Pardiñas, et al. 2007. Resolution of Some Problematic Type Localities for Sigmodontine Rodents...

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resoluTion of some ProBlemaTic TyPe localiTies for sigmodonTine rodenTs (criceTidae, sigmodonTinae)

resolución de algunos ProBlemas relaTivos a localidades TíPicas de roedores sigmodonTinos

(criceTidae, sigmodonTinae)

Ulyses F. J. Pardiñas, Pablo teta, Guillermo D’Elía, Sebastián Cirignoli, and Pablo E. Ortiz

ABSTRACT

Herein we comment on the location of 8 argentinean type localities for 22 sigmodontine taxa (Cricetidae: Sigmodontinae). this study is based on information gathered from original publications, labels of type specimens, field catalogs and notes housed in museums, and historical records and maps. these localities (arranged in order of increasing latitude) are: Higuerilla (Province of Jujuy), Caraguatay and río Paranay (Province of Misiones), río de Oro (Province of Chaco), Otro Cerro (Province of Catamarca), “Norte de Entre ríos” (Province of Entre ríos), Isla Ella (Province of Buenos aires), Neuquén, río Limay (Province of Neuquén), and Valle del Lago Blanco or Valle/región Koslowsky (Province of Chubut). We clarify the geographic location, provide current names, and/or restrict these type localities. Finally, we also pose comments on the taxonomic significance of these actions.

Key words: argentina, Sigmodontinae, type localities, taxonomy

RESUMEn

Sobre la base de una revisión de las fuentes originales, etiquetas de los holotipos, notas de museos y catálogos, reseñas históricas y consulta cartográfica, se discute la ubicación de 8 localidades típicas argentinas de roedores sigmodontinos (Cricetidae: Sigmodontinae). Estas localidades (ordenadas por latitud creciente) son: Higuerilla (Provincia de Jujuy), Caraguatay y río Paranay (Provincia de Misiones), río de Oro (Provincia del Chaco), Otro Cerro (Provincia de Catamarca), “Norte de Entre ríos” (Provincia de Entre ríos), Isla Ella (Provincia de Buenos aires), Neuquén, río Limay (Provincia del Neuquén) y Valle del Lago Blanco o Valle/región Koslowsky (Provincia del Chubut). En todos los casos se comentan las implicaciones taxonómicas y principales problemáticas relacionadas.

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Palabras clave: argentina, Sigmodontinae, localidades tipo, taxonomía

inTRoDUCTion

according to the International Code of Zoological Nomenclature (ICZN, 1999) the geographical place of capture or collection of the name-bearing type of a nominal species or subspecies is the type locality of the taxon in question. Often, when attempting to resolve taxonomic problems, it is of vital importance to collect additional specimens (topotypes) at a particular type locality. Whatever the reasons this material is needed, these specimens must be collected at the type locality of the taxa in question. as such, the value of knowing the exact place of collection of what later become type specimens is clear for all those interested in taxonomy.

Approximately 129 named taxa, either species or subspecies, have been based on sigmodontine rodents (Cricetidae, Sigmodontinae) collected in argentina. these specimens were collected at 90 localities (Fig. 1). the distribution of these type localities is highly asymmetrical, with most concentrated in those provinces extensively surveyed during the early 19th and late 20th centuries. Buenos aires Province contains the maximum number of sigmodontine type localities (16), followed by tucumán Province (10), and Jujuy Province (8), while several provinces (e.g., Santiago del Estero, Santa Fe) have none. at least 3 periods of taxonomic descriptions can be recognized in the history of the study of argentine sigmodontine rodents (Fig. 2). the first, early in the 19th century, involved the pioneer work of azara, Darwin, and Waterhouse. the second and most prolific period, from 1890 to 1930, had as main protagonists Emilio Budin in the field and Oldfield thomas at the British Museum of Natural History. Finally, a third period, beginning around 1970, included the work of several contemporary mammalogists (e.g., Massoia, Contreras, Mares, and Braun). this third period is characterized by the fact that, in general, collectors and describers are the same person.

Unfortunately, several argentine sigmodontine type localities are difficult or impossible to locate. this situation, which evidently hampers several taxonomic studies, is due to a large array of problems ranging from an original definition involving vast or ambiguously defined areas (e.g., many of the Waterhouse references), to the use of names never recorded in maps (e.g., Higuerilla), and to the changing of some geographic names (e.g., Bonifacio = Laguna alsina). these problems are rooted in several sources, most notably: a) lack of maps or detailed maps at the time of collection; b) large temporal gaps between the collection of specimens and their description, which led to the publication of incorrect, misinterpreted, or ambiguous references; c) changes or restrictions of several type localities proposed by authorities unfamiliar with the geography of the country and/or based on misinterpretations of the original or alternative information sources.

Concern about these problematic type localities was manifested as early as the first decades of the last century when thomas (1919c, 1920) discussed the location and the real existence of some of them. Later, comments in relation to these issues were posed mainly as isolated notes in taxonomic or biogeographic contributions (e.g., Pearson, 1958; Hershkovitz, 1962; Massoia and Fornes, 1964; Myers et al., 1990; Pardiñas, 1996; Pardiñas and Galliari, 1998; Díaz, 1999). However, 3 studies directly focused on these problematic type localities. First, Pearson and Lagiglia (1992) specifically discussed, based on historical sources and newly trapped material, the correctness of Fuerte

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Figure 1. argentinean sigmodontine type localities. the numbers refer to localities discussed in the text and detailed in the maps of Figs. 3, 4, and 5.

