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Mammalia, Didelphimorphia and Rodentia, Southwest of the province of Mendoza, .

Ulyses F. J. Pardiñas 1 Pablo Teta 2 Daniel Udrizar Sauthier 1

1 Centro Nacional Patagónico. Casilla de Correo 128. Puerto Madryn, Chubut, Argentina.

2 Departamento de Ecología, Genética y Evolución, Facultad de Ciencias Exactas y Naturales, Universidad de Buenos Aires, Avenida Intendente Güiraldes 2160, Ciudad Universitaria, Pabellón II, 4to piso, C1428EHA. Ciudad Autónoma de Buenos Aires, Argentina. E-mail: [email protected]

Abstract: We documented terrestrial micromammal assemblages at five localities of southwestern Mendoza province, Argentina. We added new localities for several of the most uncommon small species of this region (e.g. Loxodontomys micropus and Tympanoctomys barrerae). Two main groups of terrestrial non-volant micromammals are represented in southwestern province of Mendoza: one group is related to the South American arid diagonal, and includes species typically adapted to the xeric environments of the Monte ; a second group includes Patagonian and High Andean species. Some Patagonian species reached in the study area their northernmost distributional records (e.g. longipilis, A. olivaceus, macronyx, Loxodontomys micropus).

Introduction With the exception of a handful references in the Decrease in rainfall from west to east determines literature, small mammal communities from the the main vegetational types: bunchgrass, shrub southwest of the province of Mendoza, Argentina, steppe, and creosote bush communities. High are still poorly known. Available information elevations of the Andean Cordillera are corresponds mostly to general descriptions of the characterized by cold-dry climate, strong nightly assemblages without vouchers or localities clearly frost, hard soils of fine rock debris, and sparse stated (e.g. Roig 1965) or isolated records from shrubby vegetation; climax communities includes sporadic trappings or owl pellets analyses (e.g. bunchgrass of the Festuca, Poa, and Stipa. Thomas 1894; 1912; Massoia 1981; Massoia et al. Patagonian semidesert, locally known as Payunia, 1994; 1997; Ojeda et al. 2005; Gasco et al. 2006; occupies intermediate elevations, mostly in Nabte et al. 2006). The continuous discoveries of volcanoes slopes. Vegetation types include both new taxa and new locality records for this bunchgrass plains and areas of low shrub and province (e.g. Jayat et al. 2006) demonstrate how brush in basaltic or sandy soil. Patagonian steppes little is known about the small mammal fauna of grade eastwards into the arid , this region. In this note we document the characterized by grasslands and scattered terrestrial micromammal assemblages at five evergreen creosote scrubs of Larrea spp. (Cabrera localities from the southwestern Mendoza, adding 1976). new localities for several of the most uncommon small mammal species of this region. We also Studied assemblages, mostly constituted by made some comments regarding species craniodental remains recovered from pellets of abundance at the different localities and discuss Bubo magellanicus (Aves, Strigidae) and Tyto the biogeographic implications of some of these alba (Aves, Tytonidae), were collected in the and previously known records. following localities (from north to south; between parenthesis we provide the collection numbers Materials and methods under which the material is housed at the The southwest of the province of Mendoza is Colección de Material de Egagrópilas y Afines characterized by a cold temperate climate. "Elio Massoia" del Centro Nacional Patagónico, Rainfall ranges from 600 mm in the Andean Puerto Madryn, Chubut, Argentina [CNP-E]): (1) Cordillera to 300 mm in the Central Monte Desert. Laguna de la Niña Encantada (35º09'38" S,

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69º52'09" W; 1,826 m; CNP-E.55-6); (2) Cerro The employed here follows Pardiñas et Morocho, Laguna Llancanelo (35º36' S, 69º12' W; al. (2003). 1,300 m; CNP-E.19); (3) 10 km W Bardas Blancas on Ruta Provincial 145 (35º52'28" S, Results and discussion 69º52'46" W; 1,152 m; CNP-E.289); (4) La Assemblage compositions per locality and Pasarela (conjunction Río Grande and RN 40, previously known assemblages for this region are 36º18'46" S, 69º40'03" W; 1,250 m; CNP-E.198), given in Table 1. Figure 1 depicts the geographic and (5) Los Frisos (36º28'20" S, 69º38'54" W; location of all recording localities mentioned in 1,101 m; CNP-E.199). Taxonomic identifications this contribution and the main phytogeographic were made by comparisons with reference material. units.

