed, most museum men tags; however, we usually remeasured collections of mammals from Andean cloud hindfoot length on dried skins to check the forests remain incompletely or inaccurately accuracy of values recorded by the collector, identi®ed, with the result that patterns of spe- and we used our values whenever large dis- cies richness and endemism in this fauna are crepancies were found. All external measure- effectively unknown. ments are reported to the nearest millimeter This report contributes to the analysis of (mm). mammalian diversity in the Andean cloud- Craniodental measurements were taken forest biota by diagnosing a previously un- with digital calipers and recorded to the near- recognized clade of murid rodents from the est 0.01 mm, but values reported herein are western and central cordilleras of Colombia. Two species, both described early in the 20th rounded to the nearest 0.1 mm. The follow- century and long referred to different genera, ing were measured as illustrated and de®ned are here placed in a new genus because of by Voss (1988, 1991): CIL, condylo-incisive their uniquely shared combination of mor- length; LD, length of diastema; LM, occlusal phological characters. By doing so, we sim- length of the maxillary molar row; BM1, plify the taxonomic context for two major breadth of the ®rst maxillary molar; LIF, phylogenetic studies currently in progress, length of one incisive foramen; BIF, breadth and we draw attention to a distinctive lineage across both incisive foramina; BPB, breadth whose restricted ecogeographic distribution of the palatal bridge (between the M1 pro- provides evidence for patterns of endemism tocones); BZP, breadth of the zygomatic that may be shared with other cloud-forest plate; LIB, least interorbital breadth; ZB, zy- organisms. gomatic breadth. In addition, we measured nasal length (NL) as the greatest longitudinal MATERIALS AND METHODS dimension of either the right or left nasal bone, and interparietal breadth (IPB) as the The material we examined consists of greatest transverse dimension of the interpa- standard skin-and- preparations or ¯uid- rietal bone. preserved specimens deposited in the follow- Except as noted below, all analyzed char- ing institutional collections: AMNH, Amer- acter data were obtained from nonsenescent ican Museum of Natural History (New adults as determined by dental and pelage York); BMNH, Natural History Museum criteria. A specimen was judged to be adult (formerly the British Museum of Natural if the molar dentition was fully erupted (with History, London); FMNH, Field Museum of M3 exhibiting at least some wear) and if the Natural History (Chicago); ICN, Museo de pelage was mature; old adults (with molars Historia Natural del Instituto de Ciencias Na- turales de la Universidad Nacional de Co- worn below the widest part of the crown) lombia (BogotaÂ); IND-M, Instituto Alexan- were not measured or scored for qualitative der von Humboldt (formerly INDERENA, characters. In effect, the specimens used for Villa de Leiva); MBUCV, Museo de Biolo- our statistical analyses approximately corre- gõÂa, Universidad Central de (Ca- spond to toothwear classes 2±4 as de®ned by racas); MLS, Museo del Instituto La Salle Voss (1991). (BogotaÂ); UMMZ, Qualitative character variation is described Museum of Zoology (Ann Arbor); USNM, herein using anatomical terminology that is National Museum of Natural History (Wash- explained or referenced by Reig (1977), Voss ington, D.C.). (1988), Carleton and Musser (1989), Voss We transcribed total length (TL) and (1993), Voss and Carleton (1993), and Step- length of tail (LT) from specimen tags and pan (1995). 4 AMERICAN MUSEUM NOVITATES NO. 3373

SYSTEMATICS neric assignment at least made biogeographic sense. However, with all of the known ma- TAXONOMIC HISTORY terial of fuscatus in New York, and the types The ®rst species treated in this report was of lugens and vulcani in London, no other originally described as Aepeomys fuscatus by taxonomic assessment was possible given the Allen (1912) on the basis of 12 specimens incompleteness of Allen's description. collected by W. B. Richardson near the vil- Although Aepeomys was recognized as a lage of San Antonio in the western Andes of valid genus by Gyldenstolpe (1932) and Tate Colombia. An ornithologist by training, Al- (1932b), Osgood (1933) argued that the type len focused his description on pelage texture, species (lugens) was insuf®ciently divergent pigmentation, and other features of the pre- from the many forms then referred to Tho- served skins he examined, while ignoring all masomys Coues, 1884, to maintain this usage qualitative craniodental characters (p. 89): and concluded that Aepeomys was just a ju- nior synonym. Osgood's judgment was sub- Pelage long, thick, soft and velvety. Upperparts blackish, almost clear black over the median dorsal sequently endorsed by Ellerman (1941), who region, with a faint wash of grayish brown over the recognized Thomasomys fuscatus, T. lugens, shoulders and on the sides of the body, almost im- and T. vulcani as valid species in his in¯u- perceptible except in favorable lights, when the ex- ential monograph on classi®cation. treme tips of the hairs are seen to be grayish bistre; underparts dark gray, the pelage being slaty with the Cabrera (1961) likewise followed Osgood's extreme tips of the hairs lighter or pale drab-gray; inclusive sense of Thomasomys, but treated ears brown, thickly clothed with soft black hairs on all of the taxa formerly referred to Aepeomys the basal third externally, the rest nearly naked on as conspeci®c, listing fuscatus and vulcani as both surfaces; feet ¯esh color, very thinly haired; tail subspecies of T. lugens. light gray brown, nearly unicolor, naked. Apparently, neither Osgood, Ellerman, nor From this description and accompanying Cabrera actually examined specimens of fus- measurements, all that a reader can infer was catus, and no new character information that Allen's new species was a small, soft- about this taxon was subsequently published furred, dark-colored mouse with a naked tail until Gardner and Patton's (1976) review of approximately the same length as its head karyotypic variation among Neotropical mu- and body. Unfortunately, these traits are rids. In that report, the authors analyzed shared by many species of cloud-forest mu- chromosomal preparations from fuscatus and rids, few of which can be reliably distin- six other species of Thomasomys (sensu Os- guished by the additional details that Allen good) and remarked that Allen's form was provided. The omission of qualitative cran- highly distinctive. They summarized their iodental character information, already wide- conclusions in the context of suprageneric ly recognized as important for muroid rodent (tribal-level) assemblages as then understood systematics (e.g., by Thomas, 1906; Osgood, by taxonomists (p. 32): 1909; Goldman, 1910), was particularly un- The thomasomyine group . . . includes Thomasomys, fortunate. In justifying his generic assign- Rhipidomys, Nyctomys, Otonyctomys, and Phaeno- ment, Allen (1912: 89) stated only that fus- mys. Members of this group represented by our ma- catus ``. . . is evidently related to Aepeomys terial . . . include Rhipidomys latimanus and at least vulcani Thomas, from which it appears to six species of Thomasomys (sensu lato). Aepeomys (ϭ differ in relatively longer tail and somewhat Thomasomys) fuscatus (2n ϭ 54, FN ϭ 62) is kar- yotypically the most aberrant representative and ac- in coloration.'' tually may not belong in the Rhipidomys and Tho- The genus Aepeomys was originally masomys complex. Therefore, we are using the ge- named by Thomas (1898) to contain two neric name Aepeomys for this form. Chromosomally taxa, of which the designated type species it falls within the ®rst oryzomyine group as exempli- was A. lugens (Thomas, 1896) from Vene- ®ed by Oryzomys palustris, a group sharing chro- mosomal characters with various other groups . . . zuela. Because the second referred taxon, A. vulcani Thomas (1898), was based on a spec- Most late 20th-century checklists (e.g., Hon- imen from Ecuador, the known range of Ae- acki et al., 1982; Alberico, 1983; Cuervo- peomys (sensu Thomas) bracketed the type DõÂaz et al., 1986; Musser and Carleton, 1993; locality of fuscatus, and Allen's (1912) ge- Alberico et al., 2000) have followed Gardner 2002 VOSS ET AL.: NEW GENUS OF ANDEAN RODENTS 5 and Patton (1976) in ranking Aepeomys as a from the 1970s, when the late Charles O. valid genus, but the implicit hypothesis that Handley, Jr. (then curator of mammals at the its member species (fuscatus, lugens, and National Museum of Natural History) and vulcani) form a natural (monophyletic) group RSV (then a graduate student at the Univer- remains effectively untested by any morpho- sity of Michigan) independently examined logical, karyotypic, or biochemical data. the type series of Aepeomys fuscatus and The second species treated in this report concluded that it had been misclassi®ed by was originally described as Oryzomys intec- Allen (1912) and by all subsequent compilers tus by Thomas (1921) on the basis of three of Neotropical murid checklists. Whereas A. specimens collected by NiceÂforo MarõÂa near lugens (the type species of Aepeomys) was Santa Elena in the central Andes of Colom- clearly allied with Thomasomys and certain bia. Unlike Allen's description of fuscatus, other ``thomasomyines'', fuscatus exhibited Thomas's description of intectus included equally unambiguous similarities to oryzo- several relevant details of craniodental mor- myines. Rather than belonging to Oryzomys phology (p. 356): or other allied genera, however, fuscatus seemed to be a highly divergent form. Han- Skull peculiarly short, broad, and rounded, with broad interorbital region. Indeed, it is almost precisely like dley and RSV compared notes and agreed to that of a , with the important exception collaborate on a joint report, but other pro- that there is no trace of the supraorbital beading so jects intervened and only a few colleagues conspicuous in that group. Brain-case similarly low, were informed about their results. In the smooth, and without ridges. Palatal foramina short, meantime, Guy G. Musser examined the type about the length of the tooth-row. Molars stout and heavy, large for the size of the , their structure series of Oryzomys intectus in London, rec- more like that in Melanomys than in the smaller spe- ognized that it represented the same clade cies of Oryzomys, but many of the larger species of discovered by Handley and RSV, and gen- Oryzomys also have quite similar molars. erously provided them with copies of his From these and other comparisons, Thomas notes; RSV followed this up by examining concluded that (p. 357): intectus himself on a subsequent visit to the Natural History Museum. This is a remarkably distinct species, whose system- Other independent discoveries of essen- atic position is not at present easy to determine. Its tially the same facts about fuscatus and in- peculiarly broad low skull distinguishes it from any tectus were made by MGL and VP; their re- Oryzomys known to me, while the entire absence of supraorbital ridges separates it from Melanomys,to sults were based on new specimens and/or which its short tail and the general shape of the skull new characters (not previously considered by perhaps indicate some af®nity. Many Oryzomys, how- Handley and RSV), resulting in this collab- ever, have no supraorbital ridges, and I therefore pro- oration. With the recent death of Charles visionally place it in that genus. Handley, it seems appropriate to name the As the heterogeneous contents of Oryzo- new genus for him in recognition of his past mys were gradually sorted into species participation in our work and in tribute to his groups and subgenera by later generations of many important contributions over the course mammalogists, intectus remained an enigma, of a long and productive career in Neotrop- left unallocated by some authors (Ellerman, ical mammalogy (®g. 1). 1941) or referred to the nominate subgenus by default (Tate, 1932a; Cabrera, 1961). De- Handleyomys, new genus spite recent advances in oryzomyine system- Figures 2±11 atics that have resulted in several former sub- genera of Oryzomys being elevated to gener- TYPE SPECIES: Aepeomys fuscatus Allen, ic rank (e.g., , , 1912. ; see Carleton and Musser [1989], GEOGRAPHIC DISTRIBUTION: Known from Musser and Carleton [1993]), the classi®ca- the western and central Andean cordilleras of tion of intectus has not been addressed by Colombia at elevations from ca. 1500 to any published analysis or discussion of char- 2800 m above sea level (®g. 12). acter data. CONTENTS: Two species as redescribed be- The chronology of the present study dates low. 6 AMERICAN MUSEUM NOVITATES NO. 3373

Fig. 1. Charles O. Handley, Jr. (right) with Carl Johnson (Director of the Gorgas Memorial Labo- ratory, left) at their camp on the RõÂo Tacarcuna, Provincia DarieÂn, Panama, in 1964. Photograph courtesy of the Smithsonian Institution Archives (Alexander Wetmore Papers, box 188: photograph 8793).