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de San rafael (Mendoza Province) as the type locality of 3 sigmodontines. Similarly, Contreras (1992) and Contreras and teta (2003), reinterpreting the original works of Rengger and Azara, discussed the placement of the type localities of Calomys callosus and Oxymycterus rufus, respectively.

We have 2 objectives in this report. First, we want to call attention of those working with sigmodontine rodents to the fact that the geographic origin of several type specimens is uncertain. Second, we discuss 8 of the most problematic argentine sigmodontine type localities, to clarify their probable locations. In addition, we comment on the taxonomic uncertainties remaining with respect to several of the taxa involved.

MatErIaLS aND MEtHODS

this paper is based on the following sources of information: a) original publications, b) public historical records, c) catalogues and field notes housed in museums, d) original labels attached to holotypes, and e) consultation with mammalogists, historians, and cartographers. as cartographic source we used historical and contemporaneous topographic maps, the latter produced mainly by the Instituto Geográfico Militar (IGM, argentina). Most of the geographical coordinates provided in this paper (at degree and minute resolution) were obtained from the atlas de la república argentina (IGM, 1972, 1979). In several cases, we used GPS data obtained by us. For each locality, we also provide the names of the departments and provinces.

We discuss the type localities arranged in order of increasing latitude. In all cases, we respected the original spelling of the taxa. Finally, we followed the article 76 of

Figure 2. New taxa (binomials and trinomials) of sigmodontine rodents described per decade on the basis of Argentinean specimens.

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the International Code of Zoological Nomenclature (ICZN, 1999), particularly, the recommendation 76a that states that “in ascertaining or clarifying a type locality…an author should take into account: 76a.1.1. data accompanying the original material; 76a.1.2. collector’s notes, itineraries, or personal communications; 76a.1.3. the original description of the taxon; and 76a.1.4. as a last resort, and without prejudice to other clarification, localities within the known range of the taxon or from which specimens referred to the taxon had been taken.”

PRoBLEMATiC LoCALiTiES

1. the phantom type locality called Higuerilla

History: Emilio Budin collected in this locality in 1921. thomas (1921b, p 609) offered the original geographic reference as “Higuerilla.- 2000 m., in the Department of Valle Grande, about 10 km. east of the Zenta range and 20 km. of the town of tilcara.” Cabrera (1961, pp 451 and 466) added 2 alternative interpretations: “Higuerilla, Jujuy, 10 kilómetros al este de la Sierra de Zenta” and “Higuerilla, provincia de Jujuy, a unos 20 kilómetros al este de tilcara y a 2.000 metros de altura.” Subsequent authors repeated thomas’s reference with minor variations (e.g., Hershkovitz, 1994; Musser and Carleton, 1993; Díaz, 1999).

Problem: there is not a single place with this name on any available map of central Jujuy Province.

Present Location (Fig. 3a): Valle Grande (23° 28’ S, 64° 59’ W, 2313 m, Valle Grande, Jujuy) is the main city in the Department of Valle Grande. the region is characterized by the scarcity of roads; only provincial route 83 crosses the Calilegua National Park connecting the cities of Libertador General San Martín (23° 49’ S, 64° 47’ W, 1551 m, Ledesma, Jujuy) and Valle Grande. Searches carried out in the area recording local place names from old settlers indicate that Higuerilla was the name of an abandoned ranch (23° 33’ 35” S, 65° 00’ 37” W, 1804 m, Valle Grande, Jujuy) where the small village of Pampichuela is now located. It seems likely that Budin trapped in the vicinity of this ranch and recorded its name as the locality of his collections. therefore we equate Higuerilla with the modern locality Pampichuela (23° 32’ S, 65° 02’ W, 1735 m, Valle Grande, Jujuy; IGM 1989a). Pampichuela is about 15 km (by road) south of Valle Grande, and ca. 33 km east of tilcara (23° 34’ S, 65° 22’ W, 2461 m, tilcara, Jujuy).

Taxonomic Implications: Higuerilla is the type locality of the sigmodontines Phyllotis nogalaris thomas, 1921; Hypsimys deceptor thomas, 1921; and Oxymycterus akodontius thomas, 1921. although the first species was included in several taxonomic studies of the genus Phyllotis (e.g., Pearson, 1958; Hershkovitz, 1962), the taxonomic status of H. deceptor and O. akodontius remains unclear. the differences between H. deceptor and H. budini thomas, 1918 (type locality: León, 24° 03’ S, 65° 26’ W, 1754 m, Dr. Manuel Belgrano, Jujuy) were dismissed by Cabrera (1961). Now both forms are considered synonyms and placed in the genus Akodon (see Myers and Patton, 1989). Oxymycterus akodontius has been retained as a valid species but with the widespread view that it must be critically reevaluated (e.g., Cabrera, 1961; Galliari et al., 1996; Musser and

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Carleton, 2005). Holotypes and original series of both species were never restudied, and no new specimens from the area of the type are available.

Figure 3. Detailed maps of type localities discussed in the text (numbers refer to the order of presentation in the main text; see also Fig. 1): a (Site 1 in Fig. 1). Higuerilla (Jujuy Province), B (Site 2 in Fig. 1). Caraguatay and Paranay (Misiones Province), C (Site 3 in Fig. 1). río de Oro (Chaco Province).

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2. Where are Caraguatay and Paranay?

