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CITATION

READS

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196

5

authors, including:

105 PUBLICATIONS 892 CITATIONS SEE PROFILE

59 PUBLICATIONS 409 CITATIONS SEE PROFILE

See discussions, stats, and author profiles for this publication at: <a href=https://www.researchgate.net/publication/236903904 Phylogenetic position and morphology of Abrothrix illutea Thomas, 1925, with comments on the incongruence between gene trees of Abrothrix (Rodentia, Cricetidae) and their implicati... Article · January 2011 CITATION READS 1 196 5 authors , including: Pablo Teta Centro Nacional Patagonico 105 PUBLICATIONS 892 CITATIONS SEE PROFILE Pablo Jayat Universidad Nacional de Chilecito 59 PUBLICATIONS 409 CITATIONS SEE PROFILE Guillermo D'Elía Universidad Austral de Chile 112 PUBLICATIONS 1,266 CITATIONS SEE PROFILE All in-text references underlined in blue are linked to publications on ResearchGate, letting you access and read them immediately. Available from: Guillermo D'Elía Retrieved on: 17 June 2016 " id="pdf-obj-0-56" src="pdf-obj-0-56.jpg">

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Zoosyst. Evol. 87 (2) 2011, 227 241 / DOI 10.1002/zoos.201100005

Zoosyst. Evol. 87 (2) 2011, 227 – 241 / DOI 10.1002/zoos.201100005 Phylogenetic position and morphology ofSmith et al. 2001; Rodr guez et al. 2008; Lessa et al. 2010; Feijoo et al. 2010 ), is still incomplete. One of the least known species of Abrothrix is the Argentinean endemic A. illutea Thomas, 1925. This mouse, poorly represented in museum collections, has a small geographic distribution in northwestern Argentina (Teta et al. 2006). The generic placement of this species Hornos islands and adjacent lowlands in Patagonia; has shifted in accordance with the status given to Abro- Musser & Carleton 2005 with modifications). The knowledge of this genus, despite increased data in re- thrix , from a member of the genus Akodon when Abro- thrix was considered a synonym (Ellerman 1941) or as * Corresponding author, e-mail: guille.delia@gmail.com # 2011 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim " id="pdf-obj-1-8" src="pdf-obj-1-8.jpg">
Zoosyst. Evol. 87 (2) 2011, 227 – 241 / DOI 10.1002/zoos.201100005 Phylogenetic position and morphology ofSmith et al. 2001; Rodr guez et al. 2008; Lessa et al. 2010; Feijoo et al. 2010 ), is still incomplete. One of the least known species of Abrothrix is the Argentinean endemic A. illutea Thomas, 1925. This mouse, poorly represented in museum collections, has a small geographic distribution in northwestern Argentina (Teta et al. 2006). The generic placement of this species Hornos islands and adjacent lowlands in Patagonia; has shifted in accordance with the status given to Abro- Musser & Carleton 2005 with modifications). The knowledge of this genus, despite increased data in re- thrix , from a member of the genus Akodon when Abro- thrix was considered a synonym (Ellerman 1941) or as * Corresponding author, e-mail: guille.delia@gmail.com # 2011 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim " id="pdf-obj-1-10" src="pdf-obj-1-10.jpg">

Phylogenetic position and morphology of Abrothrix illutea Thomas, 1925, with comments on the incongruence between gene trees of Abrothrix (Rodentia, Cricetidae) and their implications for the delimitation of the genus

Pablo Teta 1 , Guillermo D’El a * , 2 , Ulyses F. J. Pardi as 3 , J. Pablo Jayat 4 and Pablo E. Ort z 5

  • 1 Museo Argentino de Ciencias Naturales “Bernardino Rivadavia”, Avenida ngel Gallardo 470, (C1405DJR) Buenos Aires, Argentina

  • 2 Instituto de Ciencias de la Tierra Evoluci n, Universidad Austral de Chile, Campus Isla Teja s/n, casilla 567, Valdivia, Chile

  • 3 Unidad de Investigaci n Diversidad, Sistem tica y Evoluci n, Centro Nacional Patag nico, Casilla de Correo 128, 9120 Puerto Madryn, Chubut, Argentina

  • 4 Laboratorio de Investigaciones Ecol gicas de las Yungas (LIEY), Facultad de Ciencias Naturales e Instituto Miguel Lillo, Universidad Nacional de Tucum n, Casilla de Correo 34, 4107 Yerba Buena, Tucum n, Argentina

  • 5 C tedra de Paleontolog a, Facultad de Ciencias Naturales e Instituto Miguel Lillo, Universidad Nacional de Tucum n, Miguel Lillo 205, 4000 San Miguel de Tucum n, Tucum n, Argentina

Received 29 August 2010 Accepted 7 February 2011 Published 23 September 2011

Key Words

Abrotrichini

Argentina

gene tree

lineage sorting

Sigmodontinae

species tree

Abstract

Abrothrix illutea Thomas, 1925 is a poorly known sigmodontine rodent of the recently diagnosed tribe Abrotrichini. Here we report new locality records in northwestern Ar- gentina, describe in detail several morphological traits of this mouse and provide an emended diagnosis for the species. Abrothrix illutea is characterized by a unique com- bination of characters, including uniform dark olive-grey dorsal coloration; skull ro- bust, with the interorbital constriction hourglass shaped and smoothly rounded; muzzle elongate; nasals and premaxillae slightly projected anterior to the incisors; palate end- ing at about the plane of the last upper molars; upper incisors broad and orthodont; molars hypsodont, M1 with anteroloph and mesoloph well developed; stomach unilocu- lar-hemiglandular; terminal crater of the phallus ventrally directed; cartilaginous distal baculum reduced, composed of two lateral digits. These morphological features are dis- cussed at the light of phylogenetic analyses of nuclear and mitochondrial DNA se- quences. Results of these analyses, the first to include A. illutea, are incongruent be- tween loci. It is suggested that the nuclear based tree better reflects the pattern of morphological variation, and presumably the species tree, than the mitochondrial tree. These aspects are discussed in reference to the contents and limits of the genus Abro- thrix.

