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A New Montane Species of Spiny Pocket Mouse (Rodentia: Heteromyidae: Heteromys) from Northwestern Costa Rica PDF

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Preview A New Montane Species of Spiny Pocket Mouse (Rodentia: Heteromyidae: Heteromys) from Northwestern Costa Rica

PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, NY 10024 Number 3509, 38 pp., 10 figures, 4 tables March 16, 2006 A New Montane Species of Spiny Pocket Mouse (Rodentia: Heteromyidae: Heteromys) from Northwestern Costa Rica ROBERT P. ANDERSON,1,2 AND ROBERT M. TIMM3 ABSTRACT Recent taxonomic works have recognized only two species of spiny pocket mice of the genus Heteromys (Rodentia: Heteromyidae) from Costa Rica. Within Costa Rica, the widespread H. desmarestianusisconsideredtooccurthroughoutthewetCaribbeanlowlands,aswellasatmiddle andhighelevationsontheCaribbeanandPacificslopesofthecountry’smainmontanesystems.In contrast,H.oresterusisknownfromonlyafewlocalitiesathighelevationsinthewesternportion of the Cordillera de Talamanca in central Costa Rica. Our morphological and morphometric analysesofspecimensfromnorthwesternCostaRicarevealthepresenceofanundescribedspecies of the genus, which we describe as Heteromys nubicolens. This new species ranges from 750 to 1840m in elevation in the Cordillera de Tilara´n and Cordillera de Guanacaste. Heteromys desmarestianusisfoundinthesurroundingmesiclowlandsandfoothills.Externally,bothspecies possessdarkbrowndorsalpelage,butH.nubicolensdiffersbyoveralllargersizeandbydistinctive cranialproportions.Inmostcranialmeasurements,H.nubicolensislargerthanH.desmarestianus; however, H. desmarestianus has a wider interorbital region and a wider braincase. Known populations of H. nubicolens occur in three highland areas (Monteverde, Volca´n Rinco´n de la Vieja–Volca´n Santa Mar´ıa, and Cerro Cacao), but populations in these areas are probably disjunct, being separated by intervening lowlands. Heteromys nubicolens is likely widespread throughouttheCordilleradeTilara´nandCordilleradeGuanacaste,butitspresenceinotherareas ofthecountryisunlikely.EvenafterrecognizingH.nubicolensasdistinctfromH.desmarestianus, morphological,karyological,andgeneticdataindicatethatH.desmarestianusrepresentsaspecies complex. Further research is therefore necessary to evaluate the taxonomic status of this species complexinotherregionsofthecountryandotherpartsofitswidespreadgeographicdistribution. RESUMEN Recientestrabajostaxono´micoshanconsideradoqueso´lodosespeciesderatonesdeabazones (5rato´nbolsero,ratonbolso´n,rato´nmochilero)delge´neroHeteromys(Rodentia:Heteromyidae) 1DivisionofVertebrateZoology(Mammalogy),AmericanMuseumofNaturalHistory([email protected]). 2DepartmentofBiology,CityCollegeoftheCityUniversityofNewYork,NewYork,NY10031([email protected]. cuny.edu). 3NaturalHistoryMuseumandDepartmentofEcology&EvolutionaryBiology,UniversityofKansas,Lawrence,KS 66045([email protected]). CopyrightEAmericanMuseumofNaturalHistory2006 ISSN0003-0082 2 AMERICAN MUSEUMNOVITATES NO. 3509 habitanenCostaRica.Heteromysdesmarestianus,conunaampliadistribucio´ngeneral,habitaen esepa´ısenlastierrasbajashu´medasdelCaribe,as´ıcomotambie´nenlasestribacionescariben˜asy pac´ıficasdelasprincipalescadenasmontan˜osas,enaltitudesmediasyelevadas.Porelcontrario, H.oresterusseconoceso´lodeunaspocaslocalidadesdealtamontan˜aenlaparteoccidentaldela Cordillera de Talamanca, en la regio´n central de Costa Rica. Nuestros ana´lisis morfolo´gicos y morfome´tricos de ejemplares provenientes del noroccidente costarricense revelan la presencia de unaespecienodescritadelge´nero,lacualdescribimosaca´ comoHeteromysnubicolens.Estanueva especiehabitaentrelos750ylos1840mdealtitudenlaCordilleradeTilara´nylaCordillerade Guanacaste. Heteromys desmarestianus se encuentra en el piedemonte y en las tierras bajas