History: Collin Campbell Sanborn sampled these localities during the Captain Marshall Field Brazilian Expedition (Field Museum of Natural History - reports, 1927). the first published reference was given by Sanborn himself (1931, p 1) when describing Oxymycterus misionalis, as “rio Paranay, an affluent of the rio Parana, near Caraguatay, about 100 miles south of rio Iguassu, Misiones territory, argentina.” two years later, Osgood (1933) annotated “Caraguatay, rio Parana, 100 miles south of rio Iguassu, Misiones, argentina” in reference to the type locality of his Thomasomys pictipes. Cabrera (1961, p 467) changed the type locality of O. misionalis, suppressing the reference to “rio Paranay”, writing “Caraguatay, rio Paraná, Misiones, argentina.” More recently, Gonzalez (2000, p 4), misinterpreting a personal communication of Pardiñas, explained (in reference to the type locality of T. pictipes) that “Caraguata-i es una isla del río Paraná frente a la localidad de Monte Carlo, Provincia de Misiones, argentina.”

Problem: Published geographic references do not provide enough information to unambiguously locate the type locality of these 2 sigmodontines.

Present Location (Fig. 3B): Information of the Field Museum of Natural History - reports (1927, p 69) help to identify Sanborn’s itinerary more precisely. the narration of the itinerary of Captain Marshall Field Brazilian Expedition indicates that “from asuncion [Paraguay], the remaining members of the expedition went by rail to Posadas, Missiones [Misiones] territory, argentine republic. Here they took a steamer on the upper Paraná for Iguazu Falls. En route, they accepted the invitation of Mr. Carlos H. Benson to camp and collect on his plantation at Caraguatay where, midway between Posadas and Iguazu Falls, Messrs. Sanborn and Ganzon stopped…the second week in Missiones was spent in camp on the rio Paranay, a small affluent of the Paraná, where birds and amphibians were of interest but mammals and reptiles scarce.” Charles Herbert Benson was one of the owners of “La Misionera S.a.I. y C.,” a 4,000 ha yerba mate plantation established around 1915. By 1926 Benson had the main house of his property in Puerto Caraguatay, on the río Paraná. Sanborn (1931, p 2) also collected several specimens of his O. misionalis in Caraguatay, but he selected a specimen from “rio Paranay” to serve as holotype.

at least 4 places are named “Caraguatay” or “Caraguata” in central Misiones Province, all of them are close or on the río Paraná. they are (IGM, 1994a, 1994b): Paraje Caraguatay (26° 39’ S, 54° 44’ W, 200 m, Montecarlo, Misiones), Colonia Caraguatay (26° 48’ S, 55° 20’ W, Montecarlo Misiones), Cerro Caraguatay (26° 42’ S, 55° 21’ W, Montecarlo, Misiones), and Puerto Caraguatay or Puerto Caraguatá (26° 37’ S, 54° 46’ W, 192 m, Montecarlo, Misiones). the “rio Paranay,” actually río Paranay Guazú, runs from the central Sierra de Misiones to the río Paraná, about 10 km south of Puerto Caraguatay. Near the confluence is located a small village named Puerto Paranay (26° 41’ S, 54° 48’ W, 122 m, Libertador General San Martín, Misiones). the type locality of T. pictipes can be restricted to Puerto Caraguatay (see above). In the case of O. misionalis, we restrict its type locality to the confluence of the río Paranay Guazú with the río Paraná (26° 41’ S, 54° 49’ W, 122 m, Libertador General San Martín, Misiones; IGM, 1994b).

Taxonomic Implications: González (2000) erected Juliomys with Thomasomys pictipes as

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type species. this author studied mainly Brazilian specimens. recent studies have revealed an unexpected level of alpha diversity in Juliomys (see Oliveira and Bonvicino, 2002). In fact, the application of pictipes to populations in Brazil started with Pine (1980), but he pointed out several differences between the type of pictipes and the specimens from southern Brazil. regrettably, nobody has recently collected at the type locality, and no new specimens are known from Misiones Province.

Oxymycterus misionalis is considered a synonym of O. hispidus following Cabrera (1961); however, this hypothesis never was tested studying topotypes or, at least, specimens from Misiones Province. Clearly, the collection of new specimens of both J. pictipes and O. misionalis in the type area Caraguatay-río Paranay is necessary to clarify their taxonomic identities.

3. río de Oro and the missing Scapteromys chacoensis

History: the first mention of this locality was made by Gyldenstolpe (1932a) when he described Scapteromys chacoensis. Gyldenstolpe (1932a, p 2) gave the type locality as “argentine, rio de Oro, Chaco austral. (Caught in ‘Lagunas de aqua dulce’).” the holotype and only specimen known of this taxon was collected and sent to the royal Natural History Museum of Stockholm by a. ros in September, 1896 (Gyldenstolpe, 1932a, p 2).

Problem: there is no additional information about this type locality, the collector, or the circumstances related to the acquisition of this specimen in museum records or on the specimen label. the transcription on the latter is “argentina, rio de Oro, Chaco austral. Ix. 1896. a. ros.”

Present Location (Fig. 3C): the río de Oro is a tributary of the río Paraguay. With its headwaters in the vicinity of Siete arboles (ca. 26° 18’ S, 59° 30’ W, Libertador General San Martín, Chaco), the río de oro extends 140 km across the northeastern portion of Chaco Province. three small towns are settled on this river (IGM, 1967), General José de San Martín (26° 33’ S, 59° 20’ W, 57 m, Libertador General San Martín, Chaco) in the upper course, Selvas del río de Oro (26° 48’ S, 58° 57’ W, 60 m, Libertador General San Martín, Chaco) in the middle, and General Vedia (26° 56’ S, 58° 40’ W, 58 m, Bermejo, Chaco), near the mouth (27° 03’ S, 58° 33’ W, Bermejo, Chaco). at this time is not possible to restrict the type locality of Scapteromys chacoensis to any place along the río de Oro.