Introduction

The tribe Abrotrichini represents a small clade within

the radiation of the Sigmodontinae (D’El a et al. 2007). Five genera are recognized within this recently diag- nosed tribe. Abrothrix, with eight living species, is the most diverse and widely distributed (along the Andes

from central

Peru to Tierra del Fuego and Cabo de

cent years, especially for southern latitude populations (e.g., Smith et al. 2001; Rodr guez et al. 2008; Lessa et al. 2010; Feijoo et al. 2010), is still incomplete. One of the least known species of Abrothrix is the Argentinean endemic A. illutea Thomas, 1925. This mouse, poorly represented in museum collections, has a small geographic distribution in northwestern Argentina (Teta et al. 2006). The generic placement of this species

Hornos islands and adjacent lowlands in Patagonia; has shifted in accordance with the status given to Abro-

Musser & Carleton 2005 with modifications). The knowledge of this genus, despite increased data in re-

thrix, from a member of the genus Akodon when Abro- thrix was considered a synonym (Ellerman 1941) or as

* Corresponding author, e-mail: guille.delia@gmail.com

# 2011 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim

Zoosyst. Evol. 87 (2) 2011, 227 – 241 / DOI 10.1002/zoos.201100005 Phylogenetic position and morphology ofSmith et al. 2001; Rodr guez et al. 2008; Lessa et al. 2010; Feijoo et al. 2010 ), is still incomplete. One of the least known species of Abrothrix is the Argentinean endemic A. illutea Thomas, 1925. This mouse, poorly represented in museum collections, has a small geographic distribution in northwestern Argentina (Teta et al. 2006). The generic placement of this species Hornos islands and adjacent lowlands in Patagonia; has shifted in accordance with the status given to Abro- Musser & Carleton 2005 with modifications). The knowledge of this genus, despite increased data in re- thrix , from a member of the genus Akodon when Abro- thrix was considered a synonym (Ellerman 1941) or as * Corresponding author, e-mail: guille.delia@gmail.com # 2011 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim " id="pdf-obj-1-157" src="pdf-obj-1-157.jpg">

228

Teta, P. et al.: Systematics of Abrothrix (Rodentia, Cricetidae)

a subgenus Akodon (Abrothrix) (Cabrera 1961; Reig 1987). In contrast, Osgood (1943) suspected that illutea was wrongly allocated to Abrothrix and suggested a possible relationship with Hypsimys [= Akodon (Hypsi- mys); see Reig 1987; D’El a 2003]. Liascovich et al. (1989), on the basis of morphology and karyotype, re- inforced the inclusion of this species within Abrothrix, as Akodon (Abrothrix) illutea. Most of the morphologi- cal features reported by Liascovich et al. (1989: 389) were basically transcribed from Reig (1987: 367) and apply to the genus Abrothrix as a whole. Several strik- ing differences between A. illutea and other members of the genus were not indicated and deserve further in- spection. Phylogenetic relationships among some mem- bers of Abrothrix (i.e, andina, jelskii, lanosa, longipilis, olivacea) were studied by Smith & Patton (1999), D’El a (2003), Rodr guez et al. (2008), and Feijoo et al. (2010) through the analysis of nuclear and/or mitochon- drial DNA sequences. Notwithstanding, no published phylogenetic study has included representatives of A. il- lutea until now. The aim of this study is to provide detailed descrip- tions of skeletal and soft (e.g., penis, stomach) anatomy of A. illutea, and to test its phylogenetic position based on nuclear and mitochondrial DNA sequences. An emended diagnosis is provided, and the distribution of A. illutea is redefined based on new locality records, museum specimens, and literature. Finally, we make some comments about the phylogeny of the genus, especially about the incongruence reflected by the tree genes of different loci.

Materials and methods

Studied specimens. The specimens included in this work were ob- tained from systematic collections, newly trapped with Sherman-like traps baited with peanut butter and animal fat, or recovered from owl pellets (Appendix 1 and 2). The former are housed at the following Argentinean institutions: Colecci n de Mam feros del Centro Nacio- nal Patag nico (CNP), Puerto Madryn, Chubut; Colecci n de Mam - feros Lillo (CML), Universidad Nacional de Tucum n, Tucum n; Mu- seo Argentino de Ciencias Naturales “Bernardino Rivadavia” (MACN), Buenos Aires. Some newly captured specimens are num- bered in the field catalog of Jorge Pablo Jayat (JPJ). Owl pellet sam- ples are deposited in the pellet collection of Pablo E. Ortiz (PEO-e). We obtained additional records from the literature. Morphologic analyses. Standard external measures were recorded from specimen tags or field catalogs: TL total length; T tail length; HF hind foot length (including the claw); E ear length; W weight. The following skull measurements were recorded with calipers following Myers et al. (1990): MLS maximum length of the skull; CIL condyloincisive length; NL nasal length; RL rostral length; RW2 mid rostral width; ZP zygomatic plate depth; IOC interorbital constriction; ZB zygomatic breadth; BB braincase breadth; DL diastema length; MTRL maxillary toothrow length; IFL incisive foramina length; OCW occipital condyle width; ML mandible length; mTRL mandibular tooth- row length. Nomenclature used to describe the skull and its structures follows Voss (1988) and Wahlert (1985) and that of the molars corre- sponds to Reig (1977). Stomachs were dissected under a stereomicro- scope; morphological analysis and description follow Carleton (1973).

The glans penis was removed from one specimen after fixation in 10 % formalin. Terminology of the anatomical description follows Hooper & Musser (1964) and Spotorno (1992). The glans penis was cleared in 1 % KOH and stained with alcian blue and alizarine red for cartilaginous and osseous tissues, respectively (Wassersug 1976). Age classes were defined according the criteria explained by Myers (1989). Finally, for comparative purposes we studied series of other Abrothrix species (see Appendix 2). Phylogenetic analyses. Phylogenetic analyses were based on complete cytochrome-b gene (cyt b) sequences and a fragment of 759 bp of the first exon of the nuclear Interphotoreceptor Retinoid Binding Protein gene (IRBP). Taxonomic coverage is broad; we expanded the data set analyzed by Feijoo et al. (2010) to include A. illutea (CNP 1489; Ap- pendix 1) in both matrices and Chelemys megalonyx (GenBank:

EU091259) in the IRBP data set. The final matrix included 6 of the 8 species of Abrothrix (sensu Musser & Carleton 2005, as modified by Rodr guez et al. 2008) and all species belonging to the remaining ab- rotrichine genera. Representatives of the remaining sigmodontine tribes and some of the sigmodontine genera with unclear phylogenetic relationships (see D’El a et al. 2006a) were used to conform the out- group. The cyt b matrix includes a total of 34 sequences and that of the IRBP 25 sequences. Sequence acquisition was as follows. The cyt b sequences were acquired, as described in Pardi as et al. (2003), in two fragments using primers MVZ 05 MVZ 16 and MVZ 103 MVZ 14. IRPB sequences were obtained using primers A1 and F1 as in D’El a (2003) following the conditions delineated by Jansa & Voss (2000). Amplification products were purified and sequenced by Macrogen Inc., Korea. Sequences were deposited in GenBank (HQ189528, HQ189529). Sequence alignment was done with Clustal X (Thompson et al. 1997) using the default parameter values. Sigmo- dontinae cyt b sequences differ in length (see D’El a et al. 2003: 353), with differences residing in the position of the terminal stop codon. Therefore, we limited the analysis to the first 1137 bases of cyt b. Due to the different taxonomic coverage reached for each gene, aligned sequences of both regions were separately analyzed by Max- imum Parsimony (MP) (Farris 1982) and Bayesian (Rannala & Yang 1996) analyses. A third set of analyses was conducted with a matrix combining both genes, totalizing 2415 characters. Here, the matrix was composed of 22 terminal taxa including the 13 abrotrichines that cover all species analyzed in both solo analyses. Chimeras were con- structed for only two species of the ingroup (Chelemys megalonyx and Pearsonomys annectens) and for two taxa composing the out- group (Akodon and Reithrodon). We conducted an incongruence- length difference test (ILD; Farris et al. 1994) in PAUP * 4 (Swofford 2000) with 10 thousand replicates to assess the extent of character conflict between both gene partitions (for an examination of the inter- pretation of the ILD as a test of data combinability see Baker & Lut- zoni 2002). MP analyses were done in PAUP * 4 with characters treat- ed as unordered and equally weighted, 200 replicates of heuristic search with random addition of sequences, and tree bisection recon- nection (TBR) branch swapping. Relative support of the recovered clades was assessed by performing 1000 jackknife (JK) replications each with 3 replicates of random addition of sequences, TBR branch swapping, and a deletion of 33 % of the characters. For the combined analysis we also calculated partitioned Bremer support values (PBS; a measure of the positive or negative contribution of each partition to the character support for a particular node in the simultaneous analy- sis; Baker & DeSalle 1997). Bayesian analyses were performed with MrBayes 3.1 (Ronquist & Huelsenbeck 2003), by means of two inde- pendent runs with 3 heated and 1 cold Markov chains each. A model with six categories of base substitution, a gamma-distributed rate parameter, and a proportion of invariant sites was specified; all model parameters were estimated in MrBayes. Uniform interval priors were assumed for all parameters except base composition and GTR param- eters, which assumed a Dirichlet process prior. Runs were allowed to proceed for 10 million generations; for each chain trees were sampled every 1 thousand generations. Log-likelihood values against generation time were plotted to check that the runs converged on a stable log-likelihood value. The first 25 % of the sampled trees were discarded as burn-in; the remaining trees were used to compute a

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Zoosyst. Evol. 87 (2) 2011, 227 241

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50 % majority rule consensus tree and obtain posterior probability (PP) estimates for each clade.

Results

Order Rodentia Bowdich, 1821 Family Cricetidae Fischer, 1817 Subfamily Sigmodontinae Wagner, 1843 Tribe Abrotrichini D’El a, Pardi as, Teta & Patton, 2007

Abrothrix illutea Thomas, 1925

Type locality. Abrothrix illutea was described by Thomas (1925: 582)

with type locality in “Tucuman Province

... (Thomas 1925: 583). Later, Thomas (1929: 41) modified the terra typica of this species indicating “Mr. Morgensen (sic) has captured four further specimens [of illutea] on Aconquija at 3000 4000

Concepcion, 400 m.”

metres, and I think there is no doubt that the original specimen [the

type of illutea] was also obtained at that altitude

...

and that it does

not occur at the low-level Concepcion at all.” In addition, Thomas wrote in the original label of the type specimen “No doubt Aconquija, 3 4,000 m.” (Liascovich et al. 1989: 388). Thomas (1929: 41) argued that Concepci n was unlikely the type locality of A. illutea because in his opinion its general landscape of humid subtropical forest (the Yungas) was an “unsuitable” environment for a species of Abrothrix. However, as Liascovich et al. (1989) first demonstrated, several re- cording localities for A. illutea are in areas of the Yungas (see below). When Stewart Shipton collected the type of illutea, Concepci n was already a relatively large city, yet was surely surrounded by patches of Yungas. Then, according to the available information the type lo- cality of A. illutea should be fixed, as Thomas (1925:583) first stated, in Concepci n (27.34 S, 65.59 W), in the Chicligasta Department, Tucum n Province, Argentina.

Emended diagnosis. A member of the genus Abrothrix of the Sigmodontinae, characterized by a unique combi- nation of characters including: dorsal coloration uni- form dark olive-grey; ventral surface ashy grey, with small patches of white hairs under the chin and in the inguinal area; skull robust, with the interorbital con- striction hourglass-shaped and the braincase well rounded; rostrum elongate; nasals and premaxillae slightly projected anterior to the incisors; naso-frontal suture U-shape, not surpassing posteriorly the plane de- fined by the lacrimals; zygomatic plate slender with the anterior margin straight and slightly oriented backward; palate ending at about the plane defined by the poster- ior faces of the last molars; upper incisors broad, yel- lowish, ungrooved and mostly orthodont; molars hypso-

2008a). The presence of A. illutea at elevations of 3000 m and 4000 m mentioned by Thomas (1929) for the Aconquija range is, as discussed in the Type local- ity heading, doubtful given the uncertainties about the exact place of collection. In addition, there is no other record of A. illutea over 2500 m of elevation (cf. Jayat et al. 2008a). The presence of A. illutea in Jujuy pro- vince is not yet well documented. Olrog (1979) re- corded the species from Cerro Calilegua (Duraznillar sensu Heinonen & Bosso [1994]), Jujuy, but those spe- cimens once housed at the CML are lost (see D az 1999). In the most recent treatment of the Jujuy mam- mal fauna (D az & Barquez 2007), A. illutea was not listed. Abrothrix illutea is mainly associated to the “Bosque Montano” (Brown et al. 2001), the upper ele- vational belt of the Yungas forest, where “aliso” (Alnus acuminata) and “pino del cerro” (Podocarpus parlato- rei) are the dominant tree species. However, a few re- cords of A. illutea are known from the “Selva Montana” between 700 and 1000 m. At its upper elevational limit, it inhabits tall grass and brushy habitats along streams (Fonollat 1984). Despite its small range, A. illutea has been found in sympatry with several other sigmodon- tines, including Akodon caenosus, A. simulator, A. spe- gazzinii, Andinomys edax, Calomys fecundus, C. muscu- linus, Graomys chacoensis, Holochilus chacarius, Necromys lactens, N. lasiurus, Oligoryzomys cf. flaves- cens, Oligoryzomys sp., Oxymycterus paramensis, O. wayku, Phyllotis anitae, and P. osilae. At least two other species of sigmodontine rodents, Oxymycterus wayku and Phyllotis anitae (Jayat et al. 2007, 2008b), three caviomorph (Ctenomys tuconax, C. tucumanus, C. viperinus), one fish (Astyanax puka), one amphibian (Gastrotheca gracilis), several mollusks (e.g. Adelopoma tucma, Scolodonta semperi, Epiphrag- mophora argentina, E. birabeni, E. hemiclausa, E. hiero- nymi, E. parodizi, E. puella), and one plant (Dunalia lorentzii) are endemic to the southern portion of the Yungas forests, sharing a similar distribution pattern with A. illutea (Brown 1986; Lavilla & Manzano 1995; Lavilla et al. 2000; Barquez et al. 2006; Mirande et al. 2007). Although this geographical sector has been char- acterized as an impoverished version of the northern Yungas, climatic and isolation history have been men- tioned as clues in determining its biological composi- tional pattern (Brown et al. 2001; Ojeda et al. 2008).

dont, with tendency to develop enamel islands in young Fossil record. Fossil remains of A. illutea have been

individuals; M1 without anteromedian flexus in adult individuals; parastyle and mesoloph well developed, the latter usually fused to the paracone; stomach unilocu- lar-hemiglandular; terminal crater of the phallus ven-

found in an exceptionally rich rodent assemblage of Middle-Upper Pleistocene age at La Angostura (Tafi Valley, Tucum n; Ortiz & Pardi as 2001). Other sigmo- dontines found in the same fossil assemblage include

trally directed; cartilaginous distal baculum reduced, Akodon spegazzinii, Andinomys edax, Calomys cf.

composed of two lateral digits.