hu´medasadyacentesaestascordilleras,rodeandoladistribucio´ndeH.nubicolens.Externamente ambas especies poseen un pelaje dorsal de coloracio´n parda oscura, pero H. nubicolens se caracterizaportenerengeneralunmayortaman˜ocorporalyproporcionescraneanasdistintivas. En la mayor´ıa de las medidas craneanas, H. nubicolens posee un mayor taman˜o que H. desmarestianus;sinembargo,H.desmarestianustieneunamayoranchuradelaregio´ninterorbitaly delacajacraneana.LaspoblacionesconocidasdeH.nubicolensseencuentranentresa´reasdealta montan˜a(Monteverde,elVolca´nRinco´ndelaVieja–Volca´nSantaMar´ıayelCerroCacao),pero estaspoblacionesprobablementeposeendistribucionesdisyuntas,estandoseparadasporlastierras bajasqueseinterponenentreellas.EsprobablequeH.nubicolenstengaunaampliadistribucio´nen laCordilleradeTilara´nylaCordilleradeGuanacaste,perosupresenciaespocoprobableenotras regionesdelpa´ıs.InclusoconelreconocimientodeH.nubicolenscomounaespeciediferenteaH. desmarestianus, datos morfolo´gicos, cariolo´gicos y gene´ticos indican que H. desmarestianus constituyeuncomplejodeespecies.Porlotanto,senecesitanestudiosadicionalesparaevaluarla taxonom´ıa de este complejo de especies en otras partes del pa´ıs y en otras regiones de su amplia distribucio´ngeogra´fica. INTRODUCTION Talamanca.Nearlycontinuousatanelevation of 1500 m, it contains several peaks higher Although Costa Rica encompasses an area than 2500 m. A broad connection, the Valle of only ca. 51,000 km2, the region’s complex Central (or Meseta Central), joins the geological history and highly variable topog- Cordillera Central and the northwestern por- raphy and climate have contributed to a di- tion of the Cordillera de Talamanca at verse fauna and flora (Janzen, 1983; elevations of ca. 1000–1500 m. To the north- McPherson 1985, 1986; Wilson et al., 2002). west of the Cordillera Central, an older range This portion of the Central American isthmus of lower (,2000 m) Tertiary volcanic peaks was formed by anextremely complexseries of and ridges forms the Cordillera de Tilara´n, events related to the subduction of the Cocos which is continuous at an elevation of ca. Plate under the Caribbean Plate (Coates and 1200 m.Finally,theCordilleradeGuanacaste Obando, 1996; Denyer et al., 2000; Montero- represents the northernmost mountain range P.,2000).Today,fourmajormountainranges in the country. Similar in origin to the exist in Costa Rica, each oriented diagonally Cordillera Central, it runs from the from southeast to northwest (fig. 1); these Cordillera de Tilara´n to near the Nicaraguan rangesvarygreatlyinoriginandage(Castillo- borderand iscomprisedofaseriesofisolated M., 1984; Bergoeing, 1998; Alvarado et al., Quaternary volcanoes, most of which reach 2000; Kussmaul, 2000; Salazar-Mondrago´n, 1500–2000 m.Lowpassesbetweenmostofthe 2000). The Cordillera de Talamanca repre- volcanoes of the Cordillera de Guanacaste sents the highest and most massive montane connecttheCaribbeanandPacificlowlandsat system in southern Central America. Formed elevations of 500–700 m. To the north of this primarily during the Tertiary, this range of range lie the expansive Nicaraguan lowlands. mixed geological origin extends from western Panama to central Costa Rica, with many peaksreachingmorethan3000 minelevation. SPINY POCKET MICE The Cordillera Central of Costa Rica is a younger (Quaternary) volcanic range that The rodent family Heteromyidae is com- lies to the north of the Cordillera de prised of three subfamilies: Heteromyinae 2006 ANDERSON AND TIMM:COSTA RICAN SPINYPOCKET MOUSE 3 Fig.1.MapofCostaRicaandadjacentregionsofNicaraguaandPanama,showingthepositionofmajor mountainranges.Grayshadingdenotesregionshigherthan1000minelevation,andareasshowninblack lieabove 2000m. (spiny pocket mice), Dipodomyinae (kanga- semiarid tropical and subtropical habitats roo rats and kangaroo mice), and Pero- from northern Mexico and southern Texas to gnathinae (silky pocket mice). Heteromyines Panama (Genoways, 1973; Morales and represent a well-defined monophyletic group Engstrom, 1989; Rogers and Engstrom, distinct from either of the two other living 1992; Williams et al., 1993). In contrast, subfamilies (Hafner, 1981; Hafner and species of Heteromys inhabit wetter (typically Hafner, 1983; Wahlert, 1991; see also Ryan, evergreen) forests from southern Mexico to 1989: 94–98; Brylski, 1990). Two extant western Ecuador (Williams et al., 1993; genera,HeteromysandLiomys,arerecognized Anderson, 1999; Anderson and Jarr´ın-V., in the Heteromyinae and can be distinguished 2002; Anderson, 2003b). from each other by a number of morpholog- Present taxonomy recognizes eight species ical characters (Anderson, 2003b). Species of of Heteromys, including two recently de- Liomys inhabit deciduous forests and other scribed from northern South America 4 AMERICAN MUSEUMNOVITATES NO. 3509 (Williams et al., 1993; Anderson and Jarr´ın- MATERIALS AND METHODS V., 2002; Anderson, 2003b; but see Patton, MUSEUM SPECIMENS 1993). Of these, only three species are known from southern Central America (Nicaragua, We examined 401 specimens of Heteromys Costa Rica, and Panama): H. desmarestianus, from the principal study area of northwestern H. oresterus, and H. australis (Patton, 1993; CostaRica,representingallknowntousfrom Williamsetal.,1993;Reid,1997;Wilsonetal., that region (appendix 1). For the purposes of 2002; see also Goodwin, 1946; Wainwright, thispaper,wedefinenorthwesternCostaRica 2002). As currently conceived, the widespread as those areas of the country northwest of the H. desmarestianus ranges from southern depression that separates the Cordillera de Mexico (Estado de Veracruz) to northwestern Tilara´n from the Cordillera Central,4 and Colombia in a wide variety of both lowland north of the crest of the Cordillera Central and montane habitats. In contrast, H. ores- (fig. 1). This study region includes the totality terus is endemic to the Cordillera de oftheProvinciadeGuanacaste,largeportions Talamanca in central Costa Rica. Finally, of the Provincia de Alajuela and Provincia de the primarily South American species H. Heredia, andtheextremenorthernpart ofthe australis shows a marginal distribution in Provincia de Puntarenas. Although our main extreme eastern Panama (Anderson, 1999). interest was in comparing the distinctive Although Heteromys desmarestianus is populations of Heteromys found at upper often an abundant member of small mam- elevations of the Cordillera de Tilara´n and mal communities in Mexico and Central Cordillera de Guanacaste with populations Americaand hasbeenwell studiedecological- of H. desmarestianus present in the immedi- ly at a few classical localities (e.g., Fleming, ately adjacent foothills and lowlands, delimi- 1983; Timm et al., 1989; Sa´nchez-Cordero, tation of the present principal study area 1993), several studies have indicated that it also allowed us to make comparisons with represents a complex of externally similar specimens from highland regions in the species. Across the range of the species Cordillera Central and with a much larger sample of H. desmarestianus from the complex, considerable variation exists in Caribbean lowlands. karyotypes and allozymes (Mascarello and We also examined critical specimens from Rogers, 1988; Rogers, 1989, 1990) and in other regions (appendix 2). These included all cranial morphology (Rogers, 1986). Recent Costa Rican and Panamanian voucher speci- fieldwork by several researchers in the mens from karyological and genetic studies Cordillera de Tilara´n and Cordillera de (Mascarello and Rogers 1988; Rogers 1989, Guanacaste in northwestern Costa Rica has 1990). Furthermore, we examined representa- led to new collections of a distinctive species tivesamplesofallcurrentlyrecognizedspecies of Heteromys. We herein undertake a