MORPHOLOGICAL DIAGNOSIS AND DESCRIP- (two carpal and three interdigital pads); TION: Adult body pelage ®ne and soft, uni- claws short, neither conspicuously elongated formly dull brownish-gray dorsally, usually nor unusually recurved. Pes long and narrow, darker (sometimes almost blackish) middor- with outer digits (I and V) much shorter than sally than on ¯anks; ventral pelage dark gray middle three (claw of dI extending to middle frosted with paler gray or buff, not sharply of ®rst phalange of dII, claw of dV extending countershaded. Mystacial, superciliary, ge- just beyond ®rst interphalangeal joint of nal, submental, interramal, and carpal vibris- dIV); conspicuous ungual tufts of long sil- sae present; mystacial hairs neither very very hairs rooted at bases of claws on dII± short nor very long, extending posteriorly to dV, but pedal dorsum otherwise only sparse- (but not beyond) caudal margins of pinnae ly covered with short pale or dark-banded when laid back against cheeks of properly hairs; plantar surface (including heel) naked, made-up skins. Pinnae not large but clearly weakly pigmented (grayish in life), with two visible above fur of head, sparsely covered metatarsal and four interdigital pads; indis- with short dark hairs. Manus sparsely cov- tinct squamae (scale-like tubercles) sparsely ered dorsally with short pale (whitish or sil- distributed along outer distal plantar surfac- very) hairs; ventral surface naked and unpig- es, but not in center of sole. Tail about as mented, with ®ve separate plantar tubercles long as combined length of head and body, 2002 VOSS ET AL.: NEW GENUS OF ANDEAN RODENTS 7

Fig. 2. Above: Handleyomys fuscatus (ICN 12825) from El Campamento, Municipio Santuario, Departamento Risaralda (appendix: locality 14). Below: Pastures and disturbed primary cloud forest at about 2500 m elevation near El Campamento; specimens of H. fuscatus and six other species of murid rodents were captured in the forest at this locality (see text). Both photographs were taken by Marcela Morales during the rainy season (November) of 1991. 8 AMERICAN MUSEUM NOVITATES NO. 3373

apparently naked (a sparse caudal pelage is only visible under magni®cation), and uni- colored (dark above and below). Mammae six in inguinal, abdominal, and postaxial pairs. Skull with long, tapering rostrum ¯anked by shallow but distinct zygomatic notches; interorbital region hourglass-shaped, neither greatly in¯ated nor unusually constricted, with rounded supraorbital margins; braincase moderately in¯ated and rounded, without prominent temporal crests, ridges, or beads. (in lateral view) moderately broad, its anterior edge vertical or nearly so (not sloping backward), with a rounded (nev- er angular or spinous) anterodorsal contour. Premaxillae short (not produced anteriorly beyond incisors to form a rostral tube with nasals). Incisive foramina neither very short nor greatly elongated, averaging about 60% of diastemal length (not extending posteri- orly between molar rows) and widest near premaxillary/maxillary sutures. Palatal bridge wide and long, without median ridge or deep lateral gutters; posterolateral pits usually large and often complex, typically consisting of two or more foramina recessed in a common fossa on each side. Mesopter- ygoid fossa not penetrating anteriorly be- tween molar rows; bony roof of fossa com- plete or perforated only by narrow slits, nev- er conspicuously fenestrated. Alisphenoid strut absent (buccinator-masticatory foramen and foramen ovale con¯uent). Carotid arte- rial morphology primitive, with orbitofacial circulation supplied by separate supraorbital and infraorbital branches of large stapedial artery (ϭ pattern 1 of Voss, 1988); course of supraorbital stapedial ramus marked by prominent squamosal-alisphenoid groove Fig. 3. Plantar view of left hind foot of Han- dleyomys intectus (ICN 16092). Distinctive charac- and sphenofrontal foramen. Postglenoid fo- teristics of the genus include the narrow, hairless ramen separated from large subsquamosal fe- sole; six small plantar pads including thenar (t), hy- nestra by slender hamular process of squa- pothenar (h), and four interdigital (1±4) tubercles; mosal. Tegmen tympani not overlapping very short hallux (I); long and subequal second (II), squamosal, or tegmen tympani-squamosal third (III), and fourth (IV) digits; short and non- overlap weak (not involving a distinct pos- opposable ®fth digit (V); indistinct plantar squamae terior suspensory process of the latter bone). (sq); and ungual tufts of long hairs rooted at the Bullae small; pars ¯accida of tympanic mem- bases of the claws of digits II±V. brane present, large; orbicular apophysis of malleus well developed. Mandible with well-developed falciform coronoid process; lower incisor alveolus without distinct capsular process on lateral 2002 VOSS ET AL.: NEW GENUS OF ANDEAN RODENTS 9

Fig. 4. Dorsal and ventral cranial views (ca. ϫ2) of Handleyomys, Aepeomys and Oryzomys. From left to right: H. fuscatus (ICN 12703), H. intectus (ICN 16092), A. lugens (AMNH 143670), O. palustris (AMNH 242524). mandibular surface. Basihyal morphology pairs, bunodont when unworn but quickly unknown. eroded with age to same level as other enam- Incisors ungrooved, with yellow-orange elled structures (occlusal surface of most enamel bands; upper teeth small, narrow, adult teeth more-or-less planar); labial and deeper than wide, opisthodont; lower teeth lingual reentrant folds long and interpene- unremarkable in coloration or morphology. trating (incipiently lophodont sensu Voss, Maxillary molar rows parallel; principal 1993); M1 anterocone entire, not divided cusps arranged in opposite labial/lingual into labial and lingual conules (anteromedian 10 AMERICAN MUSEUM NOVITATES NO. 3373

¯exus absent); antero¯exus very deep, ex- tending lingually beyond dental midline on M1 and M2; anterolophs and mesolophs large, fused with corresponding (antero- and meso-) styles on labial margins of M1 and M2; posterolophs distinct on M1 and M2, persisting with moderate to heavy wear; M3 subtriangular, smaller than more anterior teeth, with most of the same occlusal ele- ments but usually without distinct hypocone or posteroloph. First maxillary molar with one accessory labial root (four roots total); M2 and M3 three-rooted. Anteroconid of ®rst mandibular molar (m1) undivided by median ¯exid, fused with protolophid and/or anterolophid to enclose a persistent internal fold of uncertain homolo- gy (antero¯exid and/or proto¯exid); antero- lophid absent on m2 and m3; anterolabial cingulum absent or indistinct on m2, consis- tently absent on m3; mesolophids and pos- terolophids large and well developed on all mandibular teeth; ectolophids consistently absent. First mandibular molar with an ac- cessory labial root, occasionally also with an accessory lingual root (three to four roots to- tal); m2 and m3 each with two small anterior roots and one large posterior root (three roots total). Tuberculum of ®rst rib articulates with transverse processes of seventh cervical and ®rst thoracic vertebrae; second thoracic ver- tebra with greatly elongated neural spine; en- tepicondylar foramen of humerus absent. Thoracicolumbar vertebrae 19; sacrals 4; caudals 28±30, with or without hemal arch- es4; ribs 12. Stomach (in two dissected specimens of H. intectus) unilocular and hemiglandular, with- out any extension of glandular epithelium into corpus; bordering fold crosses lesser curvature slightly to right of incisura angu- laris (between that ¯exure and the pylorus); bordering fold crosses greater curvature op-

4 At least one complete hemal arch (for illustrations and discussion of this character, see Steppan, 1995: 48± 49) appears to be present in each of several undamaged caudal series of Handleyomys fuscatus that we exam- Fig. 5. Lateral cranial and mandibular views ined, where it occurs between the second and third (ICN (ca. ϫ2) of Handleyomys, Aepeomys, and Oryzo- 12793, 12795) or between the third and fourth vertebrae mys. From top to bottom: H. fuscatus (ICN (ICN 12725). However, only hemal processes (no arch) 12703), H. intectus (ICN 16092), A. lugens were observed in our single undamaged caudal series of (AMNH 143670), O. palustris (AMNH 242524). H. intectus (ICN 12164). 2002 VOSS ET AL.: NEW GENUS OF ANDEAN RODENTS 11

Fig. 6. Maxillary toothrows (ϫ20) of Handleyomys, Aepeomys, and Oryzomys. From left to right: Handleyomys fuscatus (USNM 507269), H. intectus (ICN 16074), Aepeomys lugens (MBUCV I-2793), and Oryzomys palustris (AMNH 234936). posite incisura angularis. Gall bladder absent Osgood, 1933), an assessment with which we (in two dissected specimens of H. intectus). completely agree. Aepeomys reigi, however, Phallic and other male reproductive charac- is a valid species distinguished from A. lu- ters undetermined. gens by subtle but consistent morphological KARYOTYPE: Gardner and Patton (1976: ta- characters and by its highly distinctive kar- ble 2) reported a diploid number (2n)of54 yotype (Ochoa et al., 2001). The fourth nom- chromosomes and a fundamental number inal taxon, vulcani, is based on a partially (FN) of 62 from three karyotyped specimens crushed holotype (BMNH 98.5.1.10) that ex- of Handleyomys fuscatus (USNM 507267± hibits none of the distinctive attributes shared 507269); both the X and Y chromosomes are by A. lugens and A. reigi; as independently submetacentrics (op. cit.). determined by C.O. Handley, Jr. (personal COMPARISONS WITH AEPEOMYS: As recog- commun.) and VP (in prep.), vulcani is re- nized prior to this report (e.g., by Musser and ferable to the genus Thomasomys (sensu Carleton, 1993; Ochoa et al., 2001), the ge- stricto), wherein its possible synonymy with nus Aepeomys contained four other nominal other nominal taxa remains to be determined. taxa in addition to fuscatus: lugens Thomas Thus restricted (excluding fuscatus and (1896), vulcani Thomas (1898), ottleyi An- vulcani), Aepeomys is a morphologically di- thony (1932), and reigi Ochoa et al. (2001). agnosable taxon that can be unambiguously Of these, ottleyi has long been considered a distinguished from Handleyomys despite a subjective junior synonym of A. lugens (see super®cial resemblance in external features. 12 AMERICAN MUSEUM NOVITATES NO. 3373

Fig. 7. Mandibular toothrows (ϫ20) of Handleyomys, Aepeomys, and Oryzomys. From left to right: Handleyomys fuscatus (USNM 507269), H. intectus (ICN 16074), Aepeomys lugens (MBUCV I-2793), and Oryzomys palustris (AMNH 234936).