Taxonomic Implications: Scapteromys chacoensis was synonymized with Kunsia fronto (Winge, 1887) by Hershkovitz (1966), under the trinomial Kunsia fronto chacoensis. Subsequently, avila-Pires (1972) described K. f. planaltensis based on a series of specimens trapped in the early 1960s by J. Moojen in Brasilia (Distrito Federal, Brazil). Kunsia fronto is one of the more poorly-known sigmodontines, having been recorded alive only on the río de Oro and at Brasilia. In both cases, despite intensive trapping efforts (Hershkovitz, 1990; Cirignoli et al., 2000), no new specimens were obtained.

Remarks: Despite intensive agriculture, cattle rising, and deforestation during the last 100 years, the río de Oro basin is moderately well preserved. Extensive palms and flooded grasslands remain in this area. the local extinction of K. fronto cannot be demonstrated until new fieldwork covering the entire river course would be made.

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However, analysis of owl pellet from 12 localities from General José de San Martín to General Vedia, which included thousands of specimens of small mammals, failed to yield remains of K. fronto (Cirignoli et al., 2000).

4. Is Otro Cerro a fancy name?

History: the first collector to work at this locality was Emilio Budin in 1918. thomas (1919c, p 489) discussed the real existence of Otro Cerro as follows “after making the collection at Chumbicha, Catamarca…Sr. Budin traveled westwards some 30 miles, over the Cerro de ambato to ‘another hill,’ or range of hills, for which he could find no recognized name…although Sr. Budin considered himself to be still in Catamarca…the distance and direction from Chumbicha show that he must have crossed the frontier into rioja. Under these circumstances I have decided to use Sr. Budin’s fancy name of ‘Otro Cerro’…the locality may be stated as 45 kilometers west of Chumbicha, the specimens being taken at an altitude of about 3000 meters” (italics ours). One year later thomas (1920, p 473) corrected himself by pointing out that Otro Cerro is located in Catamarca Province, and not in La rioja Province (= rioja, in his original spelling). He also wrote that “[Otro Cerro]…is in Catamarca, about 18 km. N.N.W. of Chumbicha.” almost all later authors used thomas’ (1920) second geographical reference (e.g., Hershkovitz, 1962; Williams and Mares, 1978; Mares et al., 1997; Díaz, 1999), but 3 exceptions deserve note. First, Pearson (1958, p 418), when discussing the status of Phyllotis ricardulus, and clearly following thomas (1919c) in the attribution of the name to Budin, wrote “Otro Cerro is a locality name coined by Budin to refer to a mountain 11 miles NNW of Chumbicha” (italics ours). Second, Cabrera (1961, p 487) stated “Contra lo que supone Pearson, Otro Cerro no es un nombre inventado (‘coined’) por Budin para dicha localidad típica, puesto que aparece en el mapa de Catamarca de Lange y Delachaux publicado por el Museo de La Plata en 1893.” Cabrera (1961, p 501) referenced this type locality as “Otro Cerro, en el extremo sur de la sierra de ambato, al noroeste de Chumbicha, Catamarca.” third, Myers et al. (1990, p 62) mentioned thomas’ (1919c) original supposition that Budin may have confused Otro Cerro and Cerro Nunorca, speculating that Otro Cerro is “…probably in La rioja Province…this mountain is probably within a range (for which we also have not found a name) that runs NNW from the Sierra de Valasco, originating west of La rioja.” Myers et al. (1990) tentatively changed the location of otro Cerro back to La Rioja Province, remarking one more time on the uncertainties about the exact location of this type locality.

Problem: In spite of the fact that Otro Cerro has appeared both in old and recent official argentine maps, uncertainties about the location of this type locality still remains (e.g., Myers et al., 1990).

Present Location (Fig. 4a): the map of Lange and Delachaux (Museo de La Plata, 1893) shows the location of Otro Cerro as a ranch in the southern end of the Sierra de ambato, Catamarca Province. In addition, this place is figured in the topographic map IGM (1966) but, regrettably, Otro Cerro has disappeared in the most recent edition (IGM, 1989b). an in situ inspection of this area revealed that local settlers of the small village of trampasacha (28° 50’ S, 66° 18’ W, 614 m, Capayán, Catamarca), the nearest habited place to Otro Cerro, know the existence of the latter as an abandoned ranch at the top of the Sierra de ambato. the exact location of Otro Cerro is 28° 45’ S, 66° 17’ W (2023

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m, Capayán, Catamarca), about 4 km SSE of Cerro Catalán (IGM, 1989b).

Taxonomic Implications: Otro Cerro is the type locality of several mammals (thomas, 1919c, 1920), including the sigmodontines Phyllotis ricardulus Thomas, 1919, Graomys edithae Thomas, 1919, Akodon alterus Thomas, 1919, Akodon orbus Thomas, 1919, and Reithrodon caurinus thomas, 1920. Of these 5 taxa, only P. ricardulus has been restudied, and now is considered a synonym of P. xanthopygus (Pearson, 1958; Hershkovitz, 1962). The status of G. edithae has been discussed by several authors. Cabrera (1961) treated it as a synonym of G. griseoflavus, without explaining his reasons. Massoia (1976) and Williams and Mares (1978) commented on the uncertainties regarding this form, but did not offer any resolution. although this taxon is currently treated as valid species (e.g., Musser and Carleton, 1993, 2005; Mares et al., 1997) nothing is known beyond the original description. Akodon alterus is alternatively considered a synonym of A. spegazzinii (Myers et al., 1990; Musser and Carleton, 1993) or as a full species (Blaustein et al., 1992; Díaz, 1999). Akodon orbus was synonymyzed with Necromys lactens by Cabrera (1961), probably on the basis of the similarities between both taxa remarked by thomas (1919c). However, the holotype of A. orbus has been never restudied or compared with other Necromys species. Finally, R. caurinus was subsumed under R. auritus, without any evidence favoring this hypothesis (Cabrera, 1961). trapping efforts at Otro Cerro are essential to clarify the status of these named forms.

