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Cricetidae, Sigmodontinae) and Additional Comments on the Distinctiveness of Pearsonomys

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Cricetidae, Sigmodontinae) and Additional Comments on the Distinctiveness of Pearsonomys ARTICLE IN PRESS www.elsevier.de/mambio Original investigation New data of the long-clawed mouse Pearsonomys annectens (Cricetidae, Sigmodontinae) and additional comments on the distinctiveness of Pearsonomys By G. D’Elı´a, Agustina A. Ojeda, F. Mondaca and M.H. Gallardo Seccio´n Evolucio´n, Facultad de Ciencias, Montevideo, Uruguay; Grupo de Investigacio´n en Biodiversidad, Instituto Argentino de Investigaciones de Zonas A´ridas, Mendoza, Argentina; and Instituto de Ecologı´ay Evolucio´n, Universidad Austral de Chile, Valdivia, Chile Receipt of Ms. 28.12.2004 Acceptance of Ms. 26.8.2005 Abstract The monotypic genus Pearsonomys is one of the least studied sigmodontine genera, so far known only from its holotype. We record four new localities from the Regio´n de Los Lagos (southern Chile) for P. annectens expanding its known distribution by approximately 217 km. Next, we present a phylogenetic analysis based on nuclear DNA sequences aimed to test the phylogenetic position of Pearsonomys. Results of this analysis corroborate that Pearsonomys belongs to the abrotrichine radiation being sister of Geoxus. Subsequently, we provide descriptions of P. annectens molars (tetralophodont pattern with reduced mesoloph/id), penis (complex type), stomach (unilocular- hemiglandular), and karyotype (2n=56; FN=62). The discussion of these new data is framed according the results of the phylogenetic analysis. r 2005 Deutsche Gesellschaft fu¨r Sa¨ugetierkunde. Published by Elsevier GmbH. All rights reserved. Key words: Pearsonomys, rodentia, Chile, valdivian forest Introduction The sigmodontine genus Pearsonomys, Patter- nature of the holotype, an old adult son 1992 belongs to a recently recognized male with advanced tooth wear, few radiation of mostly central and southern details were provided about its dental mor- Andean genera of the New World endemic phology. Similarly, the karyotype of P. subfamily Sigmodontinae, for which the annectens is so far unknown. Remarkably, informal name of ‘‘Anden clade’’ or abjro- other than the original description, only one trichines was coined (Smith and Patton 1999; additional study has analyzed samples of see also D’Elia 2003). Pearsonomys; this was a study based on Currently Pearsonomys encompass only one cytochrome b gene sequences aimed to assess species, P. annectens, Patterson 1992, which sigmodontine phylogenetic relationships was described on the basis of one specimen (Smith and Patton 1999). Consequently, of from the area of Mehuı´ n in the southern all abrotrichine genera, Pearsonomys is by far Chilean province of Valdivia. Due to the the least known; indeed, it is one of the least 1616-5047/$ - see front matter r 2005 Deutsche Gesellschaft fu¨r Sa¨ugetierkunde. Published by Elsevier GmbH. All rights reserved. doi:10.1016/j.mambio.2005.08.004 Mamm. biol. ] (]]]]) ] Á ]]]–]]] ARTICLE IN PRESS 2 G. D’Elı´aetal. studied genera of the entire sigmodontine Applied Biosystems). Sequencing reactions were radiation. run in an ABI 377 automated sequencer. In all With the goal of partially filling the gap on cases, both heavy and light DNA strands were our knowledge about the long-clawed mouse sequenced. Sequences of both strands were recon- ciled using Sequencer Navigator version 1.0.1 P. annectens, the aim of this study was to (Applied Biosystems). All sequences were deposited present new recording localities for this in GENBANK (Table 1). Sequence alignment was species, provide a description of its dentition, carried out with the program Clustal X (Thompson penis, stomach, and karyotype, as well as et al. 1997), by using the default values for all present a phylogenetic analysis based on alignment parameters. Aligned sequences were nuclear DNA sequences aimed to test the subjected to maximum parsimony analysis (MP; phylogenetic position of this taxon. Farris 1982; Kluge and Farris 1969). Characters were treated as unordered and equally weighted. PAUP* 4 (Swofford 2000) was used to perform 200 replicates of heuristic searches with random addi- Material and methods tion of sequences and tree bisection-reconnection branch swapping. Two measures of topology All voucher specimens, tissue samples, and cell support were carried out. First, 1000 parsimony suspensions are housed at the mammal collection jackknife replications (JK; Farris et al. 1996) with of the Instituto de Ecologı´ a y Evolucio´ ndela three addition sequence replicates each, the dele- Universidad Austral de Chile, Valdivia, Chile tion of one-third of the character data, and (IEEUACH). Measurements, sex, and locality data MAXTREE set to 5000 were performed. Branches were recorded from specimen tags. Geographic with less