(Both genera include drab-colored, soft- not problematic, taxonomic comparisons are furred mice with naked tails about as long as more usefully focused on their many salient heads-and-bodies; narrow hindfeet with six points of difference in skeletal and visceral plantar pads, short outer digits, and long un- comparisons. gual tufts; and six mammae.) Because ®eld In dorsal and lateral cranial views (®gs. 4, identi®cations of these allopatric5 taxa are 5, 8) the rostrum of Aepeomys appears long and tapering, with a prominent bony tube 5 To date, Aepeomys lugens and A. reigi are known only from the Venezuelan Andes, where the former has formed by the nasals and premaxillae that been collected at elevations ranging from 1990 to 3200 projects anterodorsally well beyond the in- m in the states of MeÂrida and TaÂchira, and the latter at cisors; the rostrum of Handleyomys appears 1600±3100 m in the states of Lara and Trujillo (Thomas, 1896; Anthony, 1932; Handley, 1976; Ochoa et al., short and blunt by comparison, and lacks any 2001). Reports of A. lugens from Colombia (Cuervo- trace of a nasal-premaxillary tube. Flanking DõÂaz et al., 1986; Alberico et al., 2000) appear to have the base of the rostrum on each side of the been based on misidenti®ed material; we have not ex- skull in Handleyomys is a shallow but well- amined any Colombian material referable to this species in the AMNH, BMNH, FMNH, ICN, IAvH, MLS, de®ned zygomatic notch formed by the free USNM, or other collections. anterodorsal margin of a broad, more-or-less 2002 VOSS ET AL.: NEW GENUS OF ANDEAN RODENTS 13

Fig. 8. Dorsal and lateral views of rostrum. A, Aepeomys lugens (AMNH 143670); B, Handleyomys fuscatus (ICN 12703); C, Oryzomys palustris (AMNH 242524). Abbreviations: nas, nasal; pre, pre- maxillary; zn, zygomatic notch; zp, zygomatic plate. 14 AMERICAN MUSEUM NOVITATES NO. 3373

vertically oriented zygomatic plate. By con- trast, the zygomatic notches are indistinct in Aepeomys because the zygomatic plate is slender and slopes posterodosally without a free anterodorsal projection. The interorbital region is hourglass-shaped (concave-sided in dorsal view) with rounded supraorbital mar- gins in both taxa (®g. 9A, B), but this part of the skull is relatively narrower in Handley- omys, a consequence of its less in¯ated fron- tal sinuses and olfactory bulb. The braincase is likewise more compact and globular in Handleyomys by comparison with the more in¯ated and ovoid calvarium of Aepeomys. In ventral cranial view, the two genera are most conspicuously distinguished by palatal morphology. Whereas the palate of Aepeo- mys is relatively short (not extending poste- riorly much behind the molar rows) and lacks well-developed posterolateral pits (®g. 10A), the palate of Handleyomys is long (extending posteriorly well behind the molar rows) and is perforated posterolaterally by one or more large pits between M3 and the mesoptery- goid fossa (®g. 10B). In addition, the wide mesopterygoid fossa of Aepeomys is ¯anked by long, narrow parapterygoid fossae, but the mesopterygoid fossa of Handleyomys is rel- atively narrower and ¯anked by shorter, broader parapterygoid fossae; in both genera, the mesopterygoid roof is entirely bony or is perforated only by small foramina. In Aepeomys, the bulla is ®rmly anchored to the rest of the skull anterodorsally by broad overlap between the tegmen tympani and a posterior suspensory process of the squamosal, and the postglenoid foramen (through which the eponymous vein exits the cranial lumen) is smaller than the subsqua- mosal fenestra (®g. 11A). By contrast, the bulla of Handleyomys is not ®rmly anchored to the skull anterodorsally because the squa- mosal lacks a posterior suspensory process; in consequence, the postglenoid foramen is much wider than the subsquamosal fenestra (®g. 11B). Fig. 9. Dorsal views of interorbital region. A, The upper molar dentitions of both genera Aepeomys lugens (AMNH 143670); B, Handle- are highly distinctive (®g. 6). The ®rst max- yomys fuscatus (ICN 12703); C, Oryzomys pal- illary molar of Aepeomys is relatively longer ustris (AMNH 242524). and narrower than the shorter, broader M1 of Handleyomys, and the unworn labial cusps (paracone and metacone) of M1±M2 in Ae- peomys are much taller and sharper than their 2002 VOSS ET AL.: NEW GENUS OF ANDEAN RODENTS 15 homologs in Handleyomys. With increasing toothwear, the principal molar cusps of Ae- peomys persist as distinct tubercles, whereas the molar cusps of Handleyomys are quickly worn down to form a common planar surface with other occlusal features. Differences in the length and orientation of the principal la- bial and lingual ¯exi are also striking. In Ae- peomys, the internal segments of the major labial folds (para¯exus and meta¯exus) are oriented anteroposteriorly down the midline of M1±M2, and the lingual ¯exi (proto¯exus and hypo¯exus) are shallow; longitudinal (rather than transverse) enamelled crests are therefore prominent in these teeth. By con- trast, the principal labial ¯exi of Handleyom- ys slant transversely across the midline of the tooth to interpenetrate with much longer lin- gual ¯exi, resulting in the morphology that Voss (1993: 20) termed ``incipient lophodon- ty''. The unworn anterocone of M1 is deeply divided by an anteromedian ¯exus in Aepeo- mys, whereas the M1 anterocone is undivided in Handleyomys. Corresponding differences are apparent in the lower molars as well (®g. 7). Whereas 13 ribs have been reported for Aepeomys lugens (see Steppan, 1995: table 5), all examined postcranial skeletons of Handleyomys fuscatus (ICN 12786) and H. intectus (ICN 12160, 12164) have 12 ribs. Visceral differences that we observed be- tween dissected examples of Aepeomys lu- gens (MBUCV I-2793, I-2794) and Handley- omys intectus (ICN 16092, 16093) include gastric morphology and occurrence of a gall bladder. The stomachs of both genera are unilocular (single-chambered), but gastric glandular epithelium in Aepeomys is restrict- ed to a small, pouch-like structure on the greater curvature (closely resembling the condition illustrated by Carleton [1973: ®g. 5C] for Oxymycterus rutilans). By contrast, Fig. 10. Ventral views of posterior palate and mesopterygoid fossa. A, Aepeomys lugens the entire right half (antrum) of the stomach (USNM 579520); B, Handleyomys intectus (ICN is lined with glandular epithelium in Handley- 16092); C, Oryzomys palustris (AMNH 242524). omys (closely resembling the hemiglandular Abbreviations: mpf, mesopterygoid fossa; ppf, condition illustrated by Carleton [1973: ®g. parapterygoid fossa; ppp, posterolateral palatal 2D] for brevicauda). A large pits; spv, . and distinct gall bladder is present in a cleft between the left and right halves of the cystic lobe of the liver in Aepeomys (as previously reported by Voss, 1991: table 4), but both 16 AMERICAN MUSEUM NOVITATES NO. 3373 dissected specimens of Handleyomys lack a pairs (inguinal, abdominal, postaxial, and gall bladder. pectoral), but Handleyomys has only six teats COMPARISONS WITH ORYZOMYS: Even ex- in three pairs (the pectoral pair is absent; see cluding intectus, the genus Oryzomys re- Voss and Carleton [1993: ®g. 8] for a map mains a morphologically heterogeneous col- of muroid mammary loci). lection of species that de®es meaningful di- Oryzomys is cranially distinctive by virtue agnosis. Because several systematic studies of its deep zygomatic notches (the zygomatic have recently suggested that OryzomysÐ notches are distinct but much shallower in even in its strict modern sense (Musser and Handleyomys; ®g. 8), convergent interorbital Carleton, 1993)Ðis not monophyletic (see region with beaded supraorbital margins (the Myers et al., 1995; Steppan, 1995; Weksler, interorbit is hourglass-shaped with rounded 1996; Bonvincino and Moreira, 2001), we margins in Handleyomys; ®g. 9), widely fen- base our comparisons on a small group of estrated mesopterygoid roof (the mesoptery- taxa that appear to be closely related to the goid roof is entirely bony or only narrowly type species, O. palustris (Harlan). This core perforated by sphenopalatine openings in concept of Oryzomys essentially corresponds Handleyomys; ®g. 10), and highly derived to Goldman's (1918) palustris group and ad- (pattern 3) carotid circulation (versus pattern ditionally includes O. couesi (Alston), O. 1inHandleyomys; see Voss [1988: ®g. 18] dimidiatus (Thomas), O. gorgasi Hershkov- for illustrated examples and explanations of itz, and other possibly valid species currently these morphologies). The lower incisor root regarded as synonyms of O. couesi (see SaÂn- is contained in a prominent bony capsule on chez et al., 2001). In the following para- the lateral surface of the mandible below the graphs we use Oryzomys in this narrowly de- base of the coronoid process in Oryzomys, ®ned sense, acknowledging that additional but a distinct capsular process is absent in genera must eventually be named to contain Handleyomys. Whereas the molar dentition at least some of the many other species cur- of Oryzomys is persistently tubercular and rently referred to this long-abused taxon. nonlophodont, the principal cusps of Carleton and Musser (1989) provided de- Handleyomys molars are quickly worn down tailed illustrations and descriptions of many to a ¯at (planar) occlusal surface dominated morphological traits of O. palustris that by interpenetrating labial and lingual ¯exi should be consulted for additional details of (conforming to the incipiently lophodont characters mentioned below. morphotype de®ned by Voss [1993: 20]). Species of Oryzomys are much more bold- Despite the many points of external and ly marked externallyÐwith abruptly coun- craniodental difference noted above, Oryzo- tershaded heads and bodies and sharply bi- mys and Handleyomys resemble one another colored tailsÐthan species of Handleyomys, in all postcranial-skeletal and visceral char- which lack distinct countershading and have acters scored for this report. In particular, unicolored-dark tails. Whereas the plantar both taxa have 12 ribs and unilocular-hem- surface of the hindfoot is almost entirely cov- iglandular stomachs; the gall bladder is like- ered with squamae and the hypothenar (lat- wise absent in each genus. eral tarsal) pad is indistinct or absent in Ory- PHYLOGENETIC RELATIONSHIPS AND OTHER zomys (see Carleton and Musser, 1989: ®g. COMPARISONS: Pentalophodont sigmodonti- 9A), squamae are only present on the outer nesÐthose with well-developed meso- distal margins of the sole and the hypothenar loph(id)sÐhave traditionally been referred to pad is distinct in Handleyomys (®g. 3). Al- one or the other of two suprageneric groups: though well-developed fringes of silvery one cluster including Thomasomys and its hairs are present along the plantar margins of putative allies (``thomasomyines''), the other the hindfoot, the claws are naked in Oryzo- Oryzomys and its supposed relatives (oryzo- mys; by contrast, no conspicuous hairy fring- myines; see Voss [1993: 21±25] for a histor- es occur along the plantar margins, but the ical review of this dichotomy). Although any claws on digits II±V are concealed by long discussion of the relationships of Handle- tufts of ungual hairs in Handleyomys. Addi- yomys in this report will soon be superseded tionally, Oryzomys has eight mammae in four by taxon-dense phylogenetic analyses of 2002 VOSS ET AL.: NEW GENUS OF ANDEAN RODENTS 17

Fig. 11. Left lateral views of auditory region. A, Aepeomys lugens (MBUCV 2793); B, Handleyomys intectus (ICN 16079); C, Oryzomys palustris (AMNH 242519). Abbreviations: ect, ectotympanic; exo, exoccipital; hp, hamular process of squamosal; per, periotic capsule of petrosal; pgf, postglenoid fo- ramen; psp, posterior suspensory process of squamosal; sq, squamosal; ssf, subsquamosal fenestra; tt, tegmen tympani (of petrosal).

