Mastozoología Neotropical, 26(1):99-116, Mendoza, 2019 Copyright ©SAREM, 2019 Versión on-line ISSN 1666-0536 http://www.sarem.org.ar https://doi.org/10.31687/saremMN.19.26.1.0.17 http://www.sbmz.com.br

Artículo

RICHNESS, ENDEMISM AND CONSERVATION OF SIGMODONTINE IN ARGENTINA

Anahí Formoso1 and Pablo Teta2

1 Centro para el Estudio de Sistemas Marinos (CESIMAR-CENPAT-CONICET). Puerto Madryn, Chubut, Argentina. 2 División Mastozoología, Museo Argentino de Ciencias Naturales “Bernardino Rivadavia” Buenos Aires, Argentina. [Correspondence: Pablo Teta ]

ABSTRACT. Sigmodontine rodents, with 86 genera and ~430 living , constitute one of the most successful radiations of Neotropical . In this contribution, we studied the distributional ranges of 108 sigmodontine species in Argentina. Our objectives were (i) to establish geographical patterns of species richness and endemism, and (ii) to evaluate the regional conservation status of these taxa. We constructed a minimum convex polygon for each species, using information from literature and biological collections. Individual maps were superimposed on a map of Argentina divided into cells of 25 km on each side. For each cell, we calculated the species rich- ness, which varied between 1 and 21 species, and its degree of endemism, which fluctuated between 0.001 and 3.28. There were 30 species of sigmodontine rodents distributed almost exclusively in Argentina, most of them restricted to forested areas (Southern Andean Yungas) or to arid and semiarid environments (High and Low Monte and Patagonian Steppe). Areas with high species richness and endemism scores corresponded grossly with the Southern Andean Yungas, the Humid Chaco plus the Paraná flooded savannas, the Alto Parana Atlantic forests plus the Araucaria moist forests, the High Monte and the ecotone between the Patagonian steppe and the Valdivian temperate forests. A reassessment of the conservation status of sigmodontine rodents distributed in Argentina retrieved 2 extinct species, 7 endangered, 7 vulnerable, 6 near threatened and 13 data deficient. These numbers suggest a much more serious situation than the expressed by previous evaluations, highlighting the urgent need to establish conservation measures for the protection of this group.

RESUMEN. Riqueza, endemismo y conservación de roedores sigmodontinos en Argentina. Los roedores sigmodontinos, con 86 géneros y ~430 especies vivientes, constituyen una de las radiaciones más exitosas de mamíferos neotropicales. En esta contribución, estudiamos los rangos de distribución de 108 especies de sigmo- dontinos en Argentina. Nuestros objetivos fueron (i) establecer patrones geográficos de riqueza de especies y endemismo y (ii) evaluar el estado de conservación regional de estos taxones. Construimos un polígono convexo mínimo para cada especie, utilizando información de la literatura y colecciones biológicas. Los mapas indivi- duales fueron superpuestos en un mapa de Argentina dividido en celdas de 25 km de lado. Para cada celda, calculamos la riqueza de especies, que varió entre 1 y 21, y su grado de endemismo, que fluctuó entre 0.001 y 3.28. Hubo 30 especies de roedores sigmodontinos distribuidos casi exclusivamente en Argentina, la mayoría de ellos restringidos a áreas boscosas (Yungas andinas del sur) o a ambientes áridos y semiáridos (Monte alto y bajo y Estepa Patagónica). Las áreas con mayor riqueza de especies y valores más altos de endemismo se correspondieron groseramente con las Yungas andinas del sur, el Chaco húmedo más las Sabanas inundadas de Paraná, el Bosque Atlántico del Alto Paraná más los Bosques húmedos de araucaria, el Monte alto y el ecotono entre la Estepa Patagónica y los Bosques templados valdivianos. Una reevaluación del estado de conservación de los roedores sigmodontinos distribuidos en Argentina recuperó 2 especies extintas, 7 en peligro, 7 vulnera- bles, 6 casi amenazadas y 13 con datos deficientes. Estas cifras sugieren una situación mucho más grave que la

Recibido 13 abril 2018. Aceptado 24 diciembre 2018. Editor asociado: G. D’Elia 100 Mastozoología Neotropical, 26(1):99-116, Mendoza, 2019 A. Formoso & P. Teta http://www.sarem.org.ar - http://www.sbmz.com.br expresada en evaluaciones anteriores, destacando la necesidad urgente de establecer medidas de conservación para la protección de este grupo.

Key words: , endangered species, extinction, Muroidea, Rodentia.

Palabras clave: Cricetidae, especie en peligro, extinción, Muroidea, Rodentia.

