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Rodentia, Echimyidae) AMERICAN MUSEUM Novitautes PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, N.Y. 10024 Number 2817, pp. 1-14, figs. 1-4, tables 1-6 June 11, 1985 A Review of the Genus Isothrix (Rodentia, Echimyidae) JAMES L. PATTON' AND LOUISE H. EMMONS2 ABSTRACT Two species of the echimyid rodent genus Iso- geographic races (bistriata [including villosa De- thrix are recognized within the Amazon and upper ville, negrensis Thomas, molliae Thomas, and bo- Orinoco basins of South America based on color liviensis Petter and Cuenca Aguirre] and orinoci and color pattern, bacular morphology, cranial Thomas), and I. pagurus Wagner. No localities of morphometrics, and karyotype. These represent sympatric contact between these two species are significant departures in the previously recognized known, although their ranges come very close in name combinations: I. bistriata Wagner, with two the general vicinity of Manaus in central Brazil. INTRODUCTION The genus Isothrix contains arboreal rats transverse plates which persist into advanced ofmoderate body size that are a well-marked age. component of the morphologically and eco- Despite the relative distinctness ofIsothrix logically diverse neotropical rodent family as a generic unit, however, no clear under- Echimyidae. Erected by Wagner in 1845 to standing of its component taxa is currently accommodate three species, it is character- available. Specimens ofthis genus are rare in ized by a pelage lacking spines, a bushy tail, collections, a state which has resulted in the and teeth with complex, irregular re-entrant description ofgeographic units at the specific angles and enamel folds. The combination of or subspecific levels without benefit ofproper soft-haired pelage and bushy tail differen- comparisons to permit a realistic assessment tiates Isothrix from other echimyid rodents ofindividual relative to geographic variation. that have similar occlusal patterns (for ex- The present paper was motivated by the ex- ample, Echimys Cuvier [1809] and AMesomys istence of large series of Isothrix from three Wagner [1845]). Diplomys Thomas (1916a), general areas of the Amazon Basin and ad- also a soft-haired form without spines, differs jacent Venezuela, mostly in the collections of from Isothrix by its rather simple teeth with the American Museum of Natural History. I Research Associate, Department of Mammalogy, American Museum of Natural History; Curator of Mammals, Museum of Vertebrate Zoology, University of California, Berkeley, California 94720. 2 Research Associate, Division ofMammals, National Museum ofNatural History, Smithsonian Institution, Wash- ington, D.C. 20560. Copyright © American Museum of Natural History 1985 ISSN 0003-0082 / Price $1.70 2 AMERICAN MUSEUM NOVITATES NO. 2817 These allow us to examine for the first time bistriata negrensis Thomas, 1920 both individual variation and geographic Isothrix villosa villosa (Deville, 1852) trends, and thus to make a better assessment villosa molliae Thomas, 1924 of the taxonomic diversity within the genus. Isothrix pictus (Pictet, 1843) TAXONOMIC HISTORY OF This view has been accepted even though ISOTHRIX Oldfield Thomas suggested that several of Nine names have been proposed as taxa these named forms most likely represented within the genus Isothrix. Wagner (1845), in the same taxon. In 1928, for example, he the original definition ofthe genus, described stated that L v. villosa and L v. molliae from three species: bistriata, with type locality Rio northern Peru were the same entity, and both Guapore, Mato Grosso, Brazil; pachyura, type in 1916 and 1928 he suggested that L villosa locality not identified (but subsequently des- might in fact grade into L bistriata and thus ignated as Cuiaba, Mato Grosso, Brazil by would represent only an upper Amazon sub- Cabrera [1961]); and pagurus, type locality species of the latter. Borba, Rio Madeira, Amazonas, Brazil. In 1848 Wagner renamed pachyurus as crassi- METHODS AND ABBREVIATIONS cauda without reason. A fourth species, vil- losa, was proposed by Deville (1852), based Ninety-six specimens of Isothrix from on Castlenau's specimen from Sarayacu Mis- eastern Peru, eastern Bolivia, southern Ven- sion, Rio Ucayali (not Rio Urubamba as giv- ezuela, and central Brazil were examined in en in Cabrera [1961]), Loreto, Peru. Subse- this study. Eighty-four of these were consid- quently, Oldfield Thomas described three ered adults by the criteria given below. Stan- forms as subspecies ofeither bistriata (orinoci dard external measurements were missing [Thomas, 1899] and negrensis [Thomas, from the labels of most specimens, so anal- 1920]) or villosa (molliae [Thomas, 1924]). yses of individual and geographic variation Finally, the name boliviensis was recently were based on external color and color pat- proposed as a Bolivian subspecies ofbistriata tern and morphometric variables of the cra- by Petter and Cuenca Aguirre (1982). nium. For the cranium, the following 