``thomasomyines'' (by VP) and oryzomyines cestry between Aepeomys and Handleyomys (by M. Weksler) that are currently nearing among characters that we do not tabulate, completion, some preliminary remarks about most of which have less well-de®ned states the data at hand are appropriate. and/or more ambiguous polarity. Among those morphological characters In contrast, Handleyomys shares an im- with well-de®ned states for which hypothe- pressive number of shared-derived resem- ses of polarity have been proposed in the lit- blances with Oryzomys, including (1) a long erature (table 1), it is noteworthy that none palate with large posterolateral pits; (2) ab- supports a close relationship between Aepeo- sence of overlap between the tegmen tym- mys (a ``thomasomyine'') and Handleyomys. pani and a posterior suspensory process of Instead, all of the tabulated character states squamosal; (3) absence of an anteromedian shared by these taxa appear to be plesiomor- ¯exus on M1; (4) 12 ribs; and (5) absence of phic on the basis of current assessments of a gall bladder. Because traits 1, 2, 4, and 5 evolutionary transformations in sigmodon- are currently thought to be oryzomyine syn- tine morphology. Likewise, we found no apomorphies (Voss and Carleton, 1993; Step- compelling evidence of recent common an- pan, 1995), we hypothesize that Handleyo- 18 AMERICAN MUSEUM NOVITATES NO. 3373

TABLE 1 Morphological Comparisons Among Aepeomys, Handleyomys, and Oryzomys

mys is a member of that clade, within which and illustrations). In fact, we have not found other taxonomic comparisons are appropri- a single trait in which Melanomys and Han- ate. Three oryzomyine genera deserve partic- dleyomys resemble one another that is not ular attention. also shared with many other small oryzo- In his original description of Oryzomys in- myines. Prima facie, they do not appear to tectus, Thomas (1921) compared it with Mel- be closely related. anomys, remarking general similarities in Ellerman (1941: 341) astutely noted the cranial shape and molar morphology. How- ``highly aberrant dentition'' of Oryzomys in- ever, species of Melanomys differ from Han- tectus with respect to other congeners, and dleyomys by their coarser, glossier, more remarked that ``. . . this species should, I richly colored (usually chestnut- or reddish- think, be transferred to the genus .'' brown) pelage; relatively shorter tail (LT Ͻ Although the incipiently lophodont molars of HBL); shorter hindfeet with blackish soles Handleyomys (®gs. 6, 7) and Nectomys (see and large, almost-naked claws; mammary Hershkovitz, 1944: ®g. 4; GoÂmez-Laverde et complement of eight teats; relatively shorter al., 1999: ®g. 4) are a noteworthy point of rostrum; relatively broader, convergent-bead- shared-derived resemblance, these genera are ed interorbit; relatively much narrower para- dissimilar in most other respects. Species of pterygoid fossae; highly derived (pattern 3) Nectomys are much larger rats (adult HBL Ͼ carotid circulation; lack of subsquamosal fe- 160 mm) that differ from Handleyomys in nestrae; and narrower, nonlophodont molars many external and craniodental traits, includ- (see Goldman [1918, 1920] for descriptions ing their coarser, glossier, water-repellent fur; 2002 VOSS ET AL.: NEW GENUS OF ANDEAN RODENTS 19

Fig. 12. Geographic distribution of Handleyomys in the western Andes (Cordillera Occidental) and central Andes (Cordillera Central) of Colombia based on specimens examined. Numbered points cor- respond to collection localities listed in the gazetteer (appendix). The 1000 m contour is indicated with a broken line; elevations above 2000 m are densely stippled. The western Andes are west of the RõÂo Cauca, and the central Andes are between the RõÂo Cauca and the RõÂo Magdalena; the mountains east of the RõÂo Magdalena are the eastern Andes (Cordillera Oriental). The frame of reference is approxi- mately 3Њ10Ј±9Њ00Ј N and 73Њ30Ј±78Њ20ЈW. Locality 16 is adjacent to localities 17±20 but lacks de®nite geographic coordinates and is not plotted.

relatively longer tail (LT Ͼ HBL); very large mammary complement of eight teats; rela- hind feet with prominent plantar squamae, tively shorter rostrum; much deeper zygo- well-developed , and na- matic notches and correspondingly broader ked claws (Hershkovitz, 1944: ®g. 2c, d); zygomatic plates; convergent-beaded inter- 20 AMERICAN MUSEUM NOVITATES NO. 3373 orbit; highly derived (pattern 3) carotid cir- and one that remains to be determined by culation; and lack of subsquamosal fenestrae. future phylogenetic studies. That Ellerman did not, in fact, transfer intec- tus from Oryzomys to Nectomys probably re- GEOGRAPHIC VARIATION AND SPECIES LIMITS sulted from the lack of supporting evidence from other (nonmolar) characters. All of the material of Handleyomys that Although Voss and Carleton (1993) stated we examined comes from elevations above that all oryzomyines possess eight mammae 1500 m in the western Andes (Cordillera Oc- in four pairs (inguinal, abdominal, postaxial, cidental) and central Andes (Cordillera Cen- and pectoral), subsequent research has shown tral) of Colombia. Because the cool, humid that the oryzomyine genus is char- habitats that prevail above 1500 m in these acterized by having only six teats because the mountain ranges are separated by the hot, pectoral mammary pair is absent (Patton and semi-arid lowlands of the upper RõÂo Cauca da Silva, 1995). Handleyomys likewise has (®g. 12), we tested the hypothesis that pop- only six teats (lacking pectoral mammae), ulations of Handleyomys in the western An- but the phylogenetic interpretation of this des (including the type locality of fuscatus) similarity is ambiguous. Six mammae are are morphologically distinct from allopatric thought to represent the plesiomorphic con- populations in the central Andes (including dition among Neotropical sigmodontines (ac- the type locality of intectus). cording to the ingroup and outgroup assump- Close visual inspection of skins and tions explained by Voss, 1993: 12), so the from both regions revealed three qualitative shared possession of this primitive feature by characters that exhibit signi®cant geographic Scolomys and Handleyomys suggests that variation in trait frequencies. (1) The short these genera might be basal to an oryzomy- hairs that cover the dorsal surface of the hind ine radiation of eight-mammate taxa. Alter- feet are dark-banded (with dense concentra- natively, if Scolomys and Handleyomys are tions of melanin that are distinctly visible un- descended from an oryzomyine ancestor with der low magni®cation) in all examined spec- eight teats, their shared loss of pectoral mam- imens from the western Andes, but most mae provides evidence of a sister-group re- specimens from the central Andes have hind lationship. Obviously, distinguishing be- feet that are covered dorsally with pale tween these (and other) alternatives is im- (pure-white or indistinctly pigmented) hairs. possible in the absence of a comprehensive (2) The nasal bones are long (extending pos- analysis of oryzomyine phylogeny, but it is teriorly well beyond the premaxillary-max- relevant that no other special similarities sug- illary suture; ®g. 13B) in most examined gest a close relationship between Scolomys specimens from the central Andes, whereas and Handleyomys. most specimens from the western Andes Indeed, no oryzomyine genus appears to have short nasals (truncated posteriorly at or exhibit any compelling pattern of synapo- near the premaxillary-maxillary suture; ®g. morphies with Handleyomys apart from those 13A). (3) The incisive foramina are anteri- that support tribal monophyly. Likewise, orly constricted (with lateral margins that are none of the morphologically diverse species abruptly narrowed at or near the premaxil- currently referred to Oryzomys (sensu Mus- lary-maxillary suture; ®g. 14A) in most spec- ser and Carleton, 1993) seems to show any imens from the western Andes, but these special morphological similarity with H. fus- diastemal openings are smoothly tapering catus or H. intectus. Given the Linnaean con- (with evenly rounded lateral margins like pa- vention of binomial nomenclature and the rentheses; ®g. 14B) in most specimens from absence of evidence for close relationship to the central Andes. Although none of these other oryzomyines, we treat Handleyomys as characters exhibits ®xed differences between a genus, but we acknowledge that assigning western and central Andean samples (table ranks to supraspeci®c clades is a biologically 2), it is noteworthy that no examined speci- arbitrary exercise. Determining the hierar- men is geographically atypical in all three chical position of this taxon with respect to scored traits. Thus, no specimen from the other sigmodontine lineages is the key issue, western Andes has pale hind feet, long nasal 2002 VOSS ET AL.: NEW GENUS OF ANDEAN RODENTS 21

logically distinctive populations of Handle- yomys. We also calculated summary statistics for external and craniodental measurements of adult Handleyomys to assess univariate pat- terns of morphometric divergence between populations from the western and central An- des (table 3). External measurement means appear to be similar from the two cordilleras, but we did not apply statistical tests because specimens were measured in the ®eld by many different persons (including inexperi- enced undergraduates) using unknown pro- tocols. We used one-way Analyses of Vari- ance, however, to test for sample differences in craniodental measurements, all of which were taken by us using identical methods. No signi®cant differences were found between cordilleran samples in Condylo-incisive Length, Length of Diastema, Breadth of In- cisive Foramina, Breadth of Palatal Bridge, and Zygomatic Breadth. Other craniodental measurements showed statistically signi®cant differences, but some of these are not empir- ically compelling (e.g., BM1 and LIB, with mean differences Ͻ 0.01 mm). The most vi- sually obvious differences indicated by these tests are in Length of Nasals (already scored as a qualitative trait; table 2, ®g. 12), and Interparietal Breadth. Although the latter measurement exhibits slight overlap between western and central Andean specimens (8.2± 10.2 mm versus 6.2±8.8 mm, respectively), the mean morphological difference is obvi- ous when skulls are viewed side-by-side (®g. 15). Contrasts between western and central Fig. 13. Typical morphology of the nasal Andean skulls in Molar Length, Length of bones in Handleyomys fuscatus (A, ICN 12701) the Incisive Foramina, and Breadth of the and H. intectus (B, ICN 16074). In H. fuscatus, Zygomatic Plate are visually subtle in some the nasals (nas) are usually short because they are pairwise comparisons of local population truncated abruptly at or near the sutures between samples but visually conspicuous in others the premaxillary (pre) and maxillary (max) (see below). bones. In H. intectus, the nasals are usually longer To summarize geographic patterns of mul- because they extend posteriorly beyond the pre- maxillary-maxillary sutures. tivariate morphometric variation, we calcu- lated Mahalanobis distances (D) among ®ve geographic samples, two from the western Andes and three from the central Andes (ta- bones, and smoothly tapering incisive foram- ble 4). In general, morphometric distances ina; nor does any specimen from the central between geographic samples from the same Andes have dark hind feet, short nasal bones, cordillera are smaller than distances between and anteriorly constricted incisive foramina. samples from different cordilleras, a pattern Therefore, these aggregate data suggest that that can be heuristically summarized by clus- the two cordilleras are inhabited by morpho- ter analysis (®g. 16). Because some sample 22 AMERICAN MUSEUM NOVITATES NO. 3373

western and Central Andean samples (ca. 6.5 within-sample multivariate standard devia- tions) reinforces the conclusion previously based on qualitative character data that these mountain ranges are occupied by morpholog- ically differentiated populations of Handley- omys. A valid criticism of cluster-analytic sum- maries of geographic variation is that hier- archical patterns are forced on data that may not, in fact, be hierarchically structured (de Queiroz and Good, 1997). In particular, dis- continuous variation (as resulting from spe- ciation) cannot be distinguished by cluster analysis from intraspeci®c clinal variation without taking the geographic distribution of the analyzed samples into consideration (op. cit.). Intraspeci®c clinal variation is not, however, a plausible explanation for our re- sults, because geographic and morphometric distances are not correlated in Handleyomys. The QuindõÂo/eastern Risaralda sample, for example, clusters with geographically distant samples from Antioquia rather than with the geographically adjacent sample from western Risaralda (just across the Cauca valley). Nevertheless, it is possible that such geo- graphically proximate but morphometrically divergent populations of Handleyomys might show some degree of intergradation. To further assess the distinctness of adja- cent western Andean versus central Andean populations, we used principal components Fig. 14. Typical morphology of the incisive analysis to summarize patterns of multivari- foramina in Handleyomys fuscatus (A, ICN ate craniometric variation in our samples 12701) and H. intectus (B, ICN 16074). The fo- ramina are usually abruptly constricted anteriorly from El Campamento (in western Risaralda; (at or near the premaxillary-maxillary sutures) in locality 14) and La Suiza (in eastern Risar- H. fuscatus, but the foraminal margins of H. in- alda; locality 12). The ®rst two principal tectus are usually smoothly tapering (resembling components account for about 75% of the to- parentheses), without any abrupt change of cur- tal variance (table 5). Projected specimen vature. An intermediate morphology is exhibited scores (®g. 17) indicate complete sample by occasional specimens of both species. separation on the ®rst component, the coef- ®cients of which suggest that La Suiza spec- imens differ from El Campamento specimens by their longer incisive foramina, narrower sizes are small, clustering sequences deter- zygomatic plates, longer nasals, and narrow- mined by small differences in computed dis- er interparietals (essentially the same pattern tances (e.g., within the central Andean clus- of morphometric divergence previously iden- ter) are probably not biologically signi®cant, ti®ed by our univariate comparisons of cen- and some large values (e.g., between Valle tral versus western Andean series). The sec- del Cauca and Western Risaralda) might be ond principal component appears to represent artifactually in¯ated. Nevertheless, the sub- a general size factor, with coef®cients that stantial estimated mean distance between resemble the usual muroid pattern of multi- 2002 VOSS ET AL.: NEW GENUS OF ANDEAN RODENTS 23