INTRODUCTION most successful radiations of Neotropical mam- mals (Parada et al. 2015, Maestri et al. 2016). During the last 500 years, almost 100 species of Species of this group are mostly distributed in mammals became extinct, species being and, to a lesser extent, in Middle more than a half of them (Turvey 2009). Only and North America (D’Elía & Pardiñas 2015). in South and Central America, including both Sigmodontine rodents occur from the humid continental and insular areas, ~32 species of rainforests of the Amazon to the extremely rodents disappeared in the last five centuries dry desert of Atacama (Maestri & Patterson (Turvey 2009, Teta et al. 2014). If we consider 2016). Their diets are usually omnivorous, but rodent extinctions across the Holocene (i.e., there are species almost strictly herbivorous, the last 10 000 years), the global number of insectivorous and fungivorous (Maestri et al. lost species ascends to 115 (Turvey 2009). This 2016). Despite their generalized morphology, scenario contradicts the generalized perception sigmodontine rodents occupy from arboreal that rodents lack major conservation problems to cursorial, fossorial or semiaquatic niches (Lacher et al. 2017). (D’Elía & Pardiñas 2015). Like other small mammals, rodents play a At least 108 species of sigmodontine rodents fundamental role in trophic chains acting as have been recorded for Argentina, including prey of other vertebrates (Jaksic 2002). Many representatives of nine tribes and some incer- rodent species occupy specialized ecological tae sedis taxa (Patton et al. 2015, Teta et al. niches, contributing to energy and nutrient flow 2018). The tribes and Oryzomy- and providing important functions to the eco- ini were best represented in subtropical and systems, such as soil tilling and seed dispersal temperate regions, both in forested and open (Lacher et al. 2017). This situation contrasts environments. Abrotrichini and Phyllotini pre- with the lack of knowledge about this group, dominated towards high latitudes and Andean especially regarding the conservation status of ranges. Phyllotini was also dominant in arid their different members. A number of factors to semiarid open areas of western Argentina, negatively influences over this situation, making where it reached its greatest diversity. Finally, it even more serious. On one hand, rodents Andinomyini and Euneomyini were distributed are one of the most diverse and at the same mostly in high mountain areas, Reithrodontini time least known groups of Mammalia, with was widespread along open grassy areas and many species that are known only from the Thomasomyini and those taxa considered as type specimen or series and sometimes from incertae sedis occurred mostly in forested collections carried out more than 100 years habitats (cf. Patton et al. 2015). ago (Amori et al. 2016). On the other hand, In this paper, two main issues were addressed. rodents are not charismatic species, attracting First, we evaluated geographical patterns of spe- little attention from researchers and officials cies richness and endemism of sigmodontine responsible for directing scientific resources rodents in Argentina. Second, and based on and funding (Fleming & Bateman 2016). the analysis of the available data, we proposed Sigmodontine rodents, with 86 living gen­era new conservation status for the species in this and ca. 430 living species, constitute one of the group at the regional level, demonstrating a CONSERVATION OF ARGENTINIAN SIGMODONTINES 101 more serious regional situation than previ- Endemism was calculated both as categorical and ously reported. continuous variables. First, we considered a taxon as endemic to Argentina if it occurs mostly (> 98% of its MATERIALS AND METHODS distributional range) or exclusively within the limits of the country. The rationale behind this procedure Argentina covers an area of almost 2.8 million km2, is that the conservation status of these species would including both continental and insular portions. depend almost exclusively on the policies adopted This country is the second largest in the Neotropics by the Argentinean legislative bodies at national or and the eight largest in the world (Real et al. 2003). regional levels. Secondarily, we calculated endemism Although the usage of political divisions to address as a continuous variable, following the equation of biogeographical issues is often criticized because Kryštufek and Griffiths2002 ( , see also Kryštufek patterns and processes responsible for biological et al. 2009): Ej = ∑1/Ai, for all i included in Sj, diversity do not recognize artificial boundaries where Ai is the number of cells for every species (such as the frontiers among countries), we decided with i = 1 to n (the maximum number of species). to choose this approach because the application of In each geographical cell j is the set of species Sj conservation measures is usually conducted at re- found within it. The sum of the weights of 1/Ai for gional or national administrative levels (Real et al. every species found in set Sj produced an overall 2003). In addition, it is widely accepted that political measure of endemism Ej. The contribution made limits shape human activities and are not always by a species is constant for each cell (Nott & Pimm completely arbitrary; in fact, political boundaries 1997, Kryštufek & Griffiths 2002). Species with range sometimes coincide with natural barriers (e.g., the distributions above 106 km2 were excluded from the Andean Cordillera to the west of Argentina), being analysis, since their individual contribution to a cell suitable to detect some natural phenomena (Real would be < 10-3. et al. 2003). The scheme of ecoregions used in this Areas with high levels of biodiversity were consid- work follows Olson et al. (2001). ered as “hotspots,” defining them as those cells with Distributional data for 108 species of sigmodon- high species richness or high endemism scores, the tine rodents inhabiting Argentina were taken from cut-off point being the upper quartile (i.e. the top Patton et al. (2015) and updated when necessary 25% squares; for a similar procedure, see Kryštufek (Jayat et al. 2016, Pardiñas et al. 2016a, Pardiñas et & Griffiths2002 ). Expressed in this way, this is al. 2016b, Teta et al. 2017, Teta and D’Elía 2017). the inverse of the definition of rarity proposed by follows Teta et al. (2018). Point data for Gaston (1994). Those cells with high species rich- marginal records of each species were imported ness plus high endemism scores were considered into Geographic Information System (GIS) and as “top hotspots.” The statistical significance of “top transformed to minimum convex polygons (= ex- hotspots” was tested using the Getis-Ord Gi* tool tent of the occurrence). Distributional ranges were of ArcGIS (Getis & Ord 1992). This statistic allows calculated for each taxa, except for those with the identification of “hotspots”, higher or lower in only 1 locality record (IUCN Species Survival magnitude than one might expect to find by chance; Commission 2012). The polygons for those species in addition, this method compares the value for a exceeding the geographic limits of Argentina were given observation with locations in the neighbor- clipped and the range size in km2 (area) for each hood, thus providing a more explicit consideration species was calculated. Complete lists of locali- of space (Nelson & Boots 2008). ties were compiled only for those species with an Global conservation status were taken from the extent of occurrence less than 20 000 km², as this IUCN Red List (2011). New species assessments were value, combined with the number of localities, based on the categories and guidelines defined by demography and threats, is used as a cut-off point the IUCN for regional level (IUCN Species Survival by the UICN to separate between threatened and Commission 2012). This methodology considers the not-threatened categories. number of known localities in which a certain species For calculating species richness (S), a grid with was documented and the extent of its distribution, square cells of 25 km2 was created and the number plus additional information (if available) about de- of species in each cell was counted. Data projec- mographic parameters (see IUCN Species Survival tions and calculations were carried out using the Commission 2012). For some species, preliminary software ArcGIS Desktop 10.5.1 (ESRI 2017) and categorizations were changed as long as there was the projected coordinate system WGS84/UTM zone a possibility of a “rescue effect” from populations 19S (EPSG: 32719). outside the region (see IUCN Species Survival Com- 102 Mastozoología Neotropical, 26(1):99-116, Mendoza, 2019 A. Formoso & P. Teta http://www.sarem.org.ar - http://www.sbmz.com.br mission 2012). With few exceptions, species based Humid Chaco, Alto Parana Atlantic forests and on historical records or documented for only one or High Monte) and those characterized by the two localities (e.g., Andalgalomys pearsoni, presence of endemic species (e.g., Delta del amoenus), with dubious taxonomic status (e.g., Paraná, southeastern Mendoza, southwestern Graomys edithae), or both (e.g., Buenos Aires) (Fig. 3). The Spearman correla- guarani, B. misionensis, B. paradisus) were considered tion coefficient between species richness and as Data Deficient. endemism scores was 0.71 (p < 0.0001) RESULTS “Hotspots” were defined as those cells with species richness ≥ 10 or with an endemism score Geographical range sizes for 108 sigmo- ≥ 0.018, retrieving 1257 cells (25.8% of the total dontine species varied from 67.18 km2 to surface of Argentina); of these, 730 (14.9% of 1 439 529.18 km2, representing from 0.0024% the total surface of Argentina) are considered as to 51.77% of the total surface of Argentina “top hotspot”, since they outperformed in both (Table 1). Only 16 species have distributional parameters simultaneously (Fig. 4). Statistical ranges that covered more than 20% of the significance of “top hotspots” is depicted on total area of the country, while the other 92 Supplement 1. species occupied less than 19% each, with The reassessment of the conservation status 52 species (this is 48.14% of total number of Argentinean sigmodontine rodents retrieved of species) occurring in an area smaller than 2 Extinct species (1 Akodontini, 1 ), 1% (Table 1). Species with a range size less 7 Endangered (3 Akodontini, 4 Phyllotini), than 5501.58 km2 (= upper limit for the lower 7 Vulnerable (2 Akodontini, 1 Oryzomyini, quartile of the species ranges annotated in 1 Phyllotini, 3 incertae sedis), 6 Near Threat- Table 1) were considered rare (N = 27), accord- ened (4 Akodontini, 2 Phyllotini), 73 Least ing to the definition of Gaston (1994). Species Concern and 13 Data Deficient (see Table 1; richness varied between 1 and 21 species per Fig. 5). Endangered species were mostly grid cell (mean = 8.14). The highest values (i.e., distributed in forested, arid to semiarid, and S ≥ 14) mostly corresponded to five main areas: grassy environments (Fig. 6). Justification for 1) Southern Andean Yungas, 2) Humid Chaco each individual case is provided on Table 2. plus Paraná flooded savannas, 3) Alto Parana Atlantic forests plus Araucaria moist forests, 4) DISCUSSION High Monte, and 5) the ecotone between the Patagonian steppe and the Valdivian temperate The study and comprehension of geographical forests (Fig. 1). patterns of species richness and endemism is There are 30 species almost exclusively dis- crucial for the conservation of biodiversity (e.g., tributed in Argentina, 13 belonging to Phyl- Ceballos et al. 2005). Understanding how bio- lotini, 13 to Akodontini, 2 to Abrotrichini and diversity is changing and responding to global 2 to Oryzomyini. Most species in this category change allows us to update, when necessary, were distributed towards the western portion of the conservation status of species, in order to the country (most cells ≥ 7 endemic species), contribute preserving their populations (e.g., both in forested areas (e.g., Southern Yungas) Lacher et al. 2017; this work). In this work, we and open, arid to semiarid, areas (e.g., High analyzed the geographic distribution, species and Low Monte, Patagonian steppe) (Fig. 2). richness, endemism, and conservation status The distributional range size of endemic spe- of 108 sigmodontine rodents that inhabit Ar- cies varied between 67.18 and 894 733.81 km2, gentina. The main results of our contribution with almost the half of them being distributed are: (i) species richness, endemism scores and in less than 1000 km2 (Table 1). “hotspots” were moderately correlated and Endemism scores varied between 0.001 and correspond to five main areas (see below); 3.28 (mean = 0.02). Highest values for this (ii) almost half of the studied species occupies, variable mostly correspond to cells with high each, less than 1% of the country surface; species richness (i.e, Southern Andean Yungas, (iii) the re-evaluation of conservation status CONSERVATION OF ARGENTINIAN SIGMODONTINES 103 EX EX NT NT DD DD DD DD DD DD DD DD DD This work This EN B1ab(iii) EN EN B1ab(iii) EN EN B1ab(iii) EN B1ab(iii) EN B1ab(iii) EN EN B1ab(iii) EN EN B1ab(iii) EN EN B1ab(iii) EN VU B1ab(iii) VU VU B1ab(iii) VU LC EX DD DD DD DD DD DD DD DD DD DD not evaluated not not evaluated not evaluated not evaluated not not evaluated not not evaluated not not evaluated not evaluated not VU B1a,b(iii) VU EN B1a,b(i, iii) B1a,b(i, EN VU B2ab(ii, iii) B2ab(ii, VU Teta & Pardiñas 2012 Pardiñas & Teta LC LC LC LC LC LC LC NT NT DD DD DD DD DD DD EN B1ab(iii) B1ab(iii) EN Status IUCN Status not evaluated not evaluated not not evaluated not not evaluated not not evaluated not evaluated not evaluated not ) 2 ------141.09 957.50 734.37 848.81 329.84 523.57 67.18 81.40