than 50% of support were allowed to distances were estimated, on the basis of the collapse. Second, Bremer support values (BS; latitude and longitude data recorded in specimen Bremer 1994) were computed for each node in tags. PAUPÃ using command files written in TreeRot version 2 (Sorenson 1999). Phylogenetic analysis We have included representatives of all sigmodon- Karyotypic analysis tine tribes as well as several sigmodontine genera Four specimens (IEEUACH 1064, 1745, 3851, whose phylogenetic relationships are not clear (see 7057) of P. annectens were karyotyped using D’Elı´ a 2003). Considering previous findings that standard chromosome techniques. Mitotic meta- place Pearsonomys within the abrotrichine group phases from direct bone-marrow were obtained (Smith and Patton 1999), taxonomic coverage is following a colchicine-hypotonic technique (Verma particularly broad with regards to abrotrichine and Babu 1995). Bone marrow was incubated in taxa, for which, when possible, more than one 0.1 ml (0.05%) colchicine plus 9.9 ml KCl (0.075 M) individual per species was included. We have for 40 min at 371C and subsequently fixed in 3:1 included outgroups (Nixon and Carpenter 1993) methanol: glacial acetic acid. Non-differential that are representative of each of the other primary chromosome staining was performed in phos- clades that, together with the subfamily Sigmo- phate-buffered Giemsa (pH=6.8). Ten metha- dontinae comprise the family Cricetidae (Steppan phases spreads were counted for each specimen. et al. 2004). A fragment of 1186 bp of the first exon Fundamental Numbers (FN) refers only to auto- of the nuclear gene Interphotoreceptor Retinoid somes (Patton 1967). Binding Protein (hereafter IRBP) was used as evidence for the phylogenetic analysis. For some specimens a shorter fragment was used. Specimens Morphology included in the phylogenetic analysis, and source and length of their sequences are listed in Table 1. Dental description followed Reig (1977) and was IRBP sequences acquired here were amplified and based on three specimens (IEEUACH 1063, 1745, sequenced in one or two fragments using the 3851) from Fundo San Martin, Valdivia Province primers A1–F1 and E1–D reported by Jansa and (see below). Penis description followed Hopper and Voss (2000) and following a protocol presented by Musser (1964) and was based on excised phalli these authors. Negative controls were included in from four formalin-preserved specimens all experiments. Amplification products were se- (IEEUACH 5427, 5461, 5463 5464). A ventral cut quenced in both directions with the amplification exposed the urethral processes and the anterior primers and dye-labeled nucleotides (Big Dye, mounds of the baculum; a second cut exposed the ARTICLE IN PRESS New data of Pearsonomys annectens (Sigmodontinae) 3 Table 1. Species included in the phylogenetic analysis (1–32: ingroup; 33–39: outgroup). GENBANK accession numbers are provided for all sequences; for those shorter than 1186 bp their length is indicated between parentheses. For the sequences acquired in this study catalog numbers and the localities of the corresponding specimens are provided. An * indicates sequences whose first 747 bp (approximately) were acquired in D’Elı´a (2003) and completed here. An # indicates sequences completely acquired here. Voucher specimens for the individuals whose sequences were partially or completely acquired in this study are catalogued in the following collections: Argentina, Coleccio´n de Mamı´feros del Centro Nacional Patago´nico (CNP); Chile, Coleccio´nde Mamı´feros del Instituto de Ecologı´a y Evolucio´n, Universidad Austral de Chile (IEEUACH); United States, Museum of Vertebrate Zoology, University of California, Berkeley (MVZ). Species Catalog number Locality Source 1 Abrothrix andinus AY277418 (747) 2 Abrothrix longipilis AY277419 (747) 3 Abrothrix longipilis AY163577 4 Abrothrix jelskii AY277442 (747) 5 Abrothrix olivaceus AY277420 (747) 6 Abrothrix olivaceus CNP 813 Argentina, Neuque´n, Volca´n ÃAY277421 Tromen 7 Akodon azarae AY163578 8 Bibimys chacoensis CNP 756 Argentina, Chaco, Dpto. ÃAY277435 Bermejo, Cancha Larga S2710403700 W5814305100 (1078) 9 Blarinomys breviceps CIT 1391 ÃAY277437 10 Calomys lepidus AY163580 11 Chelemys macronyx CNP 753 Argentina, Neuque´n, Laguna DQ069311 Epulafquen S3614901000 W7110404700 # (746) 12 Chelemys macronyx MVZ 155800 AY277441 (747) 13 Delomys sublineatus AY163582 14 Geoxus valdivianus CNP 812 Argentina, Neuque´n, Caviahue ÃAY277448 S3715302500 W7110304700 15 Holochilus chacarius AY163586 16 Irenomys tarsalis AY163587 17 Juliomys pictipes AY163588 18 Nectomys squamipes AY163598 19 Notiomys edwardsii AY163602 20 Oligoryzomys nigripes AY163612 21 Oryzomys AY163621 megacephalus 22 Oxymycterus nasutus MVZ 182701 Uruguay, Maldonado, El ÃAY277468 Pen˜asco 23 Pearsonomys IEEUACH #AY851749 annectens 7056 (1127) 24 Phyllotis xanthopygus
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