TABLE 2 variate craniodental growth allometry (Voss Comparisons of Qualitative Character-State 6 et al., 1990; Voss and Marcus, 1992). Frequencies Between Handleyomys Samples We interpret these results as indicating the from the Central and Western Andesa presence of two species of Handleyomys, one in the western Andes and the other in the central Andes, despite the apparent absence of any completely ®xed qualitative character difference. Fortunately, names are already available for both taxa: H. fuscatus for the populations inhabiting the Cordillera Occi- dental, and H. intectus for populations in the Cordillera Central. We brie¯y summarize the distinguishing attributes of each taxon in the following accounts.

Handleyomys fuscatus (J.A. Allen, 1912) Aepeomys fuscatus J.A. Allen, 1912: 89. Thomasomys fuscatus: Ellerman, 1941: 369 (new name combination). Thomasomys lugens fuscatus: Cabrera, 1961: 431 (new name combination).

TYPE MATERIAL AND TYPE LOCALITY: Al- len's (1912) original description was based on the holotype (AMNH 32230, preserved as a skin and skull) and 11 paratypes, all of which were collected between 7000 and 8000 ft near San Antonio, Valle del Cauca, posteriorly at or near the premaxillary-max- Colombia, by W.B. Richardson from 5 Jan- illary suture; incisive foramina with lateral uary to 31 March 1911. margins that are usually constricted abruptly GEOGRAPHIC DISTRIBUTION: All referred anteriorly; and wide-shallow interparietals. specimens of Handleyomys fuscatus are from VARIATION: All of the material at hand of the western Andes (Cordillera Occidental) of Handleyomys fuscatus comes from two clus- Colombia between 1700 and 2580 m above ters of adjacent localities, one of which is sea level in the departments of Valle del Cau- near Cali in the department of Valle del Cau- ca and Risaralda. ca and the other in western Risaralda de- EMENDED DIAGNOSIS: A species of Handle- partment. Although univariate mean differ- yomys distinguished by hindfeet that are cov- ences between samples from these regions ered dorsally with dark-banded (never pure- are not impressive (table 6), it is noteworthy white) hairs; nasals that are usually truncated that the pattern of divergence is not attrib- utable to a simple size factor. Whereas spec- 6 In most principal components analyses of closely re- lated muroid species, neither PC1 nor PC2 (computed imens from Valle del Cauca average larger from the total covariance matrix of log-transformed than specimens from western Risaralda in measurements) are biologically interpretable because several dimensions (e.g., CIL, LD, LIB, ZB, taxa usually differ in both size and shape. Therefore, IPB), the same specimens have absolutely factors representing general size (the result of postwean- and relatively smaller incisive foramina (LIF, ing growth) and growth-invariant shape differences are typically oblique to the principal component axes and BIF), more slender zygomatic plates (BZP), must be modeled explicitly (see Voss et al. [1990] and and shorter nasals (NL). Because only a few Voss and Marcus [1992] for relevant discussions and measurable adults are available from Valle examples). In the present application, explicit modeling del Cauca, it is possible that some of these is unnecessary because Handleyomys fuscatus and H. intectus appear to differ only in growth-invariant cranial differences are artifactual, but the estimated proportions, which clearly account for the largest axis multivariate distance between Valle del Cau- of dispersion in the total covariance matrix. ca and western Risaralda (Ͼ5 within-sample 24 AMERICAN MUSEUM NOVITATES NO. 3373

TABLE 3 Summary Statisticsa for External and Craniodental Measurements (in millimeters) and Weights (in grams) of Adult Handleyomys from the Western and Central Andes

standard deviations; ®g. 16) is larger than Handleyomys intectus (Thomas, 1921) that observed between any other conspeci®c pair of samples analyzed in this report and Oryzomys intectus Thomas, 1921: 356. Oryzomys (Oryzomys) intectus: Tate, 1932a: 16 suggests that substantial genetic variation (new name combination). might exist among Handleyomys populations ?Nectomys intectus: Ellerman, 1941: 351 (sug- from the Cordillera Occidental. In the current gested name combination). absence of other supporting evidence, it does not seem necessary or useful to recognize TYPE MATERIAL AND TYPE LOCALITY: more than a single taxon in the western An- Thomas's (1921) original description was des, but future karyotypic and molecular da- based on the holotype (BMNH 21.7.1.17, tasets should be carefully assessed for geo- preserved as a skin and skull) and two para- graphic variation among populations here re- types, all of which were collected at Santa ferred to H. fuscatus. Elena, near MedellõÂn, Colombia, by NiceÂforo SPECIMENS EXAMINED: Risaralda, El Cam- MarõÂa in December 1919 and January 1920. pamento (ICN 12799±12827), El Empalado DISTRIBUTION: All referred specimens of (ICN 12700±12703, 12705±12710), La Jalea Handleyomys intectus are from the central (ICN 12704), Los Planes (ICN 12783± Andes (Cordillera Central) of Colombia be- 12793, 12795±12798), Mampay (ICN tween 1500 and 2800 m above sea level in 15277), Siato (ICN 12208, 12725±12727); the departments of Antioquia, QuindõÂo, and Valle del Cauca, Campamento Corea (IND- Risaralda. M 3657, 3659), El Queremal (ICN 6903), EMENDED DIAGNOSIS: A species of Finca la Playa (ICN 4376, 4381, 4383, 4388, Handleyomys distinguished by hindfeet that 4389), PenÄas Blancas (USNM 507267± are usually covered dorsally by pure-white (or 507269), San Antonio (AMNH 32227± indistinctly pigmented) hairs; nasals that usu- 32233, 32236±32239; FMNH 20109). ally extend posteriorly well beyond the pre- 2002 VOSS ET AL.: NEW GENUS OF ANDEAN RODENTS 25

TABLE 4 Composition of Geographic Samples Used to Calculate Mahalanobis Distances

Fig. 15. Typical morphology of the occipital region in Handleyomys fuscatus (A, ICN 12703) and H. intectus (B, ICN 16079). In H. intectus, (FMNH 70296±70306), Santa Elena (BMNH the interparietal (ip) is narrow (transverse dimen- 21.7.1.17±21.7.1.19 [the type series], AMNH sion) relative to its depth (antero-posterior dimen- 37734), Ventanas (FMNH 70332±70338); sion), and the ¯anking sutures between the pari- QuindõÂo, El Roble (AMNH 32928, 32931± etal (par) and exoccipital (exo) are extensive. By 32933, 32937, 32940, 33021), Salento contrast, the interparietal of H. fuscatus is broad (AMNH 32939); Risaralda, La Suiza (ICN relative to its depth, and the parietal-exoccipital 12104, 12158±12179, 12891). sutures are correspondingly shorter. NATURAL HISTORY AND maxillary-maxillary suture; incisive foramina BIOGEOGRAPHY that usually have smoothly tapering (never The predominant type of natural vegeta- abruptly constricted) lateral margins; and nar- tion in the Colombian Andes between 1500 row-deep interparietals. and 2800 m above sea levelÐthe known ele- VARIATION: Available samples of Handley- vational range of HandleyomysÐis cloud omys intectus are remarkably similar forest,7 but anthropogenic habitats (second- throughout the known range of the species, ary growth, croplands, pastures) are also with measurement means that seldom differ widespread at these elevations, and xero- geographically by more than about 0.1 mm morphic formations (dry forests) can occur (table 7). The only noteworthy exception to on rain-shadowed slopes. Explicit descrip- this tendency are the interparietals of speci- tions of macro- and microhabitats where mens from QuindõÂo and eastern Risaralda, which are markedly narrower, on average, 7 We use the nontechnical term ``cloud forest'' for than those of specimens collected in Antio- montane rain forests generally, including the formations quia. Two specimens collected near El Retiro that Grubb (1977) usefully de®ned as Lower Montane Rain Forest, Upper Montane Rain Forest, and Subalpine (ICN 16091, 16093) in the department of Rain Forest. Nevertheless, we cite other synonyms used Antioquia are unusually small but do not ap- by Colombian advocates of Holdridge's (1947) life-zone pear to be exceptional in other respects. nomenclature because these are widely used and pub- SPECIMENS EXAMINED: Antioquia, Finca lished maps are available (Espinal and Montenegro, 1963). For a critical review of ``cloud forest'' and its CanÄaveral (ICN 16091±16095), La Ceja technical synonyms from a ®eld zoologist's point of (AMNH 61575; MLS 187vii, 188vii), La view, see Myers (1969); for another (botanical) perspec- Forzosa (ICN 16073±16079), RõÂo Negrito tive, see Webster (1995). 26 AMERICAN MUSEUM NOVITATES NO. 3373

Fig. 16. Results of UPGMA clustering of Mahalanobis distances (D) among ®ve geographic samples of Handleyomys de®ned in table 4. The samples labeled ``northern Antioquia'', ``QuindõÂo & eastern Risaralda'', and ``southern Antioquia'' are in the central Andes (Cordillera Central) and correspond to the species we recognize as H. intectus. The groups labelled ``western Risaralda'' and ``Valle del Cauca'' are in the western Andes (Cordillera Occidental) and correspond to the species we recognize as H. fuscatus.

TABLE 5 Principal Components Analysis of Craniodental specimens were captured are therefore im- Measurement Variation Among Adult Handleyomys portant for determining the ecological cir- from La Suiza and El Campamentoa cumstances in which small Andean mammals actually occur. Because such descriptions are not available for most historical collecting localities (e.g., ®gs. 18, 19), the following accounts provide ecological information from several sites where Handleyomys has recently been taken by Colombian research- ers. These data, together with additional in- formation summarized in the appendix, pro- vide the basis for subsequent generalizations about the ecogeographic distribution of Han- dleyomys.