revision of Heteromys, including almost all holotypes of Heteromys in northwestern Costa Rica, and lectotypes representing nominal taxa describe a species new to science, provide currently referred to the genus. The only detailed morphological comparisons between holotype or lectotype in the genus that we it and adjacent populations of H. desmares- did not examine is that of H. desmarestianus tianus, and summarize the natural history psakastus;inlieuoftheholotype,weexamined and biogeographic information available eight paratypes. Apparently, no holotype for the new species. This work represents exists for H. thompsonii, a name that Lesson a step toward characterizing the morpholo- (1827)clearlyusedtorefertothespinypocket gy and distributions of H. desmarestianus mouse from Trinidad named by Thompson and the species currently confused with it. We hope that it will facilitate future studies 4The southern slope of the division between the by other researchers, leading to a more CordilleradeTilara´nandtheCordilleraCentralismarked complete taxonomic understanding of approximately by the course of the R´ıo Barranca, these common rodents so characteristic of although the hills of the Montes del Aguacate to the southeastofthatriver(nearSanRamo´n)aresimilartothe rainforest habitats in the northern Cordillera de Tilara´n in geological origin (Bergoeing, Neotropics. 1998;Kussmaul,2000). 2006 ANDERSON AND TIMM:COSTA RICAN SPINYPOCKET MOUSE 5 (1815)asMusanomalus,nowreferredtoasH. UMMZ*University of Michigan Museum of anomalus. In considerations of species bound- Zoology,AnnArbor aries, we apply morphological and (when USNM United States National Museum of available) karyotypic and/or genetic data to NaturalHistory,Washington,DC allow evaluation under the evolutionary spe- cies concept (Wiley, 1978). We examined external and cranial morpho- Localities and specimens examined from logical characters, making comparisons northwestern Costa Rica are detailed in the among specimens of approximately the same Gazetteer (appendix 1), and other specimens age. Cranial nomenclature follows Wahlert examined are listed separately (appendix 2). (1985), Anderson (1999, 2003b), and Locality information not provided by the Anderson and Jarr´ın-V. (2002). Specimens collector appears in brackets and, where appli- were assigned to the age classes of Rogers cable, is followed by the source. Where the and Schmidly (1982) based on patterns of original elevation was reported in feet, we tooth eruption, toothwear, and molt. Age provide that datum as well as the metric classes 1–3 represent juveniles and subadults; equivalent to the nearest whole number. classes 4–6 are progressively older adults. Specimensexaminedarehousedinthefollowing Age classes do not constitute a continuous museumcollections(whereapplicable,abbrevia- variable (such as absolute age), but rather are tions follow Hafner et al., 1997—except for categories roughly corresponding to relative UCR, see Savage, 2002). An asterisk denotes age within population samples (Voss et al., museums with material from northwestern 1990). CostaRica. MEASUREMENTS AMNH American Museum of Natural History,NewYork Standard cranial measurements for ANSP Academy of Natural Sciences of Heteromys (fig. 2) follow Anderson and Philadelphia,Philadelphia Jarr´ın-V.