19 mea- The named form pictus (Pictet, 1843), a surements were taken from each skull with soft-haired echimyid from coastal Bahia, dial calipers reading to 0.05 mm: (1) greatest eastern Brazil, has been varyingly placed in length of skull [GSL]; (2) basilar length of the genus Isothrix (for example, Waterhouse, Hensel [BaL]; (3) zygomatic breadth [ZB]; (4) 1848; Ellerman, 1940; Cabrera, 1961), in a mastoid breadth [MB]; (5) least interorbital monotypic genus Nelomys (Pictet, 1843; constriction [IOC]; (6) rostral length [RL]; (7) Goldman, 1916; Thomas, 1916a), or in Echi- nasal length [NL]; (8) rostral width [RB]; (9) mys (Tate, 1935; Moojen, 1952). This dif- rostral depth [RD]; (10) diastema length [D]; ference of opinion results from pictus com- (11) palatal length A [PLA]; (12) palatal length bining simple laminated upper cheek teeth B [PLB]; (13) maxillary tooth row length (characteristic ofDiplomys) with the complex [MTRL]; (14) maxillary breadth [MaxB]; (15) lower teeth of Isothrix and Echimys. We fol- incisive foramen length [IFL]; (16) bullar low Cabrera (1961) and Honaki, Kinman, and length [BuL]; (17) post palatal length [PPL]; Koeppl (1982) in placing pictus within Iso- (18) cranial depth [CD]; and (19) mesopter- thrix, but do not consider it further. ygoid fossa width [MPFW]. Each of these Most recent authors (for example, Honaki, measurements is described and figured by Kinman, and Koeppl, 1982; Petter and Cuen- Patton and Rogers (1983) for the related ge- ca Aguirre, 1982) have followed the taxo- nus Proechimys. nomic arrangement given by Cabrera (1961) Variation in- bacular dimensions was ex- in amined from cleared and stained glandes, recognizing three species, as follows: prepared as in Hooper (1959). Specimens were Isothrix bistriata bistriata Wagner, 1845 aged according to nine categories of tooth bistriata pachyura Wagner, 1845 eruption and wear patterns: (I) dP4 erupted, bistriata pagurus Wagner, 1845 M' erupted but below occlusal level; (II) MI bistriata orinoci Thomas, 1899 to occlusal level; M2 erupting; (III) M3 erupt- 1985 PATTON AND EMMONS: ISOTHRIX 3 FIG. 1. Sampled localities of Isothrix specimens examined in this report; numbered localities are as identified in the text. The type localities for all named forms are indicated by asterisks. ing; (IV) median re-entrant fold confluent with degree of character difference between the primary fold on dP4 to M3; (V) median fold named forms recognized by Cabrera (1961) separate on dP4, open on MI through M3; (VI) and others. Finally, overall geographic trends median fold open on M2 through M3; (VII) and degrees of differentiation between local- median fold open only on M3; (VIII) median ities were examined by the multivariate pro- fold separate from primary fold on all teeth; cedure ofdiscriminant function analysis. The and (IX) entire tooth row worn, all folds iso- statistical routines of SAS (Barr et al., 1982) lated. Age classes V through IX were consid- and SPSS (Nie et al., 1975) were used for all ered to represent adult individuals, although analyses. Samples from each locality were no data are available on individual repro- considered as separate entities in the exam- ductive status in each ofthe various age classes ination of both univariate and multivariate recognized. geographic trends. The degree ofnon-geographic variation due Karyotypes from two individuals, one from both to age and sex was assessed by two-way northern Peru and one from central Brazil, analysis of variance (ANOVA) following the were prepared following the technique ofPat- suggestion ofLeamy (1983). One-way ANO- ton (1967). VAs were utilized to assess the significance Specimens forming the basis of this report of single character variation within a taxon are in the collections of the American Mu- over a geographic area and to examine the seum ofNatural History, New York (AMNH), 4 AMERICAN MUSEUM NOVITATES NO. 2817 TABLE 1 Amorin (5, AMNH), Rio Tapajoz, Limoal Non-geographic Variation for Cranial Variables (1, AMNH). in a Combined Sample of Isothrix from Northern Locality 12-BRAZIL: Amazonas; 80 km Peru (Localities 31 to 34)a N (by rd) Manaus (1, USNM). Variableb Age Sex Age x Sex Residual Locality 21-BOLIVIA: Beni; Rio Itenez at Costa Marques (1, AMNH). GSL nsc (24.6) ns (7.9) ns (0.0) (67.5) Locality 31-PERU: Loreto; Pto. Indiana, BaL ns (22.6) ns(ll.5) ns (0.0) (65.9) Rio Amazonas (25, AMNH). ZB *d (15.9) * (13.5) ns (11.0) (59.6) MB ns (0.0) * (7.2) * (33.6) (59.2) Locality 32-PERU: Loreto; Orosa, Rio IOC * (28.3) ns (4.2) ns (0.0) (67.5) Amazonas (5, AMNH). RL * (28.0) ns (1.2) ns (0.0) (70.8) Locality 33 -PERU: Loreto; Sarayacu, Rio NL ns (16.7) ns (4.1) ns (0.0) (82.2) Ucayali (14, AMNH). RW ns (11.8) ns (6.0) ns (0.0) (82.2) Locality 34-PERU: Loreto; Boca Rio RD ns (1.2) ns (0.0) ns (0.0) (98.8) Curaray (1, AMNH). D ns (15.0) ns (9.6) ns (0.0) (75.4) Locality 35-PERU: Loreto; Rio Itaya near PLA ns (12.2) ns (3.9) ns (0.0) (83.9) Iquitos (1, AMNH).
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