HABITAT DESCRIPTIONS Departamento Antioquia, Municipio AnorõÂ (9 km S AnorõÂ), Vereda Roble Ar- riba, bosque La Forzosa, 1775 m: The pro- tected area known as La Forzosa (®g. 12: locality 1) is a ca. 450 ha fragment of pri- mary premontane wet forest (``bosque muy huÂmedo Subtropical'' of Espinal and Mon- tenegro, 1963) on the northeastern ¯anks of 2002 VOSS ET AL.: NEW GENUS OF ANDEAN RODENTS 27

Fig. 17. Specimen scores of Handleyomys intectus (open triangles, from La Suiza [locality 12]) and H. fuscatus (open circles, from El Campamento [locality 14]) on the ®rst two principal components extracted from the variance-covariance matrix of log-transformed craniodental measurements (see text). Variable coef®cients and eigenvalues (scaled as percentages of the total variance) are provided in table 5. the central Andes east of the upper RõÂo NechõÂ imals were taken in Sherman live traps set in (Cuervo et al., 2001). Frequent mists, high mature forest festooned with bromeliads, annual precipitation (2600±3500 mm), and ferns, moss, and orchids; no specimens were high relative humidity (77±95%) characterize taken in traps set along the forest edge or in the local climate, which includes a short dry clearings. Traps that took H. intectus at La season from December to February and a Forzosa were placed on the ground among longer wet season from May to October (op. the elevated roots of palms and other trees, cit.). The local topography is complex, beneath the large leaves of low-growing un- . . . dissected by steep river valleys and gorges. Pri- derstory plants, and inside rotting logs. All mary forests are characterized by a heterogeneous captured individuals apparently entered the canopy, from 6±7 m on the ridges, 15±17 m on the traps at night. Sympatric rodents trapped at slopes, increasing to 20 m along watercourses, with the same locality included Heteromys aus- occasional emergent trees up to 30 m. Understorey tralis (voucher: ICN 16072), Rhipidomys la- cover is densest on the steep slopes, dominated by terrestrial herbaceous plants and by epiphytes on ridg- timanus (ICN 16087, 16088), Oryzomys al- es. The forests are particularly dynamic, with a high bigularis (ICN 16084±16086), and Melano- natural treefall rate and landslides caused by high pre- mys caliginosus (ICN 16080±16083). cipitation on steep slopes with shallow soils. (Cuervo Departamento Antioquia, Municipio El et al., 2001: 361) Retiro (4 km S El Retiro), Vereda Puente Carlos A. Delgado V. collected seven PelaÂez, Finca CanÄaveral, 2100 m: Perched specimens of Handleyomys intectus at this on the western ¯anks of the central Andes in site between 7 and 11 January 2001. The an- the valley of the RõÂo La Miel, the landscape 28 AMERICAN MUSEUM NOVITATES NO. 3373

TABLE 6 Summary Statisticsa for External and Craniodental Measurements (in millimeters) and Weights (in grams) of Adult Handleyomys fuscatus

of Finca CanÄaveral (®g. 12: locality 2) is La Suiza, 1900±1950 m: The area known as dominated by anthropogenic habitats, prin- La Suiza (®g. 12: locality 12) is a protected cipally pastures (for milk cattle) and agricul- area in the valley of the RõÂo OtuÂn on the tural ®elds, but relictual forest (``bosque huÂ- western slope of the Cordillera Central. Al- medo Subtropical'' according to Espinal and though the biological station at La Suiza is Montenegro's [1963] map) survives along surrounded by pastures, agricultural ®elds, the margins of streams that descend from and alder (Alnus acuminata) plantations, continuous forest on steeper slopes above the some primary cloud forest (``bosque muy huÂ- valley ¯oor. Carlos A. Delgado V. collected medo Montano Bajo'' of Espinal and Mon- three specimens of Handleyomys intectus on tenego, 1963; including large specimens of two brief visits to this site, once in August Quercus humboldti) persists along the nearby 2000 and again in March 2001. The Quebrada La Hacienda and on surrounding were trapped at night in dense forest (includ- hillsides. Local rainfall is bimodally distrib- ing Cecropia sp., Quercus humboldti, Inga uted, with average monthly maxima in June sp., Schef¯era sp., Guatteria goudotiana, (237 mm) and November (290 mm); January Vismia sp., Piperaceae, Rubiaceae, ferns, or- is the driest month on average (121 mm), and chids, and bromeliads) along the banks of a the mean annual total is 2038 mm (Aguilar small unnamed tributary of the RõÂo La Miel. and Rangel, 1994). The median annual tem- Other rodents taken in the same traplines as perature is about 24ЊC, with average daily Handleyomys at this site were Akodon af®nis maxima and minima of 33Њ and 16Њ, respec- (vouchers: ICN 16089, 16090), Microryzo- tively (op. cit.). mys minutus (ICN 16096), and Reithrodon- The small fauna at La Suiza was tomys mexicanus (ICN 16097±16100). sampled by ICN researchers on two separate Departamento Risaralda, Municipio Pe- occasions in 1992. On the ®rst visit to this reira, Corregimiento La Florida, Vereda site, MGL and AngeÂlica PenÄuela trapped for 2002 VOSS ET AL.: NEW GENUS OF ANDEAN RODENTS 29

TABLE 7 Summary Statisticsa for External and Craniodental Measurements (in millimeters) and Weights (in grams) of Adult Handleyomys intectus

®ve consecutive nights (22±27 February) in covered at dawn and appeared to have been the dry season; on the second, MGL and Ro- captured during the previous night. cõÂo Polanco trapped for another ®ve consec- Departamento Risaralda, Municipio utive nights (8±13 June) in the rainy season. Pueblo Rico, Vereda SiatoÂ, 1520±1620 m: The equipment used on both visits consisted The steep-sided western foothills of the Cor- of medium-size (ca. 80 ϫ 90 ϫ 230 mm) dillera Occidental near Pueblo Rico (®g. 12: Sherman live traps, large (ca. 100 ϫ 115 ϫ locality 13) have been largely cleared for ag- 380 mm) Sherman live traps, and small snap riculture and animal husbandry, but frag- traps (Museum Specials), all of which were ments of cloud-forest vegetation (``bosque placed on the ground and similarly baited. muy huÂmedo Montano Bajo'' of Espinal and Traplines were located along the margins of Montenegro, 1963) persist along the margins forest clearings, along narrow trails through of local streams. Quantitative climatic data the forest, and along the banks of Quebrada are unavailable, but local precipitation is bi- La Hacienda. Altogether, these faunal-sam- modally distributed with maxima in April± pling efforts amounted to 1163 trap-nights, June and October±November and minima in of which 603 were in February and 560 in January and July. The local fauna of small June. Handleyomys intectus was the most mammals was sampled in the valley of the commonly trapped species at La Suiza on RõÂo Siato (several kilometers SE Pueblo both visits, accounting for 50% of total cap- Rico), where MGL and 33 undergraduate tures (table 8). All trapped animals were re- students from the Universidad Nacional set 30 AMERICAN MUSEUM NOVITATES NO. 3373

Fig. 18. Cloud forest at San Antonio, Departamento Valle del Cauca, Colombia. The holotype and 11 paratypes of Handleyomys fuscatus were collected near San Antonio by William B. Richardson, a professional collector employed by the 1911 AMNH expedition to Colombia. Richardson's skin labels record local captures of H. fuscatus at elevations ranging from 7000 to 8000 ft (ca. 2130±2440 m), but provide no additional ecological details. Although other habitats were present near San Antonio, most of the collecting efforts by AMNH staff at this locality were focused on the forest (see Chapman, 1917: 24), which is indicated as ``bosque pluvial Montano Bajo'' on Espinal and Montenegro's (1963) vege- 2002 VOSS ET AL.: NEW GENUS OF ANDEAN RODENTS 31 medium-size Sherman live traps, National Akodon af®nis (ICN 12614, 12615), Reith- wire live traps (ca. 145 ϫ 145 ϫ 410 mm), rodontomys mexicanus (ICN 12723), Thom- and Museum Specials along the forested asomys cinereiventer (ICN 12623), and T. banks of the Quebrada La Cristalina for eight aureus (unvouchered). nights (20±29 September) in the late-dry/ear- Departamento Risaralda, Municipio ly-wet season of 1991. All traps were set on Santuario, Vereda Los Planes, 2530 m: the ground. A total of 21 captures were re- The eastern slopes of the Cordillera Occi- corded in 405 trap-nights, including 4 spec- dental from the municipal capital of Santu- imens of Handleyomys fuscatus,10Akodon ario (ca. 1500 m) to the vicinity of Los af®nis (vouchers: ICN 12765±12767), 4 Ory- Planes (®g. 12: locality 15) have been almost zomys albigularis (ICN 12715, 12716, completely deforested for agriculture and an- 12719), 2 (ICN imal husbandry, with only fragments of nat- 12717), and 1 (ICN ural vegetation remaining along watercours- 12718). All trapped animals were encoun- es. Above 2500 m, however, is an extensive tered at dawn and were assumed to have landscape of primary cloud forest (``bosque been captured during the previous night. muy huÂmedo Montano Bajo'' of Espinal and Departamento Risaralda, Municipio Montenegro, 1963) that is protected by the Santuario, Vereda El Campamento, 2400± Parque Nacional Natural TatamaÂ. The ICN 2500 m: This locality on the eastern slope of base camp at Los Planes was an abandoned the Cordillera Occidental (®g. 12: locality shelter about an hour and a half by mule 14) is characterized by steep terrain with from ®nca Las Delicias (ca. 2100 m) via a patches of disturbed (selectively logged) pri- dirt road that parallels the RõÂo San Rafael. mary cloud forest (``bosque muy huÂmedo Collections were made just inside the park Subtropical'' of Espinal and Montenegro, boundary, about 500 m above the south bank 1963), agricultural ®elds, and pastures (®g. of the river, in a zone of transition between 2). The local climate is presumably similar the denuded lower slopes and the primary to that described by SaÂnchez-PaÂezetal. forest. The eastern slopes of Tatama are said (1991) from the nearby Parque Nacional Nat- to have a median annual temperature of ural Tatama (see Los Planes, below). ICN re- about 17ЊC and an annual precipitation of searchers Pedro SaÂnchez Palomino and Mar- 2000±2800 mm; precipitation is bimodal, cela Morales trapped small mammals at El with maxima in May and October±Novem- Campamento on six nights (19±24 and 26 ber and minima in February and July (SaÂn- November) during the rainy season of 1991. chez-PaÂez et al., 1991). All traps (medium-size Shermans and Mu- Small mammals were trapped at Los seum Specials) were set on the ground inside Planes on eight consecutive nights (27 Oc- the forest fragments. Seventy traps were set tober to 4 November) in the rainy season of each night, for a total of 420 trap-nights. 1991 by an ICN team consisting of Pedro Handleyomys fuscatus was the most abun- SaÂnchez Palomino, MGL, and Yaneth Mu- dant of seven species taken at this locality, nÄoz-Saba. All of the traps (medium-size Sh- accounting for about 47% of all recorded ermans and Museum Specials) were set on captures. In order of decreasing capture fre- the ground in low, shrubby secondary vege- quency, the other taxa trapped at El Cam- tation that was said to be about ®ve years old pamento were Oryzomys albigularis (vouch- by local inhabitants; primary forest, however, ers: ICN 12001±12023, 12616, 12617), Mi- occurs in close proximity, and some traps croryzomys minutus (ICN 12720±12722), were set along the forest edge. Handleyomys

← tation map. High humidity and cool temperatures are reliably indicated by the profusion of vascular epiphytes and hemiepiphytes (aroids, bromeliads, ferns) and moss that cover tree trunks and limbs in this view and in the accompanying close-up (®g. 19). Essentially similar environments have been de- scribed by most modern collectors of Handleyomys (see text). Photographed in May 1911 by F.M. Chapman. 32 AMERICAN MUSEUM NOVITATES NO. 3373