Remarks: In contrast with Chumbicha (28° 52’ S, 66° 14’ W, 376 m, Capayán, Catamarca; another type locality very close to Otro Cerro), where the human-disturbance can be considered as severe and the original vegetation (Montane Chaco) has been almost totally replaced, Otro Cerro remains relatively undisturbed. this fortunate situation may be due to the isolation and difficult access to this locality. the high-elevation grasslands in the southern end of the Sierra de ambato are used by domestic cattle (goats), but probably several well-preserved places remain.

5. Why “Norte de Entre ríos” is not a type locality

History: azara (1802, p 94) described, under the name of “agreste” and based on a specimen collected “...en los 30 ½ grados de latitud...,” the species that Fischer (1829) later selected as the basis of his M[us]? azarae. tate (1932a, 1932b, p 26) included this species in Akodon, recording its locality as “30 ½° (= latitude of Entre ríos) argentina.” Later, Cabrera (1961, p 440) reported the type locality of A. azarae as “...los 30° 30’ de latitud sur y entre los ríos Uruguay y Paraná, o sea en el norte de la actual provincia de Entre Ríos” (italics ours). However, neither of these authors explained their selection of “Norte de Entre ríos” for azara’s collections. By 1796, azara had left asunción, Paraguay to participate in the defense of the borders of the Spaniards colonies against the Portuguese empire in the area of the current Brazilian state of rio Grande do Sul. there azara participated in the establishment of the towns of San Grabriel de Batoví and Villa Esperanza (Mariluz Urquijo, 1953; Mones and Klapenbach, 1997). the reference to the 30° 30’is recurrent in the Spanish version of azara’s (1802) work, in which several sigmodontines were described. Examples are the “Orejón” (azara, 1802, p 83), the “Blanco debaxo” (azara, 1802, p 181), and the “cola igual al cuerpo” (azara, 1802, p 87). Precisely, it was in the description of the later that azara provides an additional reference of fundamental value to locate the place of his collections “...[la

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obtuve] en un huerto campestre de la frontera del Brasil hacia los 30 ½ grados de latitud” (azara, 1802, p 97). this mention must correspond to San Gabriel de Batoví, because at that time there was no other locality at the same latitude along that frontier. as another piece of evidence regarding the “Murciélago orejón” (= Histiotus velatus, see Hershkovitz, 1987), azara (1802, p 304) wrote “...tuve dos o tres en los 30 ½ grados de latitud...” the distribution of this species does not reach the argentinean province of Entre ríos (see Barquez et al. 1999), but does include the Brazilian state of río Grande do Sul (Silva, 1985).

Problem: the selection of “Norte de Entre ríos” as the type locality of these taxa is arbitrary and it is partially at odds with azara’s itinerary between 1796 and 1801.

Present Location: San Gabriel de Batoví, today São Gabriel (30° 19’ S, 54° 19’ W, 118 m), is in the Brazilian State of rio Grande do Sul.

Taxonomic Implications: the relocation of the type locality of A. azarae to São Gabriel prompts substantial changes at the subspecific level of this complex taxon. a study of geographic variation in this species is currently underway (D’Elía, Pardiñas, and teta, in prep). Due to geographic reasons it is probable that the “Orejón,” usually included in the synonymy of Reithrodon auritus (Hershkovitz, 1959, 1987; tate, 1932a) should be considered a synonym of R. typicus (see Pardiñas and Galliari, 2001). Mus dubius Fischer, 1829, based on the literal type of the “blanco debaxo,” was included in Calomys by Cabrera (1961) and Langguth (1975), and later synonymized with Calomys laucha by Musser and Carleton (1993).

Remarks: Contreras and Justo (1996), in a meeting abstract, were the first authors to suggest that San Gabriel de Batoví is the true type locality of A. azarae. as discussed by Contreras and teta (2003), Hershkovitz (1994) restricted the type locality of Oxymycterus rufus as “north of the province of Entre ríos,” based on an erroneous interpretation of azara’s work.

6. Where is Isla Ella?

History: Collections in this island were made by robin Kemp in 1917. thomas (1917, p 95) offered the first and unique geographical reference as “…Isla Ella, in the delta of the rio Parana, at the top of the La Plata Estuary.” Subsequent authors used thomas’ reference (e.g., Gyldenstolpe, 1932b; Cabrera, 1961; Musser and Carleton, 1993). Hershkovitz (1966, p 97) and González and Pardiñas (2002, p 3) commented explicitly about the uncertainty of the exact location of Isla Ella. thomas (1917, p 98) presented additional information useful to locate Isla Ella when he explained the etymology of Akodon arenicola hunteri. this subspecies was named in honor of “Mr. James Hunter, the Managing Director of the Isla Ella Fruit and Forest Company.” according to P. Pereyra (Museo Naval de la Nación, tigre, Buenos aires, pers. comm. 2003), the Isla Ella Fruit and Forest Company was located on the arroyo Espera, a stream selected by its good navigability. Udaondo (1942, p 121), in a historical summary of this region, wrote “Merecen especial atención...el frigorífico de Hunter, donde se preparan en envases especiales las frutas destinadas a la exportación...” In addition, Gil (1895) indicated that by that time the Hunter family tended beehives on the arroyo Esperita,

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and Cuomo (1967) recorded the name “Ellan” for a house (or a boat pier) in an island on the arroyo Espera. No information is available concerning when the Isla Ella Fruit and Forest Company ceased operation. regrettably, the Public record Office (United Kingdom; http://www.pro.gov.uk) has no record of this Company.