800.10 347.95 212.94 158.09 112.11 1 1 2 2 3 3 Rangesize (km Table1 ) and updated status (this work). (this work). status updated and 2012 ) no no no no no no yes yes yes yes yes yes yes yes yes yes yes yes yes yes yes yes yes Endemic Tribe Phyllotini Phyllotini Phyllotini Phyllotini Phyllotini Phyllotini Phyllotini Phyllotini Phyllotini Akodontini Akodontini Akodontini Akodontini Akodontini Akodontini Akodontini Akodontini Akodontini Akodontini Oryzomyini Abrotrichini Incertae sedis Incertae Incertae sedis Incertae . sp torresi Bibimys Gyldenstolpiafronto sylvanus Akodon caprinus Phyllotis Necromys amoenus Necromys lagigliai Holochilus pearsoni Andalgalomys bonaeriensis Phyllotis Akodonphilipmyersi Brucepattersonius guarani Brucepattersonius Abrothrixmanni Brucepattersoniusmisionensis paradisus Brucepattersonius Tapecomys anitae Phyllotis Necromys lilloi Necromys edithae Graomys alisosiensis Phyllotis Andalgalomys olrogi Andalgalomys Delomys dorsalis Delomys Abrawayaomysruschi Species wayku Oxymycterus Phyllotis nogalaris Phyllotis Pardiñas ( Pardiñas & Teta to according Argentina in status regional 2011 ), Sigmodontine rodents from Argentina: species list (arranged by range size), distributional range (occupied area in Argentina), global status according IUCN Red according global status in Argentina), area (occupied range size), distributional range by species (arranged list Argentina: from rodents Sigmodontine List( 104 Mastozoología Neotropical, 26(1):99-116, Mendoza, 2019 A. Formoso & P. Teta http://www.sarem.org.ar - http://www.sbmz.com.br (Table 1 cont.) 1 (Table LC LC LC LC LC LC LC LC LC LC LC LC LC NT NT NT NT DD DD DD This work This VU B1ab(iii) VU B1ab(iii) VU VU B1ab(iii) VU VU B1ab(iii) VU LC LC LC LC LC LC LC LC LC LC NT DD DD DD DD DD DD DD DD DD not evaluated not evaluated not not evaluated not EN B1a,b(i,iii) EN Teta & Pardiñas 2012 Pardiñas & Teta LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC NT Status IUCN Status not evaluated not evaluated not evaluated not evaluated not ) 2 27