Fig. 19. Close-up of cloud-forest vegetation at San Antonio (see preceding ®gure caption for eco- geographic details). A local boy (standing just left of center) provides a sense of scale. Clearly shown is the lush vegetation (dominated by large monocots in this view) at ground level, where all specimens of Handleyomys accompanied by microhabitat information have been trapped at other localities. Pho- tographed by in May 1911 by F.M. Chapman. 2002 VOSS ET AL.: NEW GENUS OF ANDEAN RODENTS 33

TABLE 8 Small Mammal Trapping Results at La Suiza responding to Upper Montane Rain Forest in (Table entries are numbers of captures.) his system. In Chapman's (1917: 84±93) classi®cation of Colombian life zones, Han- dleyomys is a member of the humid-Subtrop- ical and humid-Temperate faunas. All of the other sympatric small mammals that have been collected with Handleyomys are likewise known primarily from wet for- ests. Whereas some are commonly encoun- tered in foothill landscapes (e.g., Heteromys australis, Melanomys caliginosus, Sigmo- dontomys alfari), others are typically found at middle elevations (e.g., Oryzomys albigu- laris), and a few are genuinely upper-mon- tane taxa (e.g., Caenolestes fuliginosus, Tho- masomys spp.). Conspicuously absent from all known sites where Handleyomys have been collected are any small mammals pri- marily associated with lowland forests (e.g., Proechimys spp.) or with semiarid habitats (e.g., Sigmodon hispidus, ). Available trapping records suggest that Handleyomys is predominantly terrestrial and fuscatus was the third most-abundant species nocturnal, but these data are less than con- taken in 460 trap-nights at Los Planes, where clusive because traps were not simultaneous- it comprised about 22% of all recorded cap- ly set in trees, and because trapped animals tures (table 9). All trapped animals were re- encountered at dawn might have been caught covered at dawn and were assumed to have before dark the previous day. However, ter- been captured during the previous night. restrial habits are consistent with inferences based on morphology: the long-narrow hind- SUMMARY foot of Handleyomys (with short, nonoppos- Available habitat descriptions and trapping able outer digits; ®g. 3) resembles the con- records indicate that species of Handleyomys dition seen in many other murids with better- occur in both primary cloud forests and ad- documented terrestrial behavior (e.g., Ama- jacent anthropogenic vegetation (secondary zonian species of Oryzomys; see Malcolm growth). Although climatic data are rarely [1991], Patton et al. [2000], and Voss et al. available from the exact sites where speci- [2001] for relevant trapping data). By con- mens have been collected, plotting collection trast, semiarboreal and arboreal muroids typ- localities on published ecological maps sug- ically have shorter, broader hindfeet with gests that Handleyomys is most frequently longer outer digits, of which dV is often se- encountered in relatively cool and very hu- miopposable (Voss et al., 2001: ®g. 53B, C). mid environments (12±18ЊC mean ambient The relatively short, naked-appearing tail of temperature; Ͼ2000 mm annual precipita- Handleyomys is another indication of terres- tion), corresponding to the vegetation zone trial locomotion because arboreal murids that Espinal and Montenegro (1963) desig- usually have relatively longer tails with lon- nate as ``bosque muy huÂmedo Montano ger, coarser hairs that sometimes form a ter- Bajo'' (table 10). In the classi®cation of trop- minal tuft (Voss et al., 2001: ®g. 61). Lastly, ical montane forest vegetation suggested by the dull, drab pelage of Handleyomys is sim- Grubb (1977), the natural habitat at most ilar to that of many other nocturnal montane Handleyomys collection localities would be murids and contrasts with the more richly called Lower Montane Rain Forest, but spec- pigmented fur of some taxa that are known imens have also been taken in habitats cor- to be active in the daytime (e.g., Scotinomys, 34 AMERICAN MUSEUM NOVITATES NO. 3373

TABLE 9 Small Mammal Trapping Results at Los Planes (Table entries are numbers of captures.)

Melanomys; Hooper and Carleton [1976], Unlike certain notoriously elusive Neo- Gardner [1983]). tropical mammals that can only be collected Other aspects of the natural history of in special microhabitats (e.g., ichthyomyines: Handleyomys are obscure. No noteworthy di- Stirton, 1944; Voss, 1988), species of Han- etary information is available; only a few ec- dleyomys appear to be abundant (or at least toparasite records have been published easily trapped) at most places where they (Brennan, 1968); nesting sites, litter sizes, have been found to date. Therefore, the ab- and gestation lengths are unknown; and data sence of specimens from large collections from year-long trapping programs are not obtained by general trapping programs can available to assess reproductive seasonality. be taken as a plausible indication that the ge- Obviously, the absence of concrete infor- nus does not occur in the region(s) where mation about so many biological details pro- such collections were made. It is not likely, vides considerable scope for productive fu- for example, that Handleyomys occurs in the ture ®eldwork on these animals. cloud-forested mountains along the Pana- manian frontier (SerranõÂa de Pirre, SerranõÂa DISCUSSION del DarieÂn), because these highlands have All of the 20 known collecting localities been repeatedly visited by mammalogical of Handleyomys are above 1500 m in the collectors (Anthony, 1916; Goldman, 1920; western and central Andes of Colombia (®g. Pearson, 1939; Handley, 1966, 1972). Simi- 12, appendix), but whether or not these re- larly, sizable collections have accumulated cords provide a reliable indication of the true over the years from many forested (or for- geographic range of the genus remains to be merly forested) localities surrounding Bogota considered. Because most Neotropical land- in the eastern Andes (Cordillera Oriental) of scapes are still poorly inventoried for small Colombia (material that we examined is in mammals, distinguishing patterns of ende- the AMNH, BMNH, FMNH, IAvH, ICN, mism from collecting artifacts is a signi®cant MLS, and USNM), and from the vicinity of research problem. However, some inferences Quito in the Andes of northern Ecuador (rep- about probable range limits can be made. resentative material examined is in the 2002 VOSS ET AL.: NEW GENUS OF ANDEAN RODENTS 35

TABLE 10 Summary Characteristics of Handleyomys Habitats Inferred from Ecological Maps

AMNH, BMNH, and UMMZ). The Sierra omys are unknown from dozens of other lo- Nevada de Santa Marta (on the Caribbean calities in the western and central cordilleras coast of Colombia, north of the frame of ref- that have been trapped (albeit brie¯y) by var- erence depicted in ®g. 12) was the focus of ious mammalogical collectors throughout the intensive general collecting in the late 19th 1900s. Although these negative results might and early 20th centuries (Bangs, 1900; Allen, be overturned by more intensive faunal sam- 1904), and the Cordillera de MeÂrida (in west- pling at some future date, they all tend to ern Venezuela) is mammalogically familiar support the prima facie conclusion from territory (Handley, 1976; PeÂfaur and DõÂaz de known records of positive occurrence that H. Pascual, 1985; Durant and DõÂaz, 1995; So- fuscatus might be narrowly endemic to the riano et al., 1999a, 1999b). The absence of central part of the western Andes and H. in- Handleyomys from all of these relatively tectus to the northern part of the central An- well-sampled areas is compelling evidence des. that the genus is not widely distributed across We are not aware of any other mammalian the mountainous landscapes of northwestern clade that consists of one species restricted South America. to the western cordillera and the other to the Collections from various other localities central cordillera, nor is any mammalian tax- suggest that species of Handleyomys might on currently known to be endemic to just not be widely distributed even in the moun- these two (western ϩ central) mountain rang- tain systems where they are known to occur. es. However, the apparent absence of any Thus, H. fuscatus is notably absent among mammals with the same distribution as Han- the FMNH material collected by Philip dleyomys is not surprising because there has Hershkovitz at Santa BaÂrbara (6Њ25ЈN, been almost no revisionary taxonomic re- 76Њ00ЈW) and other sites in the northern part search to date on northern Andean mammals of the western Andes, nor does it occur in (see below). Although several such studies Kjell von Sneidern's collections (also at are now in progress, only nonmammalian FMNH) from the vicinity of Cerro Munc- vertebrate distributions are currently avail- hique (ca. 2Њ32ЈN, 76Њ57ЈW) in the southern able for biogeographic comparisons. part of the same cordillera. Handleyomys in- Recent progress in alpha-taxonomic re- tectus is likewise absent from FMNH collec- search on the rich amphibian fauna of Co- tions made by near San lombia, for example, has documented rele- Antonio (1Њ57ЈN, 76Њ29ЈW) and San AgustõÂn vant distributional patterns. In his analysis of (1Њ53ЈN, 76Њ16ЈW) in the southern part of the speciation phenomena in the leptodactylid central Andes. In fact, specimens of Handley- frog genus Eleutherodactylus, Lynch (1999a) 36 AMERICAN MUSEUM NOVITATES NO. 3373 identi®ed ®ve sister-species pairs (albericoi- (Chlorochrysa nitidissima, Bangsia melan- lichenoides, deinops-torrenticola, juanchoi- ochlamys). helvolus, ptochus-suetus, ruizi-necopinus), Although the absence of compelling evi- each consisting of one member endemic to dence for western ϩ central Andean ende- the western Andes and the other to the cen- mism in the avian data suggests that distri- tral Andes. In all but one of these pairs, both butional patterns in cloud-forest birds might sister species inhabit essentially the same differ signi®cantly from those of sympatric habitat (cloud forest at middle elevations, ca. nonvolant vertebrates, the continuing discov- 1800±2700 m) as Handleyomys, and the dis- ery of new taxa (e.g., Cuervo et al., 2001) junct areas of endemism they occupy (in the underscores our still-incomplete knowledge central part of the western cordillera and the of the Colombian avifauna. Additionally, the northern part of the central cordillera) cor- possibility that much relevant distributional respond closely to the known distributions of data might be concealed among the numer- H. fuscatus and H. intectus (op. cit.: ®g. 26). ous avian ``subspecies'' discussed (but not Another array of montane amphibian spe- mapped) by Hilty and Brown (1986) merits cies are western ϩ central Andean endemics careful evaluation in any comprehensive fu- with disjunct distributions separated by the ture study of cloud-forest vertebrate ende- valley of the RõÂo Cauca. This fauna includes mism in northwestern South America. three cloud-forest centrolenids (Centrolene Because no simple scenario of vicariance grandisonae, Cochranella ruizi, C. savagei), or dispersal seems likely to account for the three cloud-forest dendrobatids (Colostethus documented complexity of Colombian mon- abditaurantius, Minyobates bombetes, M. tane vertebrate biogeography (Chapman, opisthomelas), one cloud-forest hylid (Hyla 1917, 1926; MuÈller, 1973; Haffer, 1974; columbiana), nine cloud-forest leptodactylids Duellman, 1979; Cracraft, 1985; Lynch et al., (Eleutherodactylus brevifrons, E. boulengeri, 1997), it seems premature to speculate about E. cabrerai, E. erythropleura, E. gracilis, E. historical explanations for the congruent dis- mantipus, E. palmeri, E. permixtus, E. the- tributions of Handleyomys and a few clades copternus), and two gymnophionans (Cae- of eleutherodactyline frogs. Possibly, some cilia occidentalis, C. orientalis). However, paleoecological connection between cloud- each of these amphibian species is more forest habitats in the central part of the west- widely distributed within the western and/or ern Andes and those in the northern part of central Andes than Handleyomys; none is re- the central Andes might be discovered by stricted to just the central part of the western palynological sampling in the RõÂo Cauca val- Andes plus the northern part of the central ley. A more compelling research goal for Andes (Ruiz-Carranza et al., 1996; Lynch systematic mammalogists, however, is to and Rueda-Almonacid, 1997; Lynch et al., evaluate the generality of the biogeographic 1997; Lynch, 1998, 1999b). pattern described above. Is Handleyomys the Birds are the only other Colombian ver- only mammalian clade with this distribution, tebrate group for which a comprehensive or can others be identi®ed among the several source of distributional data is available, but genera and species complexes of sympatric few avian taxa appear to be as narrowly en- small mammals that remain to be revised tax- demic as Handleyomys and the amphibians onomically? Future systematic research on mentioned above. To the best of our knowl- such neglected taxa as Akodon, Thomasomys, edge, ornithologists have not identi®ed any and the Oryzomys albigularis group may sister-species pairs of cloud-forest birds of provide answers to this and other outstanding which one member is endemic to the western questions about mammalian endemism in the Andes and the other to the central Andes, nor Colombian Andes. are many avian species endemic to just those cordilleras. Indeed, only 2 of the 1475 Col- ACKNOWLEDGMENTS ombian bird species mapped by Hilty and Brown (1986) are western ϩ central Andean We are grateful to the curators and support endemics with disjunct cloud-forest distri- staffs of numerous museums that we visited butions separated by the RõÂo Cauca valley or from which we borrowed material for this 2002 VOSS ET AL.: NEW GENUS OF ANDEAN RODENTS 37 report, especially Paula Jenkins (BMNH), remaining errors of fact or omission, how- Bruce Patterson and Bill Stanley (FMNH), ever, are our own. Hugo LoÂpez-AreÂvalo (ICN), and Mike Carle- ton and Al Gardner (USNM). We particularly REFERENCES thank Gonzalo Andrade-C. (director of the ICN) for his assistance with crucial specimen Aguilar P., M., and J.O. Rangel-Ch. 1994. Clima loans to the AMNH, and the Colombian del Parque Regional Natural UcumarõÂ y sec- tores aledanÄos. In J.O. Rangel-Ch. 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APPENDIX: Gazetteer We examined specimens of Handleyomys from ``AgustõÂn Codazzi'' (BogotaÂ), gazetteers (DMA, 20 localities in the western Andes (Cordillera Oc- 1988; Paynter, 1997), and other compilations of cidental) and central Andes (Cordillera Central) of specimen data (e.g., Anderson, 1999). Following Colombia. Numbers preceding each entry in the a semicolon, we provide the name(s) of the col- list that follows are keyed to symbols on the ac- lector(s) and the date(s) on which specimens were companying map (®g. 12). Italic type identi®es collected (in parentheses). For localities lacking Colombian departments, and boldface identi®es explicit descriptions of capture habitats (as sum- locality names as cited in the text of this report. marized in the preceding text), inferences about Geographic coordinates, seldom recorded by local vegetation were based on Espinal and Mon- ®eldworkers, were obtained from various pub- tenegro's (1963) ecological map of Colombia lished sources, including departmental topograph- (from which relevant descriptors are quoted ver- ic maps published by the Instituto Geogra®co batim; e.g., ``bosque muy huÂmedo Montano 2002 VOSS ET AL.: NEW GENUS OF ANDEAN RODENTS 41