Figure 1. The province of Mendoza (Argentina) including the localities discussed in the text and the main phytogeographic provinces (PP). References: 1, Laguna de la Niña Encantada; 2, Cerro Morocho, laguna Llancanelo; 3, 10 km W Bardas Blancas on RP 145; 4, La Pasarela (conjuction Río Grande and RN 40); 5, Los Frisos; 6, Chacras de Coria; 7, Huayquerías del Oeste; 8, 50 km N San Rafael; 9, San Rafael; 10, Las Aguaditas; 11, Arroyo Malo 3 archaeological site; 12, Valle Hermoso; 13, 6 km NW de Las Leñas; 14, 45 km ENE Volcán Peteroa; 15, Caverna de Las Brujas.

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Recorded species are discussed in the following Abrothrix olivaceus (Waterhouse, 1837) accounts: A. olivaceus ranges from northernmost south to the and the Island of Tierra del Order Didelphimorphia Fuego (Osgood 1943; Teta el al. 2006). This Family Didelphidae species occurs in a wide variety of , Lestodelphys halli (Thomas, 1921) including semi-arid scrub and bushy steppes, L. halli has a relatively homogeneous distribution bunchgrass, meadows, and dense in the Patagonian shrub steppes, from Santa Cruz forests (Pearson 1995). Locality records for to Río Negro (Udrizar Sauthier et al. 2007). Mendoza correspond to high Andean rocky areas Known locality records for this marsupial in the with low sparse shrubs and Payunia environments. province of Mendoza are from, north to south, Massoia (1981) captured and described specimens Chacras de Coria (32º45' S, 71º10' W, the from Las Aguaditas (ca. 34º43' S, 68º05' W; the northernmost record for the species; Birney et al. northernmost Argentinean record for the species). 1996), Huayquerías del Oeste (33º38' S, 68º26' W; These specimens occur in sympatry with Carlini, A., pers. comm.), 50 km N San Rafael molinae, a typical element from the Monte (34º15' S, 68º40' W; Nabte et al. 2006), and Phytogeographical Province (PP, hereafter). Caverna de Las Brujas (35º45' S, 69º49' W; Gasco et al. 2006). These four locality records are (Thomas, 1894) isolated, with an interposed gap of ca. 500 km C. macronyx is found in mesic habitats, such as with the closest southern localities. The forest or meadows from Mendoza south to the significance of this disjunction, if any, remains to Strait of Magellan (Osgood 1943, Pearson 1995). be explored. In addition, it is important to note Known localities in the province of Mendoza that one Late Holocene record has been correspond with High Andean rocky areas with mentioned for Mendoza (Arroyo Malo 3 low sparse shrubs and meadows. Ojeda et al. archaeological site; 34º51'20" S, 69º53'15" W; (2005) documented the karyotypes of topotypical Neme et al. 2002). specimens from Valle Hermoso (35º05' S, 70º06' W; 2,460 m). Thylamys cf. T. pallidior (Thomas, 1902) T. pallidior is found in creosote bush, mesquite Eligmodontia spp. forest, and areas with dense thorny vegetation in Species of Eligmodontia occur in arid and arid and semi-arid regions from south-central semiarid habitats and in both high and low and western south to central and western elevation areas (Mares et al. 2008). Eligmodontia Argentina (Braun et al. 2005). Based on molecular spp. are moderate to dominant in the studied evidences, individuals from San Rafael area samples. At least three species of Eligmodontia – (Mendoza) were referred by Braun et al. (2005) to E. morgani J. A. Allen, 1901, E. moreni Thomas, T. pallidior. We tentatively assigned our records 1896, and E. typus Cuvier, 1837 – occur in to this species; this small didelphid is present in southwestern Mendoza (Lanzone and Ojeda 2005; all studied assemblages. Mares et al. 2008), which are morphologically undistinguishable with fragmentary craniodental Order Rodentia remains. The former is mostly restricted to Family Patagonian steppes and the other two to the Monte (Waterhouse, 1837) PP. A. longipilis ranges from Coquimbo in Chile and Mendoza in Argentina south to Tierra del Fuego Euneomys chinchilloides (Waterhouse, 1839) (Osgood 1943; Teta et al. 2006). A. longipilis is E. chinchilloides lives in bare, windswept, rocky abundant in dense forests of Nothofagus, but also scree from Santiago latitude, in Chile, and occurs in most of the other habitats, such as Mendoza, in Argentina, south to Cape Horn marshes, shrubby steppes, tussock grass, or rocky (Pearson 1995). This species reaches high areas (Pearson 1995). Known localities in the densities in samples from High Andean areas. province of Mendoza appear to be restricted to Specimens from Valle Hermoso, morphologically High Andean rocky areas with low sparse shrubs. identified as E. chinchilloides, have a different

