A New Genus and Species of Phyllotine Rodent from Bolivia Author(S): Sydney Anderson and Terry L
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American Society of Mammalogists A New Genus and Species of Phyllotine Rodent from Bolivia Author(s): Sydney Anderson and Terry L. Yates Source: Journal of Mammalogy, Vol. 81, No. 1 (Feb., 2000), pp. 18-36 Published by: American Society of Mammalogists Stable URL: http://www.jstor.org/stable/1383124 . Accessed: 29/09/2011 16:57 Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at . http://www.jstor.org/page/info/about/policies/terms.jsp JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact [email protected]. American Society of Mammalogists is collaborating with JSTOR to digitize, preserve and extend access to Journal of Mammalogy. http://www.jstor.org Journal of Mammalogy, 81(1): 18-36, 2000 A NEW GENUS AND SPECIES OF PHYLLOTINE RODENT FROM BOLIVIA SYDNEY ANDERSON AND TERRY L. YATES* Department of Mammalogy, American Museum of Natural History, New York, NY 10024-5192 (SA) Museum of Southwestern Biology, University of New Mexico, Albuquerque, NM 87131 (TLY) A new species (Muridae, Sigmodontinae, Phyllotini), belonging to a new genus, is described on the basis of 2 specimens from 1 locality in the mountain forests of southeastern Bolivia. Diagnostic features are posteriorly divergent edges of supraorbital region, large and hyp- sodont molar teeth with somewhat prismatic pattern, and anterior zygomatic process not projecting as an overhanging point. The diploid chromosome number is 56 and the fun- damental number is 76. Other external, cranial, dental, and karyologic characters in the tribe and new taxon are described, illustrated, and discussed. The hypothesized phylogenetic placement of the new genus based on gene-sequence, chromosomal, and morphologic data is presented relative to evolutionary relationships of selected phyllotine taxa. Key words: Bolivia, new species, Phyllotini,rodent, Sigmodontinae,Yungas In July 1991, our Bolivian Expedition their relatively large size, noticeable hyp- camped at an elevation of 1,500 m in humid sodonty, and relatively prismatic form sug- forest at the village of Tapecua in the de- gested a condition between the smaller, partment of Tarija (Fig. 1). Here, the main less-specialized teeth of Graomys and the east-west road in Tarija crosses a divide on larger, more acutely angled teeth of Andi- the crest of 1 of 7 parallel ridges that lie nomys. Subsequent chromosomal analysis between the arid high altiplano with its revealed similarity to Andinomys in number puna vegetation (Cabrera and Willink 1980) of chromosomes, but not in number of to the west and the lowland Gran Chaco chromosomal arms. A review of these and with its xeric, thorn-scrub vegetation to the other genera of phyllotines revealed that no east. existing diagnosis of any phyllotine genus At Tapecua, we captured 2 rats with dark would accommodate the new rats without the of the hind feet. Their size areas on tops changing more than one-half of the state- Andino- and general appearance suggested ments about what characters were diagnos- mys edax. However, the pelage seemed tic. coarser and the ears slightly larger (in com- The collection of 1991 included series of to size of head and When parison body). other phyllotine species of the region (e.g., we examined the uncleaned skulls in the Phyllotis xanthopygus, P. osilae, P. capri- field, the of the posterior divergence edges nus, P. wolffsohni, Graomys domorum, and of the frontal bone in the area supraorbital G. griseoflavus, in addition to A. edax). The and its width were unlike these fea- greater new rats were quite different from any of tures in Andinomys, and suggested Gra- these and from Andalgalomys. Thus, a new omys. When the skulls were cleaned in the genus is proposed for the new species. laboratory and the teeth were examined, Steppan has studied cladistic relationships * Correspondent: tyates @sevilleta.unm.edu within the tribe Phyllotini, and in 1993 in- 18 February 2000 ANDERSON AND YATES-NEW PHYLLOTINE RODENT 19 match those of the earlier paper in all cases. This -10 69~ 66 63 60 10- makes it difficult to compare scorings for char- -- f 0 - 1000 m / acter states in studies or to add taxa in 1000 - 3000 m previous E studies. In a few cases, we were un- 12- subsequent e, >3000 m certain what exact dimensions were to be mea- sured so some variation may be present between S'-,',, ' -14 - 14- our scorings and those of these authors. Also, in assessing the state of genera in the cladogram, we believed it best to include more than 1 spe- cies for those that have more than 1. Thus, we added Andalgalomys pearsoni, Calomys callo- from -18 '18- sus, and Graomys domorum, using data *' % Braun (1993) wherever possible. Our resulting character matrix is not reproduced here but is -20 120 available on request. Polymerase chain reaction and nucleotide se- quencing.