Bajo''), taking into account geographical coordi- June 1950). Natural vegetation probably nates and known elevations of collected material. ``bosque pluvial Subtropical''. 7. QuindõÂo, [Municipio Salento], El Roble 1. Antioquia, Municipio Anorõ (9 km S AnorõÂ), [4Њ41ЈN, 75Њ36ЈW; on W slope of central An- Vereda Roble Arriba, bosque La Forzosa, des], 7200 ft [2195 m]; Leo E. Miller (9 No- 6Њ59ЈN, 75Њ08ЈW [on W slope of central An- vember 1911). Natural vegetation probably des], 1775 m; Carlos A. Delgado V. (7±11 ``bosque muy huÂmedo Montano Bajo''. January 2001). See text for a description of 8. QuindõÂo, [Municipio Salento], Salento local habitats. [4Њ38ЈN, 75Њ34ЈW; on W slope of central An- 2. Antioquia, Municipio El Retiro (4 km S El des], 7000 ft [2134 m]; Leo E. Miller (No- Retiro), Vereda Puente PelaÂez, Finca CanÄ- vember 1911). Natural vegetation probably averal, 6Њ01ЈN, 75Њ30ЈW [on E slope of cen- ``bosque muy huÂmedo Montano Bajo''. tral Andes], 2100 m; Carlos A. Delgado V. 9. Risaralda, Municipio MistratoÂ, Vereda El (11 August 2000, 14 March 2001). See text Empalado [ca. 5Њ22ЈN, 75Њ54ЈW; on E slope for a description of local habitats. of western Andes], 1700±1900 m; Marcela 3. Antioquia, [Municipio La Ceja], La Ceja Morales (29 March±3 April 1992). Included [6Њ02ЈN, 75Њ26ЈW, on E slope of central An- sublocalities are ``km 13 carretera MistratoÂ- des at 2217 m]; NiceÂforo MarõÂa (1919, 1921). Pueblo Rico'', ``km 13 carretera MistratoÂ- Natural vegetation probably ``bosque huÂmedo Costa Rica'', and ``km 13 carretera MistratoÂ- Montano Bajo''. San Antonio del ChamõÂ''. Natural vegetation 4. Antioquia, [Municipio MedellõÂn], Santa probably ``bosque muy huÂmedo Montano Elena [ca. 6Њ13ЈN, 75Њ32ЈW; in central An- Bajo''. des], 9000 ft [2743 m]; Howarth S. Boyle (19 10. Risaralda, Municipio MistratoÂ, Vereda La November 1911), NiceÂforo MarõÂa (December Jalea, km 8 carretera MistratoÂ-San Antonio 1919±January 1920). The exact provenance del Chamõ [ca. 5Њ21ЈN, 75Њ53ЈW; on E slope of Boyle's and Niceforo MarõÂa's material of western Andes], 1720 m; Marcela Morales from Santa Elena, which is said to be near (4 April 1992). Natural vegetation probably MedellõÂn (Chapman, 1917; Thomas, 1921), is ``bosque muy huÂmedo Montano Bajo''. uncertain because several localities near the 11. Risaralda, Municipio MistratoÂ, Vereda Mam- pay, km 10 carretera MistratoÂ-San Antonio city are named Santa Elena or Santa Helena de ChamõÂ[5Њ22ЈN, 75Њ53ЈW; on W slope of (phonetically equivalent names in Spanish). western Andes], 1950 m; Marcela Morales (7 Fortunately, all of them fall within a small April 1992). Natural vegetation probably region, approximately 9 ϫ 9 km, that can rep- ``bosque muy huÂmedo Montano Bajo''. resented by a single point on our map (®g. 12. Risaralda, Municipio Pereira, Corregimiento 12). The best match with Boyle's recorded La Florida, Vereda La Suiza [ca. 4Њ44ЈN, elevation (Niceforo MarõÂa did not record this 75Њ35ЈW; on W slope of central Andes], datum) corresponds to the above coordinates 1900±1950 m; Marcela GoÂmez-Laverde, An- on IGAC's (1979) 1:25,000 topographic geÂlica PenÄuela, and RocõÂo Polanco (23 Feb- sheet. Paynter's (1997) longitude for Santa ruary±13 June 1992). Included sublocalities Elena (71Њ10ЈW) is an obvious mistake. Nat- are ``al borde de la Quebrada La Hacienda'', ural vegetation at ca. 2700 m probably ``por el camino hacia el Cerro Morro Azul'', ``bosque muy huÂmedo Montano''. and ``Quebrada La Hacienda''. Specimens re- 5. Antioquia, [Municipio] SonsoÂn, 9±15 km E ported from this locality were all collected RõÂo Negrito [5Њ42ЈN, 75Њ10Ј±75Њ13ЈW; on W within the 1992 boundaries of the departmen- slope of central Andes], 1700±2100 m; Philip tally-administrated Parque Regional Natural Hershkovitz (4±19 October 1950). Natural UcumarõÂ, but the sublocalities listed above vegetation probably ``bosque muy huÂmedo are now part of the nationally-administrated Montano Bajo''. According to Dr. Robert P. Santuario de Fauna y Flora del OtuÂn-Quim- Anderson (personal commun.), it is likely that baya. See text for a description of local hab- Hershkovitz's specimen labels from this lo- itats. cality refer to measured distances from the 13. Risaralda, Municipio Pueblo Rico, Vereda municipal center (SonsoÂn) along the RõÂo Ne- Siato [5Њ14ЈN, 76Њ02ЈW; on W slope of west- grito, rather than to distances from the river ern Andes], 1520±1620 m; Marcela GoÂmez- as implied by his syntax. Laverde (22±23 September 1991). Included 6. Antioquia, [Municipio] Valdivia, Ventanas sublocalities are ``Quebrada la Cristalina'', [7Њ05ЈN, 75Њ27ЈW; on NW slope of central ``Inmediaciones de la Quebrada la Cristali- Andes], 2000 m; Philip Hershkovitz (20±27 na'', and ``borde de la Quebrada la Cristali- 42 AMERICAN MUSEUM NOVITATES NO. 3373

na''. See text for a description of local habi- W slope of western Andes], ca. 1800 m; Ser- tats. afõÂn Arango (28 July 1978). Local vegetation 14. Risaralda, Municipio Santuario, Vereda El probably ``bosque pluvial Montano Bajo''. Campamento [ca. 5Њ07ЈN, 75Њ58ЈW; on E 18. Valle del Cauca, Municipio PichindeÂ, Finca slope of western Andes], 2500 m; Pedro SaÂn- la Playa [ca. 3Њ27ЈN, 76Њ37ЈW; on E slope of chez and Marcela Morales (19±23 November western Andes], ca. 1800 m; Lucio VelaÂsquez 1991). See text for a description of local hab- (26 October±2 November 1966). Local veg- itats. etation probably ``bosque muy huÂmedo Mon- 15. Risaralda, Municipio Santuario, Vereda Los tano Bajo''. New species of chiggers (Acari- Planes, Parque Nacional Natural Tatama na: Trombiculidae) were described by Bren- [5Њ08ЈN, 76Њ04ЈW; on E slope of western An- nan (1968) from specimens of Handleyomys des], 2530 m; Marcela GoÂmez-Laverde, Pe- fuscatus collected at this locality. dro SaÂnchez, and Yaneth MunÄoz-Saba (28 19. Valle del Cauca,RõÂo PichindeÂ, PenÄas Blan- October±3 November 1991). See text for a cas [ca. 3Њ27ЈN, 76Њ43ЈW; on E slope of west- description of local habitats. ern Andes at 2000 m]; Alfred L. Gardner 16. Valle del Cauca, Campamento Corea, three (23±24 October 1974). Local vegetation hours on horseback from Cali [not located; in probably ``bosque pluvial Montano Bajo''. western Andes], 2500±2800 m; P. O. Lowe 20. Valle del Cauca, San Antonio [3Њ30ЈN, (17±18 November 1977). Based on elevation, 76Њ38ЈW; on E slope of western Andes], natural vegetation probably ``bosque pluvial 7000±8000 ft [2134±2438 m]; William B. Montano''. Richardson (5 January±31 March 1911). Lo- 17. Valle del Cauca, Municipio Dagua, El Quer- cal vegetation probably ``bosque pluvial emal, Antena de Tokio [3Њ29ЈN, 76Њ44ЈW; on Montano Bajo''. Recent issues of the Novitates may be purchased from the Museum. Lists of back issues of the Novitates and Bulletin published during the last ®ve years are available at World Wide Web site http://library.amnh.org. Or address mail orders to: American Museum of Natural History Library, Central Park West at 79th St., New York, NY 10024. TEL: (212) 769-5545. FAX: (212) 769- 5009. E-MAIL: [email protected]

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