Distribution and habitats. Corroborated records of Ab- rothrix illutea (Fig. 1) come from Catamarca and Tucu-

  • m n provinces, northern Argentina, between 700 and

2500 m (Fonollat 1984; Liascovich et al. 1989; Capl- lonch et al. 1997; Mares et al. 1997; Jayat et al. 2006,

C. laucha-musculinus, Necromys cf. N. lactens, Neo- tomys ebriosus, Oligoryzomys cf. O. flavescens, Oligory- zomys sp., Oxymycterus sp., Phyllotis sp., Reithrodon

auritus, and the extinct Tafimys powelli.

Description. Abrothrix illutea is one of the largest re- presentatives of the genus, characterized by its almost

# 2011 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim

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Teta, P. et al.: Systematics of Abrothrix (Rodentia, Cricetidae)

230 Teta, P. et al.: Systematics of Abrothrix (Rodentia, Cricetidae) Figure 1. Locality records for Abrothrix

Figure 1. Locality records for Abrothrix illutea (the numbers correspond to those of the Appendix 1) in the provinces of Catamarca and Tucum n, Argentina.

uniform coloration and a tail of about 65 67 % of head and body length combined (Table 1). The dorsal pelage is soft, woolly, and dense; it is composed almost en- tirely of grey hairs with ochraceous tips, giving a gen- eral dark olive-grey coloration. Dorsal hairs are 10 to 12 mm long, being longer in the mid rump region; guard hairs are slightly longer, projecting 2 3 mm be- yond the fur in the rump. The rhinarium and the lips are covered by short, thin, much more ochraceous deli- cate hairs. Mystacial vibrissae are abundant, bright, and moderately long, reaching the anterior portion of the ears. Interamal and submental vibrissae are short and white. Ears are short, rounded, and dark grey in colora- tion, with the margin of the inner surface covered with short and thin black hairs. Flanks are similar to the dor- sum but slightly clearer. Ventrally the colour is ashy grey, with each individual hair dark grey at their bases and white at the tip. There is a small patch or some

white hairs under the chin

and in the inguinal area.

Fore and hindfoot are covered by short brown hairs dor-

sally. Ungual tufts are whitish, shorter than the claws in

the forefeet and longer than claws on the hindfeet. Plantar pads are six, including a nearly oval and large thenar pad, a rounded hypothenar pad, and four rounded to nearly ovoid interdigital pads. The tail is slightly bicolored, dark grey dorsally and ashy grey in the venter. The skull is robust and has a moderately elongated and broad rostrum, inflated sinuses, gracile zygomatic arches, and well inflated, rounded braincase (Fig. 2). The nasals are broad, divergent forward and anteriorly expanded, covering almost entirely the rostrum in dor- sal view (Fig. 3a). The naso-frontal suture is U-shaped and does not extend beyond the plane defined by the posteriormost point of the lachrymals (Fig. 3a). The in- terorbital constriction is hourglass-shaped, smoothly rounded, without square edges or beads. The interparie- tal is small but clearly visible in dorsal view. The occi- pital region is inflated and without ridges. The gnathic process is almost absent and nasals and premaxillae ex- tend slightly beyond the incisors front plane (Fig. 3b). The nasolacrimal capsules are well developed, contri-

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Table 1. External and craniodental measurements of Abrothrix illutea from Tucum n, Argentina (age classes follows Myers 1989).

 

Age class 1 ( N ¼ 3)

Age class 2 ( N ¼ 5)

Age class 3 ( N ¼ 2)

Age class 5 ( N ¼ 2 3 a )

mean sd

range

mean sd

range

mean sd

range

mean sd

range

TL

190 3.21

188 194

195 6.93

184 202

197

192 202

209

206 212

T

83 2.00

81 85

84 5.02

76 88

83

82 83

83

80 85

HF

28 0.58

28 29

29 0.89

28 30

30

29 30

29

27 30

E

19 1.15

18 20

21 1.30

19 22

20

19 20

21

20 21

W

33.8 1.04

33.0 35.0

38.9 4.71

33.5 45.5

39.5

37.0 42.0

53.8

52.5 55.0

MLS

29.14 0.25

28.96 29.42

29.92 0.75

29.00 31.10

29.50

28.92 30.08

31.75 0.30

31.44 32.04

CIL

26.50 0.34

26.16 26.84

27.15 1.32

24.94 28.48

27.29

26.62 27.96

29.40 0.36

29.00 29.68

RL

11.07 0.30

10.82 11.40

11.46 0.34

11.02 11.92

11.28

11.04 11.52

12.06 0.10

12.00 12.18

ZB

14.36 0.12

14.28 14.50

14.68 0.39

14.16 15.08

14.60

14.32 14.88

15.37 0.27

15.10 15.64

BB

13.46 0.09

13.38 13.56

13.49 0.21

13.26 13.74

13.31

13.06 13.56

13.58 0.12

13.48 13.72

IOC

5.12 0.03

5.10 5.16

5.13 0.12

5.00 5.28

5.01

5.00 5.02

5.09 0.23

4.86 5.32

MTRL 4.45 0.05

4.40 4.50

4.56 0.13

4.42 4.78

4.27

4.20 4.34

4.69 0.06

4.64 4.76

NL

10.46 0.14

10.30 10.58

10.86 0.37

10.30 11.24

10.74

10.26 11.22

11.79 0.32

11.48 12.12

RW2

5.55 0.19

5.34 5.70

5.67 0.10

5.54 5.82

5.66

5.42 5.90

5.79 0.17

5.62 5.96

DL

7.38 0.15

7.22 7.52

7.79 0.35

7.40 8.30

7.70

7.52 7.88

8.37 0.11

8.24 8.44

IFL

6.96 0.11

6.84 7.04

7.23 0.46

6.92 8.04

7.12

6.94 7.30

7.53 0.22

7.30 7.74

OCW

7.21 0.08

7.12 7.28

7.27 0.08

7.16 7.36

7.30

7.18 7.42

7.49 0.16

7.30 7.60

ZP

2.15 0.08

2.06 2.20

2.12 0.04

2.10 2.20

2.25

2.10 2.40

2.43 0.05

2.38 2.48

ML

14.87 0.33

14.50 15.12

15.33 0.48

14.80 16.10

15.47

15.02 15.92

16.22 0.11

16.14 16.34

mTRL 4.60 0.06

4.54 4.66

4.71 0.08

4.62 4.82

4.69

4.66 4.72

4.85 0.12

4.72 4.96

a Measurements with two values lack standard deviation.