529 218.94 485.62 721.12 424.05 197.29 371.96 473.83 609.83 629.02 357.79 774.58 395.39 913.79 099.79 358.87

910.30 441.05 501.58 516.52 453.30 987.47 203.65 452.50

4 5 5 5 8 8 9 9 20 10 10 12 13 16 16 16 16 16 18 18 19 19 20 20 Rangesize (km no no no no no no no no no no no no no no no no no no no no no yes yes yes Endemic Tribe Phyllotini Phyllotini Phyllotini Akodontini Akodontini Akodontini Akodontini Akodontini Akodontini Akodontini Akodontini Akodontini Akodontini Akodontini Oryzomyini Oryzomyini Oryzomyini Oryzomyini Wiedomyini Abrotrichini Abrotrichini Abrotrichini Abrotrichini Thomasomyini Castoriaangustidens Bibimys jelskii Abrothrix Abrothrixxanthorhina Oxymycterus paramensis Oxymycterus polopi Akodon illutea Abrothrix obscurus Necromys Oxymycterusquaestor legatus Tapecomys primus Tapecomys Deltamyskempi budini Akodon Eligmodontiabolsonensis Nectomyssquamipes pictipes Juliomys iheringi Brucepattersonius Calomys tener tener Calomys lanosa Abrothrix Blarinomys breviceps Blarinomys Rhipidomysaustrinus Species Euryoryzomysrussatus Akodon Oecomysfranciscorum CONSERVATION OF ARGENTINIAN SIGMODONTINES 105 (Table 1 cont.) 1 (Table LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC DD This work This LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC DD DD DD DD not evaluated not evaluated not evaluated not VU B1a,b(iii) VU VU B2ab(ii, iii) B2ab(ii, VU Teta & Pardiñas 2012 Pardiñas & Teta LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC DD Status IUCN Status not evaluated not evaluated not evaluated not evaluated not ) 2 772.76 826.38 761.20 388.69 663.55 603.11 957.81 532.97 589.27 422.57 916.52 609.23 847.53 441.52 418.34 110.28 798.17 158.90 648.73 092.69 721.24 421.67 053.83 900.07