(2002)andweretakentothenearest BM(NH)Natural History Museum, London 0.01 mm with digital calipers. We measured [formerly British Museum (Natural alladultspecimensofHeteromysinageclass4 History)] with intact skulls from northwestern Costa EBRG Museo de la Estacio´n Biolo´gica de Rica, as well as from all available samples of RanchoGrande,Maracay,Aragua H.oresterus.Externalmeasurementsandmass FMNH* Field Museum, Chicago [formerly were copied from specimen tags and, when FieldMuseumofNaturalHistory] necessary, from primary field notes. ICN Instituto de Ciencias Naturales, Occipitonasal length (ONL): greatest dis- Universidad Nacional de Colombia, tance from anteriormost projection of nasal Bogota´ bones to posteriormost portion of occipital KU* UniversityofKansasNaturalHistory bone. Museum,Lawrence Zygomatic breadth (ZB): greatest width LACM* Natural History Museum of Los across zygomatic arches at right angle to AngelesCounty,LosAngeles longitudinal axis of cranium. LSUMZ*Louisiana State University Museum Rostral length (RL): greatest distance ofNaturalScience,BatonRouge from notch lateral to lacrimal bone to ante- MCZ Museum of Comparative Zoology, riormostprojectionofnasalboneonsameside HarvardUniversity,Cambridge of cranium. MNCR* Museo Nacional de Costa Rica, San Nasal length (NL): greatest distance from Jose´ anteriormost projection of one nasal bone to MVZ* Museum of Vertebrate Zoology, itsposteriormostprojection(notnecessarilyat UniversityofCalifornia,Berkeley medial suture between nasals). ROM* RoyalOntarioMuseum,Toronto Least interorbital constriction (IOC): least UCR* Museo de Zoolog´ıa, Universidad de width across interorbital constriction at right CostaRica,SanJose´ angle to longitudinal axis of cranium. 6 AMERICAN MUSEUMNOVITATES NO. 3509 Fig. 2. Dorsal, ventral, and lateral views of a cranium of Heteromys showing the methods for taking measurements.Abbreviations andmeasurements are definedin Materials andMethods. Squamosal breadth (SB): width across ietal bone, always taken along medial line of squamosals anterior to external auditory cranium even when notch present in posterior meatus at right angle to longitudinal axis of border. cranium. Parietal breadth (PB): greatest width Maxillary toothrow length (MTR): dis- across parietal crests at right angle to longi- tancefromanteriorlipofalveolusofpremolar tudinal axis of cranium. to posterior lip of alveolus of third molar. Skull depth (SD): greatest distance from Interparietal width (IW): greatest trans- dorsalmost point of braincase to horizontal verse width measured from lateralmost pro- plane passing through ventral borders of jections of interparietal bone at right angle to maxillary cheek teeth and ventral borders of longitudinal axis of cranium. occipital condyles (taken by placing skull on Interparietal length (IL): greatest dis- glass microscope slide with upper incisors tancefromanteriormostprojectionofinterpar- rested overedgeofslide,andthen subtracting ietal bone toposteriormost borderof interpar- thickness of slide). 2006 ANDERSON AND TIMM:COSTA RICAN SPINYPOCKET MOUSE 7 STATISTICS of H. oresterus with those of the montane species from northwestern Costa Rica. Specimens in age class 4 (the most abun- Finally, we conducted a principal compo- dant adult age class) were used for all nents analysis (PCA) of specimens from quantitative comparisons. Statistical analyses northwestern Costa Rica, based on the vari- were performed in Minitab (1998; release ance-covariance matrix of log -transformed e 12.1), and probabilities were compared to cranial measurements. We interpreted the a 5 0.05 for hypothesis testing. Material multivariate axes by examination of loadings from northwestern Costa Rica was used and of coefficients (elements) of the unit for the primary analyses detailed here. Using eigenvector. Loadings are Pearson product- species assignments based on our qualitative moment correlationcoefficientsbetweenspec- morphological examinations, we calculated imen scores on each axis and the log - descriptive statistics for standard external e transformed variables. measurements, mass, cranial measurements, and three derived ratios (tail length/head-and- body length; least interorbital constriction/ occipitonasal length; and parietal breadth/ SYSTEMATICS occipitonasal length). For these analyses, all localities for each species were pooled A NEW SPECIES OF HETEROMYS FROM because sample sizes were too small at