Zoosyst. Evol. 87 (2) 2011, 227 – 241 231 Table 1. External and craniodental measurements of

buting to the broadened aspect of the rostrum (Fig. 3a). The zygomatic plate is slender (Fig. 3b), with its ante- rior margin straight and slightly slanted backward (Fig. 3c), while the zygomatic notch is shallow and lat- erally compressed. The zygomatic plate has a reduced but noticeable free upper border (Fig. 3c), uncommon within Abrothrix representatives. The masseteric tuber- cle is inconspicuous but present. The zygomatic arches are weakly flared laterally; the jugal is large and the maxillary and squamosal processes of the zygoma are not in contact. The parietals have deep lateral expan- sions with crenulate suture (Fig. 3d). The hamular pro- cess of squamosal is long and narrow and distally spa- tulate (Fig. 3d). The subsquamosal fenestra and the postglenoid foramen are large, with the former larger than the latter (Fig. 3d). The mastoid has a conspicuous fenestra, not reaching the exoccipital contact area. The incisive foramina are long and broad, extending poster- iorly to the M1 protoflexus (Fig. 3e). The medial pro- cess of the incisive foramina occupies nearly of 3/4 of the opening. The palate is short, wide (sensu Hershko- vitz 1962) and flat. Molar rows are somewhat divergent anteriorly. The mesopterygoid fossa is relatively broad with nearly subparallel or slightly divergent lateral mar- gins (Fig. 3f); its anterior border is rounded or squared,

Figure 2. Dorsal (upper left), ventral (upper right) and lateral (center) views of the skull, and labial view of mandible (lower center) of Abrothrix illutea (JPJ 481). Scale ¼ 5 mm.

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reaching the posterior plane of M3. The mesopterygoid roof is totally ossified anteriorly but elongate spheno-

a sphenofrontal foramen. At the junction of the trough for the masticatory-buccinator nerve and the squamosal-

palatine vacuities are present medially (Fig. 3f). The alisphenoid groove a large perforation is consistently

parapterygoid fossae are shallow with straight and pos- teriorly divergent borders, subequals in size to the me- sopterygoid fossa (Fig. 3f). The alisphenoid strut is ab- sent. Cranial foramina associated with the cephalic arterial pattern and other osteological traits of the ali- sphenoid region include a large foramen ovale, a re- duced anterior opening of the alisphenoid canal, a broad trough for the masticatory-buccinator nerve, and a squamosal-alisphenoid groove connecting anteriorly to

present. The periotic is attached to the skull by a devel- oped tegmen tympani, which overlaps the posterior sus- pensory process of the squamosal. The auditory bullae are relatively large and have short Eustachian tubes. The manubrium of the malleus is large and distally ex- panded; a well developed orbicular apophysis is present (“parallel” type, according to Carleton [1980]). The mandible is slender and delicate (Fig. 2). The masseteric crests are moderately to well developed, ex-

232 Teta, P. et al.: Systematics of Abrothrix (Rodentia, Cricetidae) reaching the posterior plane of M3.

Figure 3. Some anatomical traits of Abrothrix illutea: a. Rostrum in dorsal view (note the posteriormost extention of nasals [na] and the moderate development of nasolacrymal capsules [nc]); b. Rostrum in lateral view (note the nasal and premaxillary [pm] slightly surpassing the incisor profile and the orthodont condition denoted by the vertical incisive plane); c. Zygomatic plate [zp] in ventrolateral view (arrow indicates the upper free border); d. Temporal region with deep lateral parietal [lp] expansion and elongate hamular process [h] distally spatulated; e. Palate (note the posterior ends of the incisive foramina [fi] even with M1 protoflexus); f. Mesopterygoid region (note the anterior border of the mesopterygoid fossa even with the posterior faces of the M3 and the elongate sphenopalatine vacuities [sv]; g. Ascending ramus of the mandible in labial view (a moderate capsular projection [cp] lying below the coronoid process); h. Mandible in dorsal view (note the well developed retromolar shelf [rs]); i. Left M1 in occlusal view showing well developed parastyle [pa] and mesoloph [m].

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tending almost to the anterior border of m1. The ante- while glandular epithelium is mostly limited to the an-

rior point of the diastema is slightly above of the alveo- trum. The bordering fold bisects the stomach on a lar plane. The mental foramen is usually large and not straight line from the incisura angularis to a point op-

obvious in lateral view. The capsular projection of the lower incisive alveolus is small but clearly visible, and

posite it on the great curvature. The phallus is elongate, with a much reduced distal

is located just below of the base of the coronoid pro- baculum. The terminal crater is somewhat ventrally di-

cess (Fig. 3g). The coronoid process is elongated, slen- rected, not distally as usual, and showed a clear ventral der, sharp-pointed, and directed posteriorly. The angular invagination that occupies ca. 25 % of phallus (Fig. 4b).

and condyloid processes are thick and separated by a

The urethral processes are wide and long; with their in-

wide and semi-circular lunate notch. The posterior sertions near the middle of the os penis (Fig. 4c). The

point of the angular process is anterior to the posterior baculum is long and smoothly curved ventrally, and is

border of the condyloid process. A retromolar shelf is well developed in the inner part of the mandible be- tween the coronoid process and m3 (Fig. 3h).

relatively free within the crater. Its proximal osseous segment consists of a wide and nearly trapezoidal base. The distal baculum is elongated and narrow, with two

Upper incisors are broad, yellowish, ungrooved and small osseous lateral expansions. The cartilaginous ba-

orthodont, but some individuals are somewhat proo- dont. Molars are hypsodont (but not as much as Akodon budini), with terraced crowns and a tendency to devel-

culum is reduced (ca. 1 mm in length) and is composed of two lateral digits (Fig. 4c).

op enamel islands in young individuals. The M1 has an Comparisons. Most populations of Abrothrix jelskii are anteromedian flexus limited to a shallow groove only in characterized by a very striking coloration, with white

young individuals; in adults the procingulum is fan- shaped. The parastyle and mesoloph are well developed

spots at the base of ears, white (or grey) ventral parts that contrast with the dark dorsum, and a fulvous or

(Fig. 3i), the latter usually fused to the paracone (with ochraceous tint of the nose, hind feet, and tail. In addi-

a paralophule participation) in subadults and adults,

tion, the skull of A. jelskii has much larger tympanic

sometimes with an enamel island on the base of para- bullae, shorter rostrum, and nasals that at the posterior-

cone belonging to a close mesoflexus. The M2 has a

most point surpass the plane defined by the posterior-

well developed anteroloph, a mesoloph slightly shorter most point of the lacrimals.