-Total genomic DNA was isolated -22 0 250 22 from frozen liver tissue following Hillis et al. 69 66 63 60 (1996). The cytochrome-b gene of the mito- A I I I chondrial DNA was amplified using the poly- FIG. 1.-Map of Bolivia showing the location merase chain reaction (PCR). The primers used of Tapecua in relation to the major elevational to amplify the cytochrome-b gene were light- belt between 1,000 and 3,000 m known as the strand (Mus 14095) 5' GAC ATG AAA AAT Yungas and Valles Region. CAT CGT TGT AAT TC 3' and heavy-strand (Mus 15398) 5' GAA TAT CAG CTT TGG GTG TTG RTG 3'. Double-stranded DNA us on this new cluded data provided by spe- (dsDNA) products were obtained with PCR am- cies (Steppan 1993). Another study (Braun plification using Taq DNA polymerase (Saiki et 1993) on phenetic and cladistic relation- al. 1986, 1988); PCR amplifications were per- ships in the tribe was completed before dis- formed in 25-lI reactions. Samples were dena- covery of this new species. tured at 950C for 45 s, primers were annealed at 500C for 40 s, and segments were extended at METHODSAND MATERIALS 700C for 2 min with 4 s added to each extension mM Cranial measurements were taken with a stage for 40 cycles; 2.5 pl of reaction buffer (500 25 mM craniometer as described by Anderson (1969). KCl; 100 mM Tris-HC1, pH 8.4-8.7; 0.5 bovine serum 4 The midfrontal width was measured at the mid- MgCl2; p1 albumin), p1 dle of the suture between the frontal bones as deoxynuclepside triphosphates (2 mM of each seen in dorsal view. We reviewed characters deoxyadenosine triphosphate, deoxycytidine tri- and used in diagnosis and comparisons of Andino- phosphate, deoxyguanosine triphosphate, in 10 mM Tris- -mys, Graomys, Salinomys, and Andalgalomys deoxythymidine triphosphate each was with other genera by various authors (Braun and HC1, pH 7.9), and 2 pmole of primer of the Mares 1995; Gyldenstolpe 1932; Hershkovitz added to every reaction. The efficiency 1962; Olds et al. 1987; Osgood 1947; Thomas amplification was verified by visualizing 5 il of 1902). Most of these characters are included in the dsDNA product on a 0.8% agarose gel the description or comparisons below. stained with ethidium bromide. We also attempted to code, for the new spe- Primers used to amplify the cytochrome-b cies, all characters used by Braun (1993) and gene also were used in the sequence reaction. In Braun and Mares (1995). Although 21 of the 25 addition, 2 internal primers (light-strand 5' GTT characters we used matched those of Braun GAA TGA ATC TGG GGC GG 3' and heavy- (1993), the character states do not match for all strand 5' AAC GAC AAG GTT AGC GAT characters. In addition, in the later paper (Braun AAA GG 3') were used to sequence the entire and Mares 1995), even fewer states were used gene. Specimens were sequenced on an Applied and ranges of values assigned to states do not Biosystems 377 DNA Sequencer (Perkin-Elmer 20 JOURNALOF MAMMALOGY Vol. 81, No. I Applied Biosystems, Foster City, California). (International Commission of Zoological Sequences were attained using the Taq Dye- Nomenclature 1985). See Olds and Ander- Deoxy@ TerminatorCycle SequencingKit (Per- son (1989) for Vorontsov's tribal reference. kin-Elmer Applied Biosystems, Warrington, United Kingdom) and following protocols pro- Tribe Phyllotini Vorontsov, 1959 vided therein. Sequences reportedin this paper The tribe was diagnosed by Olds and An- were depositedin the GenBankdatabase (acces- derson and In their sion numbersAF159283-AF159294). (1989) Steppan (1995). of the tribe Olds and Data analyses.-Maximum parsimony was diagnosis Phyllotini, used to derive a phylogeny from the nucleotide- Anderson (1989) included the following sequence data and separatelyfrom morphologic combination of characters: 1, heel hairy; 2, data. All maximum-parsimonyanalyses were ears moderate to large; 3, palate long (ex- conductedusing PAUP,version 3.1.1 (Swofford cept in Irenomys); 4, incisive foramina 1993). In the morphologic analysis, characters long; 5, parapterygoid fossa relatively were treatedas and unordered,weighted equally, broader than mesopterygoid fossa (except that was the exhaus- analysis performedusing in Punomys); 6, sphenopalatine vacuities tive-searchoption. In the analysis of nucleotide- large; 7, supraorbital region never evenly sequence data, characterswere treated as unor- curved in cross section; 8, interparietal well dered, discrete characters with four possible notch ex- states (A, C, G, and T). Analyses were per- developed; 9, zygomatic deeply cised so in teeth tetra- formed using the branch-and-boundsearch op- (less Irenomys); 10, tion. An unequalweighting scheme was used in lophodont; 11, M3 50% the length of M2.