22 23 24 26 26 33 34 42 50 52 56 57 59 61 62 63 64 66 70 72 74 78 87 91 Rangesize (km no no no no no no no no no no no no no no no no no no no no yes yes yes yes Endemic Tribe Phyllotini Phyllotini Phyllotini Phyllotini Phyllotini Phyllotini Phyllotini Phyllotini Akodontini Akodontini Akodontini Akodontini Akodontini Akodontini Akodontini Oryzomyini Oryzomyini Oryzomyini Euneomyini Euneomyini Abrotrichini Abrotrichini Andinomyini Reithrodontini Akodon fumeus Akodon Auliscomyssublimis paranaensis Akodon mordax Euneomys Thaptomys nigritaThaptomys Andalgalomysroigi tarsalis Irenomys typicus Reithrodon boliviae Calomys caenosus Akodon domorum Graomys brendae michaelseni Geoxus Sooretamysangouya Phyllotistucumanus Akodonalbiventer Akodonmontensis Salinomysdelicatus edax Andinomys lactens Necromys Oligoryzomysfornesi venustus Calomys Geoxusvaldivianus lepidus Calomys Species 106 Mastozoología Neotropical, 26(1):99-116, Mendoza, 2019 A. Formoso & P. Teta http://www.sarem.org.ar - http://www.sbmz.com.br (Table 1 cont.) 1 (Table LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC This work This LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC not evaluated not not evaluated not Teta & Pardiñas 2012 Pardiñas & Teta LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC LC Status IUCN Status not evaluated not ) 2 932.40 685.38 088.62 818.18 586.64 100.32 987.40 969.86 333.83 696.80 174.17 209.03 459.55 546.04 354.40 887.41 653.47 224.09 948.34 297.68 192.55 893.63 433.26 727.16

101 106 110 140 141 142 158 168 201 225 230 245 251 281 300 313 326 336 373 454 527 576 600 624 Rangesize (km no no no no no no no no no no no no no no no no no no no no yes yes yes yes Endemic Tribe Phyllotini Phyllotini Phyllotini Phyllotini Phyllotini Phyllotini Phyllotini Akodontini Akodontini Akodontini Akodontini Akodontini Akodontini Akodontini Oryzomyini Oryzomyini Oryzomyini Euneomyini Euneomyini Abrotrichini Abrotrichini Abrotrichini Abrotrichini Abrotrichini Eligmodontia puerulus Eligmodontia Calomyscallidus simplex simplex Pseudoryzomys Akodonsimulator lenguarum Necromys ebriosus Neotomys Eligmodontiamoreni spegazzinii Akodon edwardsii Notiomys aquaticus Oligoryzomysnigripes macronyx Paynomys toba Akodon Loxodontomysmicropus Abrothrixandina chacoensis Oligoryzomys Calomyscallosus rufus Oxymycterus Abrothrixhirta Eligmodontiamorgani olivacea Abrothrix Eligmodontiatypus iniscatus Akodon Euneomys chinchilloides Euneomys Species CONSERVATION OF ARGENTINIAN SIGMODONTINES 107 (Table 1 cont.) 1 (Table 2 7 7 6 73 13 LC LC LC LC LC LC LC LC LC LC LC LC LC This work This 1 2 4 1 60 24 LC LC LC LC LC LC LC LC LC LC LC LC not evaluated not Teta & Pardiñas 2012 Pardiñas & Teta - - 1 2 7 80 LC LC LC LC LC LC LC LC LC LC LC LC Status IUCN Status not evaluated not ) 2 186.07 263.14 173.29 573.07 529.18

956.65 486.17 469.11 702.62 211.49 599.16 293.24 733.81

090 144 211 228 439

643 712 721 783 784 844 886 894 1 1 1 1 1 Rangesize (km

no no no no no no no no no no no yes yes Endemic Tribe Phyllotini Phyllotini Phyllotini Phyllotini Phyllotini Akodontini Akodontini Akodontini Oryzomyini Oryzomyini Oryzomyini Oryzomyini Reithrodontini Holochilusvulpinus Graomys chacoensis Graomys longicaudatus Oligoryzomys griseoflavus Graomys Phyllotisxanthopygus azarae Akodon dolores Akodon Reithrodonauritus laucha Calomys lasiurus Necromys musculinus Calomys flavescens Oligoryzomys Extinct Endangered Vulnerable treatened Near Leastconcern Data deficient Data Holochilus Species 108 Mastozoología Neotropical, 26(1):99-116, Mendoza, 2019 A. Formoso & P. Teta http://www.sarem.org.ar - http://www.sbmz.com.br

Fig. 1. Map of Argentina depicting species richness of sigmodontine rodents (left) and ecoregions according to Olson et al. 2001 (right).

Fig. 2. Map of Argentina depicting the superimposed distributions of endemic species of sigmodontine rodents. CONSERVATION OF ARGENTINIAN SIGMODONTINES 109

Fig. 3. Map of Argentina depicting endemism scores for Fig. 4. Map of Argentina depicting “top hotspots” (= black sigmodontine rodents. cells) for sigmodontine rodents.

Fig. 5.Histograms of species conservation status for sigmodontine rodents as defined by Teta and Pardiñas (2012; gray bars) and this work (black bars). 110 Mastozoología Neotropical, 26(1):99-116, Mendoza, 2019 A. Formoso & P. Teta http://www.sarem.org.ar - http://www.sbmz.com.br

Fig. 6. Map of Argentina showing the super- imposed distribution of threatened species of sigmodontine rodents.