than that of the M1, and a conspicuous mesofossetus between protocone and paracone. In young specimens a faint remnant of the posteroloph is also observed. M3

Abrothrix longipilis has a sharp contrast of the dorsal and ventral coloration (usually with a reddish tint to the dorsal midline) and the dorsal surface of the manus,

is small and cylindrical, with a half of the length of and the pes covered by short and thin white to cream

M2 or less, and with a conspicuous island even present in old specimens (age class 4). The anteromedian flexid of m1 is only present in young individuals; adults have

hairs. Abrothrix sanborni is smaller and uniformly blackish, without indication of countershading, and has smaller ears. Abrothrix lanosa has a more yellowish ol-

a procingulum subcircular in outline. A protostylid ive coloration, with the dorsal surface of the manus and

(plus labiolophulid, sensu Hershkovitz 1993) and the posterolophid are well developed. The mesolophid is

pes covered by short and thin white to cream hairs, and smaller ears. The skull of A. longipilis, A. lanosa, and

short and totally fused to the entoconid. A bulbous me- A. sanborni clearly depart from that of A. illutea. In

taconid is typical. The m2 has, in all the examined

these three species, the nasals largely surpass the ante-

young specimens, protostylid and posterolophid. The rior plane of the incisors and together with the premax-

m3 is a half of the length of m2 and has a simplified bilophodont pattern. The axial skeleton includes 13 ribs, 19 thoracicolum-

bar (dorsal) vertebrae, 3

sacral vertebrae, and 24 28

illae form a trumpet-like tube; the naso-frontal suture is much more acute and the posteriormost point of the na- sals surpass the plane defined by the posteriormost point of the lacrimals. In addition, the palate is longer,

caudal vertebrae (n ¼ 4). The neural spine on the sec- with

the anterior border of the mesopterygoid fossa

ond thoracic vertebra is much longer than the spines on the adjacent vertebrae (a widespread condition for sig- modontines, Voss 1988). The tuberculum of the first rib articulates with the transverse process of both the se-

well behind the posterior plane of M3. Abrothrix andina, A. hershkovitzi, and A. olivacea (including the forms markhami and xanthorhinus; see Pearson & Smith 1999; Rodr guez et al. 2008) are

venth cervical and the first thoracic vertebrae. The ster- much smaller (length of tail < 75 mm; length of hind-

num consists of an anterior manubrium, four sterne-

foot < 25 mm), with dorsal colorations that range from

brae, and a posterior xiphoid process. In the apendicular dark brown or olive brown to grey or pale ochraceous

skeleton, the humerus lacks an entepicondylar foramen; the supratrochlear fossa is perforated. The stomach is single chambered, unilocular-hemi- glandular (Fig. 4a). The incisura angularis is shallow,

(some forms have yellow or orange spots on nose, feet and tail). The skulls of A. andina and A. olivacea are smaller (condylo-incisive length < 27 mm), and much more delicate, with short and narrow rostra, moderately

scarcely extending beyond the esophageal opening. The extended nasals and premaxillae, slightly larger palates

corpus is spacious, with a broad fornix ventricularis. The cornified epithelium is coincident with the corpus,

(at least in A. olivacea), wider parapterygoid fossae, and proportionally larger tympanic bullae.

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234 Teta, P. et al.: Systematics of Abrothrix (Rodentia, Cricetidae) Figure 4. a. Morphology of theCarleton 1973; D’El a et al. 2007 ). The unilocular-hemiglandular pattern also characterizes the thrix species ( Spotorno 1992 ). Abrothrix jelskii and abrotrichine genera Geoxus , Notiomys , and Pearson- A. olivacea have shorter and narrower urethral pro- cesses. With some specific variations, the base of the baculum in other Abrothrix species is more laterally ex- panded and nearly triangular in shape. Abrothrix longi- pilis , A. lanosa , and A. sanborni lack of cartilaginous portion in the baculum, whereas A. andina , A. jelskii , and A. olivacea , have a much reduced distal baculum with two laterals and one medial digit. omys but in these taxa the glandular epithelium covers a small area of the corpus ( Carleton 1973; D’El a et al. 2006b; Pardi as et al. 2008). Phylogenetic position . Abrotrichine phylogenetic rela- tionships have been recently studied with mitochondrial and nuclear DNA sequences ( Smith & Patton 1999; D’El a 2003; D’El a et al. 2006b; Feijoo et al. 2010 ). Among the tribe Abrotrichini, the stomach morphol- Feijoo et al. (2010) analyzed separately both mitochon- ogy depicted by A. illutea is also found in other Abro- drial and nuclear loci, finding that the topologies de- # 2011 WILEY-VCH Verlag GmbH & Co. KGaA, Weinheim " id="pdf-obj-8-10" src="pdf-obj-8-10.jpg">

Figure 4. a. Morphology of the stomach (drawn from JPJ 1408); b c. Ventral and in- ternal view, respectively, of the glans penis (based on JPJ 141) of Abrothrix illutea. Ab- breviations: a antrum; b ventral lappet; bf bordering fold; c cartilaginous bacu- lum; ce cornified squamous epithelium; co corpus; e esophagus; ge glandular epithelium; ia incisura angularis; p py- lorus; pb proximal baculum; up ure- thral processes.

Both the elongate phallus and the ventral invagina- tion of the terminal crater are also found in other Abro-

thrix species (Carleton 1973; D’El a et al. 2007). The unilocular-hemiglandular pattern also characterizes the

thrix species (Spotorno 1992). Abrothrix jelskii and abrotrichine genera Geoxus, Notiomys, and Pearson-

A. olivacea have shorter and narrower urethral pro- cesses. With some specific variations, the base of the baculum in other Abrothrix species is more laterally ex- panded and nearly triangular in shape. Abrothrix longi- pilis, A. lanosa, and A. sanborni lack of cartilaginous portion in the baculum, whereas A. andina, A. jelskii, and A. olivacea, have a much reduced distal baculum with two laterals and one medial digit.

omys but in these taxa the glandular epithelium covers a small area of the corpus (Carleton 1973; D’El a et al. 2006b; Pardi as et al. 2008).

Phylogenetic position. Abrotrichine phylogenetic rela- tionships have been recently studied with mitochondrial and nuclear DNA sequences (Smith & Patton 1999; D’El a 2003; D’El a et al. 2006b; Feijoo et al. 2010).