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diploid complement from other specimens Reithrodon auritus (Fischer 1814) previously assigned to this species (Ojeda et al. R. auritus is found in grassland habitats of 2005). Musser and Carleton (2005) restricted the southern , from the Island of Tierra name chinchilloides for Fuegian populations and del Fuego northwards to the Pampean region included the continental samples under E. (Pardiñas and Galliari 2001); north of 36º S its petersoni J. A. Allen, 1903, a taxonomic hypo- populations appear to be restricted to a few high thesis that must be further tested. altitude areas (mainly > 2,000 m). Only two records are certainly known in the province of Euneomys mordax Thomas, 1912 Mendoza, both in open vegetation areas (Jayat et E. mordax is found at high elevations (> 1,700 m) al. 2006, Gasco et al. 2006). The specimens from in the provinces of Mendoza and Neuquén, in Cerro Morocho, Laguna Llancanelo, were Argentina, and adjacent areas of Chile (Reise and previously reported by Jayat et al. (2006) as the Gallardo 1990; Pearson and Christie 1991). Ojeda first record for Mendoza. et al. (2005) documented the karyotypes of topo- typical specimens from Valle Hermoso. This Family Ctenomyidae species occurs in sympatry with E. chinchilloides Ctenomys sp. in Laguna de la Niña Encantada, Caverna de Las Taxonomy of Ctenomys populations from Brujas, and Valle Hermoso (Ojeda et al. 2005; this southwestern Mendoza Province is unclear. C. work). pontifex Thomas, 1918, has its type locality in this area. According to Pearson and Lagiglia (1992), Loxodontomys micropus (Waterhouse, 1837) the relationships between this form and the L. micropus occurs in forest, marshes, humid Chilean C. maulinus Philippi, 1872 and C. dense grasslands, or among bushes in southern maulinus brunneus Osgood, 1943 are unclear. Chile and southwestern Argentina; from about 35º Additionally, Pearson and Lagiglia (1992) S in Mendoza south to Santa Cruz (Pearson 1995). referred specimens from near San Rafael to C. Jayat et al. (2006) listed the first specimens from mendocinus Philippi, 1869. C. mendocinus occurs Mendoza, based on trapped individuals from in northern and central Mendoza from ca. 31º S to Valle Hermoso, 45 km ENE Volcán Peteroa 34º S and from Andean Precordillera eastward (35º05' S, 70º05' W), and from 6 km NW Las across arid sub-Andean regions (Rosi et al. 2005). Leñas (35º06' S, 70º07' W). Here we add a new record for Mendoza, in Laguna de la Niña Family Caviidae Encantada. Microcavia australis (I. Geoffry Saint-Hilaire & d'Orbygny, 1833) Oligoryzomys longicaudatus (Bennet, 1832) M. australis inhabits arid and semiarid lowlands O. longicaudatus occupies brushy places and edge and valleys from Jujuy, northwestern Argentina, of forests from Mendoza south to Santa Cruz, south to Santa Cruz (Cabrera 1953; Tognelli et al. where is replaced by its congeneric O. 2001). magellanicus (Cirignoli et al. 2006). Isolated population occurs in the Monte PP of Mendoza, Family Octodontidae La Pampa, and Buenos Aires (Carbajo and Tympanoctomys barrerae (Lawrence, 1941) Pardiñas 2007), mostly along the main rivers that T. barrerae is found in arid and semi-arid areas of dissect this region. central-western Argentina. This rat is known from only twelve patchy localities in salt basin xanthopygus (Waterhouse, 1837) (salares), sand dune, and open scrubland habitats P. xanthopygus occupies open rocky habitats from in the Monte PP (Díaz et al. 2000; Ojeda et al., Peru south along the to Santa Cruz and 2007). In this work we added one new locality southern Chile (Kramer et al. 1999). This species record to the seven previously known for the dominates the samples from Payunia and High province of Mendoza (cf. Ojeda et al., 2007; Andean areas. Nabte et al. 2006).

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Table 1. Assemblages of micromammals known for Southwestern province of Mendoza, Argentina (x = presence).