succession of habitats, and isolating populations, potentially lead speciation (mostly by allopatry) and contribute to generate high species richness and turnover (Parada et al. 2015, Maestri & Patterson 2016). In fact, species richness for sigmodontine rodents is strongly affected by elevation, which in South America is mostly dominated longitudinally by the (Maestri & Patterson 2016). In accordance with our results, most of these same areas are recognized as diverse and distinctive centers of mammalian endemism, such as the Brazilian Atlantic forest or the Monte desert (cf. Mares 1992, Amori et al. 2012, Maestri & Patterson 2016). Statistically significant “top hotspots” were broadly coincident with the Southern Andean Yungas and the Alto Parana Atlantic forests, highlighting the importance of forested environments as at the regional level revealed a more critical areas with high species richness and unique taxa situation than the one reported in a previous (Amori et al. 2012). Further analyses, incorpo- assessment (Teta & Pardiñas 2012). rating phylogenetic and functional diversity as High values of species richness and ende- additional variables, are much needed in order mism scores coincided mostly in five main to determine “hotspots” with more accuracy areas: 1) Southern Andean Yungas, 2) Humid (Mazel et al. 2018). Chaco plus Paraná flooded savannas, 3) Alto Understanding diversity implies to consider Parana Atlantic forests plus Araucaria moist not only species richness or endemism but forests, 4) High Monte, and 5) the ecotone also rarity (Halffter1994 ). Rare species are between the Patagonian steppe and the Valdiv- those that occur in a small distributional range ian temperate forests. Most of those areas are or with low abundances through large areas adjacent to hilly or mountain chains (i.e., the (Gaston 1994, Halffter1994 ). Species with small Southern Andean Yungas, the High Monte and geographic ranges are expected to be more the ecotone between the Patagonian steppe and vulnerable to habitat deterioration and other lo- the Valdivian temperate forest are close to the calized anthropogenic disturbances. Our results Andes; the Alto Parana Atlantic forests and showed that almost half of the sigmodontine Araucaria moist forests are influenced by the rodents distributed in Argentina (N = 52) oc- relatively high elevations of the Serra do Mar cupies, each, less than 1% of the total surface and adjacent high plateaus of eastern ), of the country; among these, 27 (including 18 that offers high environmental heterogeneity endemic) could be considered rare following (Amori et al. 2012). Mountain ranges, by es- the definition of Gaston (rarity being defined tablishing barriers to dispersal, creating vertical as the inverse of range size, i.e. the converse CONSERVATION OF ARGENTINIAN SIGMODONTINES 111 ------all individuals in < 10 locations; the extent and quality of its its of quality and the extent locations; 10 < in allindividuals 000 km²,000

). Use of term “location” follows the definition provided by the IUCN Speciesby IUCN the provided the definition follows term “location” of 2012 ). Use Table2 Justification Extent of occurrence 20 < of Extent declining. is habitat Extent of occurrence < 5000 km², all individuals in < 5 locations; the extent and quality of its habi its of quality and the extent locations; 5 < in occurrence km²,5000 < all of individuals Extent declining. is tat tat is declining. is tat Extent of occurrence < 5000 km², all individuals in < 5 locations; the extent and quality of its habi its of quality and the extent locations; 5 < in occurrence km²,5000 < all of individuals Extent Extent of occurrence < 5000 km², all individuals in < 5 locations; the extent and quality of its habi its of quality and the extent locations; 5 < in occurrence km²,5000 < all of individuals Extent tat is declining. is tat Extent of occurrence < 5000 km², all individuals in < 5 locations; the extent and quality of its habi its of quality and the extent locations; 5 < in occurrence km²,5000 < all of individuals Extent tat is declining. is tat Extent of occurrence < 5000 km², all individuals in < 5 locations; the extent and quality of its habi its of quality and the extent locations; 5 < in occurrence km²,5000 < all of individuals Extent tat is declining. is tat Extent of occurrence < 5000 km², all individuals in < 5 locations; the extent and quality of its habi its of quality and the extent locations; 5 < in occurrence km²,5000 < all of individuals Extent declining. is tat Extent of occurrence < 5000 km², all individuals in < 5 locations; the extent and quality of its habi its of quality and the extent locations; 5 < in occurrence km²,5000 < all of individuals Extent tat is declining. is tat Extent of occurrence < 50000 km², all individuals in < 5 locations; the extent and quality of its its of quality and the extent locations; 5 < in occurrence km²,50000 < all of individuals Extent declining. is habitat Extent of occurrence < 5000 km², all individuals in < 5 locations; the extent and quality of its habi its of quality and the extent locations; 5 < in occurrence km²,5000 < all of individuals Extent declining. is tat Extent of occurrence < 5000 km², all individuals in < 5 locations; the extent and quality of its habi its of quality and the extent locations; 5 < in occurrence km²,5000 < all of individuals Extent declining. is tat - - 81.40 112.11 800.10 1957.50 3329.84 2734.37 4910.30 2848.81 18774.58 Rangesize Status EN B1ab(iii) EN EN B1ab(iii) EN EN B1ab(iii) EN EN B1ab(iii) EN EN B1ab(iii) EN EN B1ab(iii) EN EN B1ab(iii) EN VU B1ab(iii) VU VU B1ab(iii)* VU VU B1ab(iii)* VU VU B1ab(iii)* VU 2012 ). Abrawayaomysruschi Species Akodonphilipmyersi Andalgalomys olrogi Andalgalomys Necromys lilloi Necromys Phyllotis anitae Phyllotis Phyllotis alisosiensis Phyllotis Phyllotis nogalaris Phyllotis Bibimys Castoriaangustidens Delomys dorsalis Delomys Juliomys pictipes Juliomys Justification of the threatened status of species of sigmodontine rodents from Argentina. An asterisk denotes a change in the conservation status on basedthe status in conservation the change a denotes asterisk An Argentina. from rodents species sigmodontine of of status of the threatened Justification SpeciesSurvival (see IUCN countries neighbor rescue effect Commission from Survival Commission ( SurvivalCommission 112 Mastozoología Neotropical, 26(1):99-116, Mendoza, 2019 A. Formoso & P. Teta http://www.sarem.org.ar - http://www.sbmz.com.br (Table 2 cont.) 2 (Table all individuals in < 10 locations; the extent and quality of its its of quality and the extent locations; 10 < in allindividuals its of quality and the extent locations; 10 < in allindividuals 000 km²,000 km²,000