most NORTHWESTERN COSTA RICA individual sites. Additionally, we conducted Our analyses indicate that two species of a series of two-tailed t-tests comparing means Heteromys are present in northwestern Costa of the two species for each measurement or Rica.Inthisregion,Heteromysdesmarestianus ratio. is widely distributed in mesic areas from low We then conducted analyses of three geo- elevations to ca. 1000 m in the Cordillera de graphic samples in northwestern Costa Rica Tilara´n and Cordillera de Guanacaste, but it withrelativelylargesamplesizes.Intwocases, ascendstomorethan2000 mintheCordillera nearby localities were pooled to create the Central. The other species is restricted to samples. La Selva (localities 18, 19; appendix middle and high elevations of the Cordillera 1),CerroCacao(locality28),andMonteverde de Tilara´n and Cordillera de Guanacaste (localities 30–32; all on the Pacific slope or (750–1840 m). It displays cranial proportions crest of the Cordillera de Tilara´n) constituted and measurements distinct from those of H. these samples. First, we calculated the same desmarestianus, as well as differences from all descriptive statistics mentioned above. Then, other recognized species of the genus. As no to test for morphological differentiation be- available name exists for this species, we tweenhighlandsamplesfromtheCordillerade describe it as: Tilara´n and the Cordillera de Guanacaste, we conducted a series of two-tailed t-tests com- paring means of the samples from Cerro Heteromys nubicolens, new species Cacao and Monteverde for those measure- Cloud-dwelling Spiny Pocket Mouse ments and ratios. We also compared the highland species Figures 3, 5, 6, 7 from northwestern Costa Rica with Heteromys oresterus (known only from the HOLOTYPE: KU 159025, nulliparous adult Cordillera de Talamanca) using the measure- female; skin, skull, and postcranial skeleton in ments and ratios mentioned above. To do so, excellent condition; plus frozen tissues origi- we first calculated standard descriptive statis- nally preserved in 95% ethanol (fig. 3). tics for specimens from the type locality of H. Collected on 16 October 2000 from COSTA oresterus (El Copey de Dota). Next, we RICA:PUNTARENAS:Monteverde,Monteverde calculated descriptive statistics for all individ- Cloud Forest Reserve, Investigator’s Trail, uals (of age class 4) that we consider to be H. 10u189N, 84u489W, at 1550 m elevation by oresterus (appendix 2). We then conducted Robert M. Timm and Christy M. McCain; a series of two-tailed t-tests comparing means originalnumberRMT4468. 8 AMERICAN MUSEUMNOVITATES NO. 3509 Fig. 3. Dorsal, ventral, and lateral views of the cranium of the holotype of Heteromys nubicolens (KU 159025), anadult femalein ageclass 4.See appendix1 forfull provenience. 2006 ANDERSON AND TIMM:COSTA RICAN SPINYPOCKET MOUSE 9 PARATYPES: We designate as paratypes the suture dipping well ventral to parietal crest following 24 specimens (adults in age classes posterior to its widest point; braincase not 4–6; skins and skulls in good condition) from inflated; interorbit narrow; rostrum long and the Pacific slope or crest of the Cordillera de cylindrical; skull large (ONL 35.72–41.02 mm Tilara´nintheMonteverderegion,andhoused in adult specimens of age class 4; tables 1, 2), in a variety of museum collections (appendix elongated and relatively narrow; body size 1): COSTA RICA: PUNTARENAS: Monteverde, average to large for genus; dorsal pelage soft 1450 m, ROM 97307; Monteverde, Arthur to moderately spiny; dorsal coloration dark Rockwell’scafetales,1400 m,UMMZ115419, brown and only faintly grizzled with thin 115420; Monteverde, Cerro Amigos, 1790 m, ochraceoushairsintermixedamongspines;no KU 142057; Monteverde, Hoge woods, lateral ochraceous band present on flanks; 1420 m, LACM 64867; Monteverde, John plantar surface of hind feet naked. Campbell’s woods, 