Assemblages Previous This paper

1

2 Los Frisos La Pasarela Cerro Morocho Caverna de Las Brujas 10 km W Bardas Blancas Laguna de la Niña Encantada Laguna de la Niña Encantada

Altitude (m a.s.l.) 1,826 1,800 1,826 1,300 1,152 1,250 1,101 Didelphimorphia Lestodelphys halli x Thylamys cf. T. pallidior x x x x x x x Rodentia Cricetidae Abrothrix longipilis x x x Abrothrix olivaceus x x x x x x Chelemys macronyx x x x Eligmodontia spp. x x x x x x x Euneomys chinchilloides x x x x x Euneomys mordax x Euneomys spp. x x Loxodontomys micropus x x Oligoryzomys longicaudatus x Phyllotis xanthopygus x x x x x x x Reithrodon auritus x x Caviidae Microcavia australis x x x x Ctenomyidae Ctenomys sp. x x x x x x Octodontidae Tympanoctomys barrerae x

1 Data from Massoia et al. (1994). 2 Data from Gasco et al. (2006).

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Two large landscape units located in southern tudes; and (4) Two species, Chelemys macronyx Mendoza (Figure 1) are characterized by the total and Loxodontomys micropus, mostly associated absence of data regarding micromammal with the Nothofagus forest, ecotonal areas, and assemblages. These are the volcanic plateau of humid microenvironments in the Andean Payunia (ca. 10,000 km2) and the Sierra del piedmont of Patagonia and High Andean Areas in Nevado chain (with heights of ca. 4,000 m). the province of Mendoza. Despite this fact, that clearly tackles our interpretation of the main zoogeographic patterns, Some species of the subgroups 3 and 4 reach their several general aspects can be addressed. Two northernmost Argentinean distributional limits in main groups of terrestrial non-volant micro- central-western or southwestern of the province of are represented in southern Mendoza. Mendoza. They are Abrothrix olivaceus (northern- One group is related to the South American arid most record in Las Aguaditas, see above), and a diagonal, and includes species adapted to the xeric pool of Patagonian and High Andean species – environments of the Monte Desert (e.g. Akodon Abrothrix longipilis, Chelemys macronyx, molinae, Calomys musculinus, Eligmodontia Euneomys chinchilloides, E. mordax, and typus, griseoflavus). Species of this Loxodontomys micropus – with northernmost re- group are widely distributed through central cord at Laguna de la Niña Encantada (35º17' S). Argentina, in arid and semiarid environments. The With the exception of Lestodelphys halli, the second group includes Patagonian and High species of the groups 3 and 4 or their presumed Andean species. Within this group, we can counterparts (e.g. Euneomys noei, Loxodontomys recognize at least four main patterns of pikumche) are also present on the other side or the distribution: (1) Phyllotis xanthopygus has a Andean chain, in Chilean territory; however, west widespread distribution from Peru to southern of the Andes, they reach more northern locations (including 18º S in the case of A. olivaceus; cf. South America, mostly related to rocky outcrops Osgood 1943). This fact invites to consider if the and partially independent from the main known northern limit in Argentina is the true environmental matrix (Kramer et al. 1999); (2) distributional boundary or just an effect caused by Euneomys mordax restricted to High Andean (> low sampling effort. Along the Andes, between 1,700 m) humid grasslands in southwestern Laguna de la Niña Encantada in the south and Río Mendoza and northern Neuquén, in Argentina, Mendoza in the north, there is a strip of abut 50 x and adjacent areas of Chile; (3) A pool of species 250 km of mostly unexplored territory, where only widely distributed in the Patagonian semiarid some minor collections were made in early XX steppes (e.g. Abrothrix spp., Euneomys century (Thomas 1920). The continuity of chinchilloides); species of this subgroup have a presumably suitable habitats for the taxa above distribution compressed against the Andes in mentioned along this gap suggests that southwestern Mendoza and northern Patagonia, populations of several of these species would that gradually expands to the east in southern lati- remain to be discovered.

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Acknowledgements Sebastián Cirignoli and Dario Podestá helped during field work conducted in southern Mendoza that retrieved several of the studied samples; Hernán Povedano collected and kindly offered the sample from Cerro Morocho. Marcela Nabte assisted during laboratory tasks. Guillermo D'Elía and Sergio Tiranti made valuable comments on an earlier version of this study. Field activities in Mendoza were authorized by Dirección de Fauna Silvestre. This research was funded by CONICET. To the mentioned persons and institutions the gratitude of the authors.

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