Justification Extent of occurrence 20 < of Extent declining. is habitat occurrence 20 < of Extent declining. is habitat and the extent km²,5,000 occurrence< and is of extent its small range, its in common Although declining. is habitat its of quality its of quality and the extent km²,20,000 occurrence< and is of extent its small range, its in Rare declining. is habitat its of quality and the extent km²,20,000 occurrence< and is of extent its small range, its in Rare declining. is habitat the km²,20,000 occurrence< and is of extent its small range, its in common apparently Although declining. is habitat its of quality and extent and the extent km²,20,000 occurrence< and is of extent its small range, its in common Although declining. is habitat its of quality and the extent km²,5,000 occurrence< and is of extent its small range, its in common Although area. protected a within included is population only known The declining. is habitat its of quality Extent of occurrence < 5000 km², all individuals in < 10 locations; the extent and quality of its its of quality and the extent locations; 10 < in occurrence km²,5000 < all of individuals Extent declining. is habitat 67.18 212.94 9203.65 9452.50 5501.58 3,523.57 12721.12 16473.83 19395.39 Rangesize NT NT NT NT NT NT Status VU B1ab(iii) VU VU B1ab(iii) VU VU B1ab(iii)* VU Oecomysfranciscorum Oxymicterus wayku Oxymicterus sylvanus Akodon chacoensis Bibimys Deltamyskempi Eligmodontiabolsonensis obscurus Necromys bonaeriensis Phyllotis Species Tapecomys primus Tapecomys CONSERVATION OF ARGENTINIAN SIGMODONTINES 113 of widespread). This situation is not surpris- Argentinian specimen was secured in 1886 ing, because prevalence of rarity is a common [Pardiñas et al. 2008, Bezerra 2011]) and the phenomenon among other taxonomic groups marsh rat Holochilus lagigliai (last seen in the in different geographical regions (Gaston 1994). decade of 1950 [Pardiñas et al. 2013]). Amori In this study, species with restricted distribu- et al. (2012), from a continental perspective, tions occur mostly in the Southern Andean reported no more than one threatened spe- Yungas, the Alto Parana Atlantic forests plus cies per region across Argentina. Here, we Araucaria moist forests, and the Monte desert. documented a much different situation, with Most species in the Southern Andean Yungas up to 3-4 threatened species in some parts of and the Monte desert are, according to current the High Monte and the Alto Parana Atlantic knowledge, truly restricted (Table 1); while forests plus the Araucaria moist forests. These most of those of the Alto Parana Atlantic for- results highlight the need of regional assess- ests plus Araucaria moist forests have reduced ments, especially when this type of information distributions in Argentina (due to the small constitutes the base for future research or is range occupied by these forests in this country), used to taking regional conservation measures although most of them are relatively widespread (see IUCN Species Survival Commission 2012). along the forested portions of eastern Paraguay Most threatened species shared some fea- and southeastern Brazil (cf. Patton et al. 2015). tures that influence its conservation status, The reassessment of the conservation status such as small distributional ranges, low of Argentinean sigmodontine rodents showed a abundances or a reduced number of locality situation more critical and severe than currently records (Table 2). In most cases, these species envisioned. Our findings differ from the evalu- occur in habitats where both their extension ation carried out by Teta & Pardiñas (2012) and quality are declining, such as the Alto who reported 1 Extinct species (1 Akodontini), Parana Atlantic forests, the Araucaria moist 2 Endangered (2 incertae sedis), 4 Vulnerable forests (e.g., Castoria angustidens, Delomys (1 Akodontini, 1 Euneomyini, 2 Phyllotini) dorsalis, Juliomys pictipes) or the grasslands of and 1 Near Threatened (1 Akodontini). Despite Humid Pampas in central-eastern Argentina some methodological issues resulting in such (e.g., Deltamys kempi, Necromys obscurus). The different figures, our results suggest a more situation of the leaf-eared mice of the worrying regional situation than that formerly Phyllotis is outstanding, since at least 4 spe- reported. Regarding this point, it is impor- cies of this taxon are considered under some tant to note that all changes recorded since threatened category. Species of Phyllotis are Teta & Pardiñas (2012) are non-genuine (i.e., mostly restricted to rocky outcrops and high those resulting—for example—from improved grassy areas, having, in some cases, extremely knowledge, taxonomic changes, or correction restricted ranges (Steppan & Ramírez 2015) of earlier errors; see IUCN Species Survival in threatened environments (e.g., Southern Commission 2012), because of new available Andean Yungas). information or mistakes in the interpretation of Our work has two potential sources of bias, some guidelines (e.g., Teta & Pardiñas [2012] do one primarily related to taxonomic issues and not consider the “rescue effect” from popula- the other linked to artifacts in species mapping tions in neighbor countries or misunderstood (Kryštufek & Griffiths 2002, Isaac et al. 2004). the concept of locality according to the IUCN). For example, Andalgalomys olrogi is here cat- We found that threatened species inhabit mostly egorized as endangered; however, this rodent forested environments, but also open, arid, was included by some researchers (e.g., Diaz semiarid to temperate, steppes and grasslands. et al. 2006) into the synonymy of A. roigi, a Among those species that we considered as rare, species categorized as of least concern. For the there are 2 listed as Extinct, 7 Endangered and genus Brucepattersonius, there is at least three 3 Vulnerable (Table 1). Extinct species include species with very restricted distributional areas the swamp rat fronto (last seen in (i.e., Brucepattersonius guarani, B. misionensis, Brazil in the decade of 1990; the only known B. paradisus), which were categorized as Data 114 Mastozoología Neotropical, 26(1):99-116, Mendoza, 2019 A. Formoso & P. Teta http://www.sarem.org.ar - http://www.sbmz.com.br