1520–1580 m, FMNH DESCRIPTION: Dorsal pelage (fig. 5) soft to 128417, 128419, 128420, 128423, 128425; KU moderately spiny and dark brown (sharply 142791; LACM 64863, 64865; MVZ 161224, contrasting with soft, pure white pelage of 161225; Monteverde, Monteverde Cloud venter), onlyfaintlygrizzled with thin ochrac- Forest Reserve, Investigator’s Trail, 1550 m, eous hairs intermixed among spines (grizzling KU 159022–159024, 159026, 159027, 159029; generally less pronounced along midline, pro- MNCR1336;Monteverde,QuebradaQuecha, ducing slight dorsal stripe); ears dark brown KU 143337; Monteverde, Stella Wallace’s to dark gray and small to medium in size; tail house, KU 143339. strongly bicolored for most of its length (then ETYMOLOGY: The adjective nubicolens, or unicolored dark distally), generally longer ‘‘cloud-dwelling,’’ derives from the Latin thanhead-and-bodylength(tables1,2);patch nubes (cloud) and colo (dwell, inhabit) and is of dark coloration present on dorsal and applied here in reference to the species’ externalsurfacesofforearms,continuouswith distributionincloudforestspresentonmoun- darkcolorationofflanks;ventralandinternal tainsthatriseabovethesurroundinglowlands surfaces of forearms white; hind feet large of northwestern Costa Rica (Brown, 1956: (35 mm or greater in adults; tables 1, 2), with 213, 478). naked plantar surface; skull (figs. 3, 6, 7) DISTRIBUTION: Known only from the moderately large for genus (tables 1, 2); Cordillera de Tilara´n and Cordillera de anteriorhalfofpremaxillaryconvex(inflated), formingasmooth(notstepped)lateralborder Guanacaste of northwestern Costa Rica of rostrum (in dorsal view); rostrum long and (fig. 4). In the Monteverde region of the cylindrical (not tapered anteriorly or with Cordillera de Tilara´n, it is found from 750 to dorsal flare anteriorly); nasals inflated anteri- 1840 montheCaribbeanslopeandfrom1350 orly; interorbital constriction narrow; brain- to 1840 m on the Pacific slope. In the case narrow,not inflated; parietaland tempo- Cordillera de Guanacaste, the species has ral crests weakly to moderately developed; been collected at 1100–1500 m on Cerro interparietal moderately wide, often with Cacao and from 800 to 1200 m on Volca´n slight anterior point; incisive foramina gener- Rinco´n de la Vieja–Volca´n Santa Mar´ıa. See ally thin and slightly tapering anteriorly; no also Sympatry and Zones of Contact with swellingatposteroventralborderofinfraorbi- Heteromys desmarestianus, below. tal foramen5; mesopterygoid fossa V-shaped, DIAGNOSIS: A species of spiny pocket with long, thin hamular processes of ptery- mouse with adults showing the following goids; shallow parapterygoid fossa; postalar combination of characters (figs. 3, 5, 6, 7): p4 (lower permanent premolar) with 3 lophs; 5Erroneously listed as ‘‘anterodorsal border of infra- P4 (permanent upper premolar) with long, orbitalforamen’’inAnderson(2003b).Inmostspeciesof curved fold in anterior border of posterior Liomys,adistinctswellingispresentattheposteroventral loph; mesopterygoid fossa V-shaped, with borderoftheinfraorbitalforamen(generallyanterodorsal long, thin hamular processes of pterygoids; totheincisiveforamina).Suchaswellingisneverpresent in species of Heteromys. Other, more salient and optic foramen small, with exterior margin consistent differences between the two genera are listed formed bystrong barofbone;parietomastoid inAnderson(2003b:11). 10 AMERICAN MUSEUMNOVITATES NO. 3509 ds anote s,en ud n ag estidin ra ah ms sy dera sG y om1). eterdix Hn e ofpp a s( ocalitiezetteer la ntG esehe rt pn rei n rclesgive Cia a.dat c Ritom. ad0 Costespone200 nrv stercorabo hwedtolie ortereack nmbbl ysinnunin mrew oao Heterorocalitiesareassh fLd alitiesolens.on,an ctionlocH.nubicnelevati e i ollofm wingcalities1000 shoelocthan potr Fig.4.Matrianglesdenregionshighe

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