Among the tribe Abrotrichini, the stomach morphol- Feijoo et al. (2010) analyzed separately both mitochon- ogy depicted by A. illutea is also found in other Abro- drial and nuclear loci, finding that the topologies de-

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235

rived from each gene are incongruent. Not surprisingly, here we found equivalent results.

ported (JK ¼ 97). Abrothrix andina, A. olivacea, A. jel- skii, and A. illutea form a clade, as in the IRBP

The IRBP matrix has 247 variable characters of topology, but one more weakly supported (JK < 50).

which 72 were parsimony informative. The analysis of Abrothrix andina and A. olivacea are sister species

this evidence generated 1188 shortest trees (328 steps;

(JK ¼ 100) while A. illutea and A. jelskii compose a

CI ¼ 0.826; RI ¼ 0.746), of which their strict consen- weakly supported clade (JK ¼ 63). These relationships

sus is shown in Figure 5a. Abrotrichine is strongly

are one of the three possible resolutions of the poly-

supported (JK ¼ 100). The dichotomy at the base of tomy found in the IBRP analysis. The difference be-

the abrotrichine clade leads to Abrothrix (JK ¼ 88) on

tween the IRBP trees (MP and BA) and the cyt b MP

one hand and to a clade (JK ¼ 72) composed of the topology is that in the latter A. lanosa is not sister to

remaining genera on the other. Within Abrothrix, A. il- A. longipilis, but to the ((A. andina, A. olivacea) (A. il-

lutea appears as part of a clade (JK ¼ 91) with a polytomy at its base formed by three main lineages:

lutea, A. jelskii)) clade. Moreover, the BA of the cyt b gene gives a third topology (Fig. 6b), with A. illutea

A. andina A. olivacea, A. jelskii, and A. illutea. The one of the four lineages that form part of a polytomy at

same topology was recovered in the Bayesian analysis (Fig. 5b).

the base of the Abrothrix clade (PP ¼ 1). Other mem- bers of this polytomy include A. andina A. olivacea

The cyt b matrix has 534 variable characters, of (PP ¼ 1), A. jelskii A. lanosa (PP ¼ 0.58), and A. long-

which 434 are parsimony informative. A total of five shortest trees (length ¼ 2265 steps; CI ¼ 0.361;

ipilis. Differences between the cyt b topologies are ne- cessarily due to the intrinsic differences of the em-

RI ¼ 0.520) were found; their strict consensus is repre- ployed methods and the use they made of the observed

sented in Figure 6a. Abrothrix appears strongly sup- variation. However, we note that these differences,

Zoosyst. Evol. 87 (2) 2011, 227 – 241 235 rived from each gene are incongruent. Not

Figure 5. Results of the phylogenetic analyses of IRBP gene sequences. Genbank accession numbers of analyzed sequences are included at terminal labels: a. Strict consensus tree of the 1182 most-parsimonious trees (length: 328; CI: 0.826; RI: 0.746) ob- tained in the maximum-parsimony analysis. Numbers on branches indicate Jackknife support values of the adjacent nodes. Only Jackknife values above 50 % are shown; b. Majority rule consensus obtained in the Bayesian analysis. Numbers on branches indi- cate posterior probability values of the adjacent nodes.

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236 Teta, P. et al.: Systematics of Abrothrix (Rodentia, Cricetidae) Figure 6. Results of the phylogenetic

Figure 6. Results of the phylogenetic analyses of cyt b gene sequences. Genbank accession numbers of analyzed sequences are in- cluded at terminal labels: a. Strict consensus tree of the 5 most-parsimonious trees (length: 2265; CI: 0.361; RI: 0.520) obtained in the maximum-parsimony analysis. Numbers on branches indicate Jackknife support values of the adjacent nodes. Only Jackknife values above 50 % are shown; b. Majority rule consensus obtained in the Bayesian analysis. Numbers on branches indicate posterior probability values of the adjacent nodes. (note that A. olivacea is recovered paraphyletic to A. markhami; see Rodriguez et al. 2008.)

while important, occur in areas of the trees that are weakly supported in both analyses. The ILD test show that the increment of homoplasy

resulting from combining

both gene partitions is not

significant (P ¼ 0.998). The combined matrix has 739 variable characters, being 476 parsimony informative. One shortest tree (length ¼ 2422 steps; CI ¼ 0.426; RI ¼ 0.393) was found (result no shown). Its topology is identical to that recovered in the MP cyt b solo ana- lysis; differences relate to support values. PBS values indicate that six of the twelve abrotrichine clades are supported by characters from both cyt b and IRBP genes (i.e., PBS values are positive for both genes). In addition, 2 clades, which are supported by cyt b charac- ters, are neither contradicted nor supported by the IRBP evidence (i.e., IRPB PBS ¼ 0). Three clades are sup-

ported by cyt b characters and contradicted by the evi- dence from IRBP (i.e., IRBP negative PBS values). Fi- nally, one clade of the combined analysis is supported only by IRBP characters (i.e., cyt b PBS ¼ 0). Overall, in the combined analysis cyt b characters support a total of eleven clades while IRBP characters support seven nodes. The topology found in the Bayesian analysis (re- sult not shown) also places A. illutea as sister to A. jel- skii (PP ¼ 1) and recovers, as the MP IRBP solo analy- sis, A. lanosa as sister to A. longipilis (PP ¼ 0.98).

Discussion

Abrothrix is a moderately diverse genus of Sigmodonti- nae, with eight species (Musser & Carleton 2005).

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Even though the genus is now relatively well known both in systematic terms and on different aspects of the natural history of some of its species, much remains to

Abrothrix by their notorious trumpet-like rostrum (see Osgood 1943; Feijoo et al. 2010). Therefore, it may be suggested that, even when Abro-

be studied. Ongoing phylogeographic studies (Lessa thrix as currently understood has been always recovered

et al. 2010; D’El a et al. in prep) indicate complex de- monophyletic and with good support (Smith & Patton

mographic histories for some of the species of Abro-

1999; D’El a 2003; D’El a et al. 2006b; Rodr guez

thrix and that the alpha diversity of the genus is not et al. 2008; Feijoo et al. 2010), a better classificatory

well reflected in current classification.

scheme would be one that reflect this large morphologi-

Similarly, phylogenetic relationships among species cal variation by recognizing more than one genus. As of Abrothrix are still uncertain because the topologies has been discussed by Patterson et al. (in press), the

inferred from two independent loci are partially incon- most reasonable alternatives to current classification

gruent. Regarding A. illutea, differences between both would be: 1) restricting Abrothrix to longipilis and re-

markers are not as large as those involving other spe- cies of Abrothrix (e.g., A. lanosa; see also Feijoo et al. 2010), as both loci when analyzed separately and to-

lated species (i.e., lanosa, sanborni), while expanding Chroeomys to, in addition to jelskii, encompass andina, olivacea and related species (i.e., hershkovitzii, markha-

gether tend to place A. illutea in a clade together with mi); or 2) while restricting Abrothrix to longipilis and

  • A. jelskii and a consistently recovered clade comprising

related forms and Chroeomys to jelskii (and perhaps il-

  • A. andina and A. olivacea. Feijoo et al. (2010) when lutea), erecting a new genus for andina, olivacea, and

discussing the locus-dependant phylogenetic position of related species. However, no classificatory innovation