Deficient due to their dubious taxonomic status cies is crucial for their conservation (Ceballos et and potential conspecificity (cf. Lanzone et al. al. 2005). The situation of the order Rodentia is 2018). Another example is the case of Phyllotis paradigmatic, because historical records suggest tucumanus from northwestern Argentina, that the species in this group are among the whose taxonomic splitting would increase the most vulnerable mammals to extinction due to endemism scores in an area already recognized human activities (Turvey 2009). Simultaneously, as a “hotspot” and also reduce the average this group includes some of the least known range sizes for the different species in this species of the world. Funding of spe- genus (see Jayat et al. 2016). Contrarily, the cific conservation projects for rodents is a first opposite effect would occur ifP. alisosiensisis step to maintaining the exceptional diversity of is included into the synonymy of P. anitae , as this order on our planet. is suggested by the available morphological and molecular evidence (see Jayat et al. 2016). ACKNOWLEDGEMENTS Regarding the species distribution datasets, our This work has benefited from the numerous discussions approach ignores that real ranges of species are held over the past years with colleagues and friends, includ- better represented as mosaics than as continu- ing G. Cassini, S. Cirignoli, E. Cuéllar, G. D’Elía, D. Flores, ous polygons in a map, which do not always N. Fracassi, C. Galliari, G. Gil, P. Jayat, C. Lanzone, S. Lu- represent the real geographic distribution of the cero, R. Ojeda, P. Ortíz, U. F. J. Pardiñas, J. Pereira, D. Po- destá and D. Udrizar. Similarly, two anonymous reviewers species. Fortunately, it is well documented that and the asociated editor (G. D’Elía) provided valuable input broad biogeographic patterns are not sensitive on an earlier version of this contribution. We also want to to grid square size (see Schall & Pianka 1978, acknowledge the help provided by P. Dell’Aciprete, I. Minoli Di Marco et al. 2013). and V. Alonso Roldán with some methodological issues. The fragmentation of habitats and the loss of biodiversity are in constant advance and LITERATURE CITED it seems to become even worse in coming Amori, G., F. Chiozza, B. D. Patterson, C. Rondinini, decades. Human activities are the main cause J. Schipper, & L. Luiselli. 2012. Species richness of habitat loss, fragmentation and species ex- and distribution of Neotropical rodents, with conservation implications. Mammalia 77:1-19. tinction, but climate change is projected to be https://doi.org/10.1515/mammalia-2012-0050 equally or even more important in the near Amori, G., G. Alari Esposito, & L. Luiselli. 2016. future (Hoffman et al.2010 , Pereira et al. 2010, Known from a handful of specimens: analyzing the Dawson et al. 2011). Global biodiversity is worldwide patterns of occurrence and conservation of rodents and shrews recorded only from the type currently under a huge pressure of change and locality. Journal of Threatened Taxa 8:8556-8563. threat (Ceballos et al. 2005). Recognizing and https://doi.org/10.11609/jott.2405.8.3.8556-8563 understanding this problem is crucial for carry- Bezerra, A. M. R. 2011. Collection records of Gyldesntolpia ing out plans for managing and protecting the planaltensis (Avile-Pires, 1972) (Rodentia, Cricetidae) suggest the local extinction of the species. Mastozoología species that more need it (Ceballos et al. 2005). Neotropical 18:119-123. For example, protected areas such as National Ceballos, G., P. R. Ehrlich, J. Soberón, I. Salazar, Parks are scattered through Argentina, covering & J. P. Fay. 2005. Global mammal conservation: ca. 13% of its total surface. This protection is what must we manage? Science 309:603-607. https://doi.org/10.1126/science.1114015 unevenly distributed, being mostly coincident Dawson, T. P., S. T. Jackson, J. L. House, C. Prentice, with areas of natural attractions or impressive & G. M. Mace. 2011.Beyond predictions: biodiversity landscapes (e.g., Alto Parana Atlantic forests, conservation in a changing climate. Science 332:53-58. Valdivian temperate forests). From our data, https://doi.org/10.1126/science.1200303 D’ Elía G. & U. F. J. Pardiñas. 2015. Subfamily it is clear that additional protected areas are Wagner, 1843. Mammals of South strongly needed in some distinctive ecoregions, America, Volume 2 Rodents (J. L. Patton, U. F. J. such as the High Monte that supports numerous Pardiñas & G. D’Elía, eds.). The University of Chicago endemic species. In many countries, and more Press, Chicago and London. Díaz, M. M., P. Teta, U. F.J . Pardiñas, & R. Barquez. especially in developing ones, funds designated 2006. Phyllotini Vorontzov, 1959. Mamíferos de to conservation are limited, even when the need Argentina: sistemática y distribución (R. Barquez, for constant updates on the knowledge of spe- CONSERVATION OF ARGENTINIAN SIGMODONTINES 115

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ONLINE SUPPLEMENTARY MATERIAL Supplement 1 https://www.sarem.org.ar/wp-content/uploads/2019/08/SAREM_MastNeotrop_26-1_Formoso-sup1.doc Fig. S1. Maps of Argentina depicting the statistical significance of “top hotspots” for sigmodontine rodents (left, species richness; right, endemism scores).