Problem: there is no single place recorded in any available map with the name of Isla Ella.

Present Location (Fig. 4B): the Delta del Paraná is a large region of islands, canals, and streams at the mouth of the río Paraná. Most of the numerous islands lack specific names. We believe, according to the information presented above, that Isla Ella is a small island (ca. 1 km2) surrounded by the streams Espera and Esperita (34° 22’ S, 58° 38’ W, Primera Sección Delta del Paraná, tigre, Buenos aires; IGM, 1959). this island is about 8 km NNE of tigre city (34° 25’ S, 58° 35’ W, 2 m, tigre, Buenos aires).

Taxonomic Implications: One genus, 3 species, and 1 subespecies were named from specimens collected in Isla Ella (thomas, 1917, 1920). these are: Deltamys kempi Thomas, 1917, Oligoryzomys delticola Thomas, 1917, Akodon arenicola hunteri Thomas, 1917, and Scapteromys aquaticus thomas, 1920. the generic status of Deltamys has been discussed by several authors (see D’Elía et al., 2003 and references therein), but no study following the original description has included specimens from this island. The status of A. a. hunteri remains uncertain. this subspecies was subsumed under Akodon azarae (see Cabrera, 1961), a widespread species of Akodon which has never been systematically reviewed (see comments on A. azarae in the “Norte de Entre ríos” account). In addition, thomas (1921a) pointed out striking resemblances between A. a. hunteri and A. sylvanus thomas, 1921, the latter from Sunchal (24° 16’ S, 64° 27’ W, 1140 m, Santa Bárbara, Jujuy). Following early observations by thomas (1921a), Cabrera (1961) synonymyzed A. sylvanus with A. azarae. In addition, Hershkovitz (1990), in a footnote, indicated the potential conspecifity of A. sylvanus and A. a. hunteri, but not with A. azarae. D’Elía and Pardiñas (2004) recently revised Scapteromys, but they did not include topotypical material. Finally, the distinctiveness of O. delticola, thought to be very close to O. longicaudatus (Osgood, 1943) has not been critically evaluated in any comprehensive review of the genus; however, recently Francés and D’Elía (in press) proposed the formal synonymy of O. delticola under O. nigripes.

7. the obvious, but neglected type locality of Akodon neocenus

History: a single specimen of Akodon trapped by E. Weiske on November 4, 1910, was made the holotype of A. neocenus by thomas (1919b). thomas (1919b, p 213) referenced its type locality as “Neuquén, rio Limay, Upper rio Negro, Patagonia.” Subsequent authors introduced in this geographic reference minor, but important, variations. Gyldenstolpe (1932b, p 103) stated “Western Patagonia, rio Limay, Upper rio Negro, Neuquén territory.” Cabrera (1961, p 449) shortened it to “río Limay, provincia de Neuquén.” Finally, Myers (1989) offered a literal transcription of thomas (1919b) early reference; he also located the place in a map (Myers, 1989: Fig. 2b), in the middle course of the río Limay. all these changes are relevant because, with the introduction of the term “territory” or “province” associated with “Neuquén,” this type locality was partially diluted. However, the original reference is clear and self-explanatory

Pardiñas et al.: Problematic Type Localities for Sigmodontine Rodents 403

Figure 4. Detailed maps of type localities discussed in the text (numbers refer to the order of presentation in the main text; see also Fig. 1): a (Site 4 in Fig. 1). Otro Cerro (Catamarca Province), B (Site 6 in Fig. 1). Isla Ella (Buenos aires Province); the star indicates the island tentatively identified as this type locality.

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without emendations. the community of Neuquén was established a few years after the “Desert Conquest” in 1880. In 1902 the railway station of Neuquén was created, and 2 years later Neuquén was selected as main city of the territory (alvarez, 1988). the río Limay forms the southern limit of the city; below its confluence with the río Neuquén (38° 59’ S, 68° 01’ W), it becomes the río Negro. the holotype of A. neocenus probably was trapped in the vicinity of Neuquén close to the río Limay. It is important to note that in 1910, the year of collection, work was begun to develop an irrigation system in the río Negro valley. In fact, the headwaters of this river are known as “alto Valle del río Negro,” the plausible origin of “Upper rio Negro” in the original geographic reference.

Problem: Successive alterations of the name of this type locality gradually increased the uncertainties concerning its location.

Present Location (Fig. 5a): Neuquén (38° 57’ S, 68° 04’ W, 259 m, Confluencia, Neuquén) is settled on the inner corner formed by the confluence of 2 main rivers, the northern río Neuquén, and the southern río Limay (IGM, 1994c).

Taxonomic Implications: Akodon neocenus was alternatively considered as a full species (e.g., thomas, 1919b, 1927; Musser and Carleton, 1993; Galliari et al., 1996; Pardiñas et al., 2003) or as a subspecies of A. varius (e.g., Gyldenstolpe, 1932b; Cabrera, 1961). this fluctuating scenario is due mainly to the fragmentary nature of the holotype (an imperfect skin and skull) and to the lack of additional specimens collected in the type locality. topotypes are fundamental to assessing not only the status of A. neocenus, but also that of the related species as A. dolores, A. molinae, and the recently described A. oenos (see Braun et al., 2000).