  • A. lanosa, argued that the IRBP based topology was ea-

should be suggested until the phylogenetic relationships

sier to reconcile than the cyt b based tree with the pat- among

species of Abrothrix are better known (e.g., is

tern of morphologic variation displayed by the species lanosa sister to longipilis? Is illutea sister to jelskii? of Abrothrix as well as with their geographic distribu- What is the phylogenetic placement of sanborni?) and tion. In addition, these authors suggested that analysis the alpha-taxonomy refined (e.g., complex olivacea- of a third gene and/or the inclusion in future phyloge- xanthorhinus; is gossei a junior synonym of andina?).

netic analyses of additional species of Abrothrix, espe- In light of our phylogenetic results, it is clear that ana- cially A. illutea, would help in clarifying the nature of lysis of additional loci together with the analysis of

the differences between the cyt b and IRBP trees. Our results show that, unfortunately, the inclusion of A. illu- tea does not reconcile these alternative topologies. More in general, our results reinforce the idea that at least one of the genes is not correctly tracking the whole species tree of Abrothrix. This discrepancy between nuclear and mitochondrial topologies may have different causes, including slightly

morphological characters is mandatory. This task is cur- rently underway. However, we emphasize that if the main radiation of Abrothrix was explosive, the elucida- tion of the phylogenetic relationships of its species may be illusory, even with the inclusion of more loci, due to different lineage sorting among loci. Regarding the biogeographic history of Abrothrix, the still unclear phylogenetic relationships among the

different sampling between both loci, convergence in species of the genus preclude advancing a solid biogeo-

one locus among no related lineages due to selection graphic hypothesis accounting for its diversification, in- (e.g., Castoe et al. 2009), episodes of introgressive hy- cluding the origin of A. illutea. However, it is interest- bridization (e.g., Patton & Smith 1994), and hemiplasy ing to note that A. illutea is not sister to its

(Avise & Robinson 2008; see also Pamilo & Nei 1988). At this time, the nature of the discrepancy between the nuclear and mitochondrial gene trees of Abrothrix re- mains unclear.

geographically closest congener A. andina, but rather to A. jelskii, a species inhabiting high elevation Puna en- vironments. Closest known records of both species are about 300 km apart. The fossil record shows that in

This topological incongruence is relevant in terms of northwestern Argentina, a region with a complex topo-

the establishment of the contents of Abrothrix. In spite graphy and with a large environmental variability (Bur-

of its moderate diversity, Abrothrix is a morphologi- cally variable genus (Gyldenstolpe 1932; Osgood 1943; Patterson et al. 1984; Spotorno et al. 1990; Feijoo et al.

2010). As currently delimited (see D’El a 2003: 317) it includes jelskii, type species of Chroeomys (a divergent form both in terms of pelage and, with minor extent, skull and molar morphology). Similarly, longipilis, type species of Abrothrix, is remarkably divergent from the other species of the genus in having, together with

  • A. lanosa and A. sanborni, a unique phallic morphology

characterized by the absence of soft-bacular mounds,

kart et al. 1999), sigmodontine distributions have chan- ged extensively from the Pleistocene to the present (e.g., Ortiz et al. 2000; Ortiz & Pardi as 2001; Teta & Ortiz 2002). Further detailed phylogeographic studies, combined with prospecting for new fossils records, may clarify the processes that prompted the differentia- tion of A. illutea. Abrothrix and the remainder Abrotrichini have had a long history in connection with the Southern Cone of

South America, with a fossil record extending back into the Pliocene (Reig 1978). Abrothrix representatives are

(e.g., Gallardo et al. 1988; Spotorno et al. 1990; Spo- the most conspicuous elements in cold high-latitude

torno 1992). Similarly, longipilis, lanosa, and sanborni forest and steppes and are fundamental components of depart in skull morphology from the other species of several trophic chains (Osgood 1943; Mann 1978; Pat-

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terson et al. 1984, 1989, 1990; Meserve et al. 1988, 1991). Clearly, attending to these and other tacit rea- sons, the systematics of Abrothrix deserves a detailed and extensive fresh approach more than 180 years since Charles Darwin secured the first specimens.

Acknowledgements

We thank L. Ferr , E. Martin, D. Miotti, G. Namen, S. Sabat , and J. Torres Dowdall for helping us during field work. We are indebted to the following curators: R. Barquez (Colecci n de Mam feros Lillo, Tucum n, Argentina), D. Romero (Museo Municipal de Ciencias Nat- urales Lorenzo Scaglia), and D. Flores and O. Vaccaro (Museo Argen- tino de Ciencias Naturales “Bernardino Rivadavia”, Buenos Aires, Ar- gentina). B. Patterson, J. Patton and an anonymous reviewer provided valuable comments on an earlier version of this paper. Financial sup- port was provided by Consejo Nacional de Investigaciones Cient ficas y Tecnol gicas (CONICET) PIP 6179 (to UFJP), Fundaci n ProYun- gas, and FONDECYT 11070157 (to GD). This work was possible thanks to the institutional support of the Laboratorio de Investiga- ciones Ecol gicas de las Yungas and C tedra de Paleontolog a, Uni- versidad Nacional de Tucum n.

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2 km antes de La Heladera (Monteros, Tucum n), 27 3 0 3.8 00 S, 65 40 0 17.1 00 W. (12). 5 km S Las Higuerillas on Hwy 9, 1173 m (Pacl n, Catamarca), 27 57 0 28.5 00 S, 65 41 0 55.5 00 W. (20). Andalgal (Andalgal , Catamarca), 27 36 0 31.5 00 S, 66 19 0 10.5 00 W.

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Apeadero Militar Gral. Mu oz, km 41, Hwy 307 (Taf del Valle, Tucum n), 26 58 0 50.5 00 S, 65 39 0 50.9 00 W. (10). Aprox. 10 km S of Hualinchay, sobre el camino a Lara, 2300 m (Trancas, Tucum n), 26 19 0 20.2 00 S, 65 36 0 45.5 00 W. (1). Concepci n, 400 m (Chicligasta, Tucum n), 27 21 0 S, 65 35 0 W.

(15).

Cuesta del Clavillo, 5 km S of Banderita (Andalgal , Catamarca), 27 29 0 S, 66 06 0 W. (16). El Naranjal (Monteros, Tucum n), 27 02 0 S, 65 41 0 W. (11).

Horco Molle (Yerba Buena, Tucum n), 26 47 0 38.4 00 S, 65 18 0 57.6 00

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Horco Molle, Parque Biologico Sierra de San Javier, senda R o Las Piedras (Yerba Buena, Tucum n), 26 46 0 06 00 S, 65 19 0 66.4 00 W.

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La Angostura (Taf del Valle, Tucum n), 26 56 0 25.5 00 S,

65 42 0 10.5 00 W. (9).

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