8. What is the Koslowsky region or Valle del Lago Blanco?

History: the first rodents from this locality were trapped by Julio Koslowsky in 1900, and sent to Oldfield thomas between 1900 and 1903. thomas (1903, 1916, 1919a) referenced this locality as “Valle del Lago Blanco, Southern Chubut (Cordillera region)” (thomas, 1903, p 241), “Valle del Lago Blanco, Cordillera region of Southern Chubut territory, Patagonia” (thomas, 1903, p 243), “Koslowsky Valley, 46° S., 71° W., Central Patagonia” (thomas, 1916, p 187), or “Valle de Lago Blanco, Koslowsky region, Patagonia, 46° S.” (thomas, 1919a, p 205). In reality, there is no map that includes these place names exactly as written by thomas. On the contrary, a small town called Lago Blanco (45° 57’ S, 71° 17’ W, 598 m, río Senguerr, Chubut) is located in the southwestern corner of the Lago Blanco, and the ranch Estancia Lago Blanco (45° 54’ S, 71° 20’ W, río Senguerr, Chubut) is on the northern margin of the same lake (IGM, 1950). However, the history of this type locality is more complicated; fortunately, the historical research by aguado (2003) helps to pinpoint its exact locality (see also aguado and Williams, 2003). Julio Koslowsky (1866-1923), the first argentinean herpetologist, funded a colony of russian, Polish, and Lithuanian families in 1898 on the banks of the arroyo Huemules. this colony only lasted 2 years, but Koslowsky lived there from 1898 to 1904. During these years, Koslowsky collected and sent to different museums numerous specimens of plants and vertebrates. Most of these specimens were labeled as “Valle del Lago Blanco.” But this geographical reference must be understood in its historical context.

Pardiñas et al.: Problematic Type Localities for Sigmodontine Rodents 405

the place where Koslowsky’s first ranch (called Estancia Los Halcones) was located is a valley (now known as Valle Huemules) that covers the area from Lago Blanco in the east to the argentine – Chilean border in the west. So, Koslowsky used the name of “Valle del Lago Blanco” as a geographic source of his collections. this situation is revealed by Koslowsky (1904) himself when he described 2 mammals obtained in this area. the title of his 1904 article is “Dos mamíferos de Patagonia cazados en el Valle del Lago Blanco [territorio del Chubut].” But, in the text, he wrote the exact provenance of one of these mammals as “El Felis guigna lo cacé el 1° de Febrero de 1902 en los

Figure 5. Detailed maps of type localities discussed in the text (numbers refer to the order of presentation in the main text; see also Fig. 1): a (Site 7 in Fig. 1). Neuquén (Neuquén Province), B (Site 8 in Fig. 1). Valle del Lago Blanco or Valle/region Koslowsky (Chubut Province).

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bosques de la Cordillera sobre el río Aysen, cuyo lecho y orillas son la continuación oeste del valle del Lago Blanco” (Koslowsky, 1904, p 131). the río aysen (presently in Chilean territory) was in the vicinity of the Koslowsky ranch, more than 25 km west of Lago Blanco. the names “Valle Koslowsky” or “region Koslowsky” are historical names that were used by the commission that surveyed the argentine- Chilean border between 1896 and 1902, and were used to reference the area occupied by Koslowsky’s ranch, i.e., the present Valle Huemules (see aguado, 2003).

Problem: there is not a single place name recorded on any available map with the name of Koslowsky region or Valle del Lago Blanco.

Present Location (Fig. 5B): the “Koslowsky region,” “Koslowsky valley,” or “Valle del Lago Blanco,” as used by both Koslowsky and thomas, are historical and obsolete names applied to the Valle Huemules. this valley includes an area of subandean grasslands, oriented approximately east and west, from the Lago Blanco on the east (western margin of Lago Blanco = 45° 55’ S, 71° 19’ W; IGM, 1950) to the Chilean border in the west (the río Simpson); the northern (45° 50’ S, 71° 31’ W) and southern (46° 03’ S, 71° 31’ W) borders of this valley are surrounded by mountain ridges or basaltic plateaus. Valle Huemules covers an area of ca. 1,200 km2. It is impossible to say where exactly in this valley Koslowsky trapped the rodents. therefore, for taxonomic and nomenclatorial purposes, we restrict this type locality to Estancia Valle Huemules (45° 57’ S, 71° 31’ W, 593 m, río Senguerr, Chubut). this selection appears to be adequate because this ranch is located in the middle of the Valle Huemules –an area certainly crossed by Koslowsky many times during his collecting days- and is identified on several maps (e.g., IGM, 1951). the first Koslowsky ranch (Estancia Los Halcones, ca. 45° 58’ S, 71° 36’ W, río Senguerr, Chubut) is now part of the Estancia Valle Huemules.

Taxonomic Implications: One subgenus, 3 species, and 1 subspecies were described from specimens collected by Koslowsky in the Lago Blanco area (see thomas, 1903, 1916, 1919a). these are: Akodon suffusus Thomas, 1903, Akodon (Chelemys) vestitus Thomas, 1903, Oryzomys magellanicus mizurus Thomas, 1916, and Akodon iniscatus Thomas, 1919. In addition, at least 1 individual of the rare sigmodontine Notiomys edwardsii was collected there (see thomas, 1919a; Pardiñas and Galliari, 1998). as currently understood, A. iniscatus, which distributes across most of Patagonia, is a complex taxon whose geographic variation and biological boundaries have not been correctly assessed. For a revision of this species, the collection of topotypes in the Valle Huemules area is needed. Akodon suffusus (= Abrothrix longipilis), Akodon (Chelemys) vestitus (= Chelemys macronyx), and Oryzomys magellanicus mizurus (= Oligoryzomys longicaudatus) are taxa similarly widely distributed in southwestern argentina and Chile, and lack formal taxonomic revisions (see Pardiñas et al., 2003; Palma et al., 2004; and references cited therein).

Remarks: the region of Valle Huemules was considered one of the richest in Patagonia, promoting several border conflicts between argentina and Chile at the beginnings of the 20th century (aguado, 2003). Photographs taken by Koslowsky between 1898 and 1910 show extensive grasslands, numerous lakes, streams, small rivers, and dense forests in the nearby mountain ridges. Intensive sheep ranging and forest extirpation started in 1908; the effects of these practices on small mammal populations have not

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been assessed. a fieldtrip in late 1980s, with the goal of obtaining new specimens of A. iniscatus, was undertaken by Orlando Scaglia (pers. comm.). He trapped, mainly in overgrazed fields, in the vicinity of the town of Lago Blanco but only captured Abrothrix longipilis. However, he found A. iniscatus remains in owl pellets gathered in this area.

SUMMarY OF FINDINGS

Type Locality: Higuerilla (Jujuy, argentina).

Action Proposed: Equate this type locality to Pampichuela (23° 32’ S, 65° 02’ W, 1735 m, Valle Grande, Jujuy, argentina).

Sigmodontine Taxa Affected: Phyllotis nogalaris thomas, 1921; Hypsimys deceptor Thomas, 1921; and Oxymycterus akodontius thomas, 1921.

Type Locality: Caraguatay (Misiones, argentina).

Action Proposed: restrict this type locality to Puerto Caraguatay (26° 37’ S, 54° 46’ W, 192 m, Montecarlo, Misiones).

Sigmodontine Taxa Affected: Thomasomys pictipes Osgood, 1933.

Type Locality: río Paranay (Misiones, argentina).

Action Proposed: restrict this type locality to the confluence of the río Paranay Guazú with the río Paraná (26° 41’ S, 54° 49’ W, 122 m, Libertador General San Martín, Misiones).

Sigmodontine Taxa Affected: Oxymycterus misionalis Sanborn, 1931.

Type Locality: río de Oro (Chaco, argentina).

Action Proposed: None. the río de Oro is located in the northeastern portion of Chaco Province.

Sigmodontine Taxa Affected: Scapteromys chacoensis Gyldenstolpe, 1932.

Type Locality: Otro Cerro (Catamarca, argentina).

Action Proposed: None. Otro Cerro is located at 28° 45’ S, 66° 17’ W (2023 m, Capayán, Catamarca).

Sigmodontine Taxa Affected: Phyllotis ricardulus thomas, 1919; Graomys edithae Thomas, 1919; Akodon alterus thomas, 1919; Akodon orbus thomas, 1919; and Reithrodon caurinus thomas, 1920.

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Type Locality: “Norte de Entre ríos” (Entre ríos, argentina).

Action Proposed: restrict this type locality to São Gabriel (30° 19’ S, 54° 19’ W, 118 m, rio Grande do Sul, Brazil).

Sigmodontine Taxa Affected: “agreste” [= Mus? azarae Fischer, 1829], and “Blanco Debaxo” [= Mus? dubius Fischer, 1829].

Type locality: Isla Ella (Buenos aires, argentina).

Action Proposed: Identify location of Isla Ella as 34° 22’ S, 58° 38’ W (Primera Sección Delta del Paraná, tigre, Buenos aires).

Sigmodontine Taxa Affected: Deltamys kempi thomas, 1917; Oligoryzomys delticola Thomas, 1917; Akodon arenicola hunteri thomas, 1917; and Scapteromys aquaticus thomas, 1920.

Type Locality: Neuquén, río Limay (Neuquén, argentina).

Action Proposed: None. Neuquén is located at 38° 57’ S, 68° 04’ W (259 m, Confluencia, Neuquén).

Sigmodontine Taxa Affected: Akodon neocenus thomas, 1919.

Type Locality: Valle del Lago Blanco or Valle/region Koslowsky (Chubut, argentina).

Action Proposed: restrict this type locality to Estancia Valle Huemules (45° 57’ S, 71° 31’ W, río Senguerr, Chubut).

Sigmodontine Taxa Affected: Akodon suffusus thomas, 1903; Akodon (Chelemys) vestitus thomas, 1903; Oryzomys magellanicus mizurus thomas, 1916; and Akodon iniscatus thomas, 1919.

aCKNOWLEDGMENtS

this contribution was possible thanks to many persons and institutions, who freely provided information, insights, or help in different ways: B. Patterson (information about C. Sanborn collections), a. aguado, and J. Williams (information about J. Koslowsky), J. Vezub, r. Hora, E. Miguez, P. Pereyra, G. Galafassi, H. Venencio, staff of La reconquista Museum, M. Cirignoli, J. alvarez Gelvez, staff of Dirección de Geodesia Departamento de Investigación Histórica y Cartográfica, staff of the Dirección Provincial de Catastro del Ministerio de Economía de la provincia de Buenos aires (history of the Isla Ella Fruit and Forest Company and search of Isla Ella), J. Contreras (discussion about Norte de Entre ríos), C. Meister (cartographic support), staff of Instituto Geográfico Militar library (cartographic support), and staff of Museo de La Plata library (cartographic support). the fieldwork involved in this research was funded by S. Steppan (NSF DEB-0108422) and Universidad Nacional de La Plata. Special thanks to D. Podestá and C. Galliari for their help during the Otro Cerro, río de

Pardiñas et al.: Problematic Type Localities for Sigmodontine Rodents 409

Oro, and Higuerilla searches. Suggestions and stylistic assistance provided by D. Kelt and an anonymous reviewer greatly improved this work. this contribution was partly funded by the Consejo Nacional de Investigaciones Científicas y técnicas and is part of the research Project 487 (Universidad Nacional de la Patagonia San Juan Bosco).

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