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Mastozoología Neotropical ISSN: 0327-9383 [email protected] Sociedad Argentina para el Estudio de los Mamíferos Argentina

Geise, Lena; Pereira, Luciana G.; Astúa, Diego; Aguieiras, Marcia; Lessa, Leonardo G.; Asfora, Paulo H.; Dourado, Francisco; Esberárd, Carlos E. L. TERRESTRIAL OF THE RIVER BASIN, : A TRANSITION AREA BETWEEN ATLANTIC FORESTAND CERRADO Mastozoología Neotropical, vol. 24, núm. 1, julio, 2017, pp. 95-119 Sociedad Argentina para el Estudio de los Mamíferos Tucumán, Argentina

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Artículo

TERRESTRIAL MAMMALS OF THE BASIN, BRAZIL: A TRANSITION AREA BETWEEN ATLANTIC FOREST AND CERRADO

Lena Geise1, Luciana G. Pereira1, Diego Astúa2, Marcia Aguieiras1, Leonardo G. Lessa3, Paulo H. Asfora1, 2, Francisco Dourado4, and Carlos E. L. Esberárd5,

1 Laboratório de Mastozoologia, Departamento de Zoologia, IB, Universidade do Estado do Rio de Janeiro, Rio de Janeiro, RJ, Brazil. [Correspondence: ] 2 Laboratório de Mastozoologia, Departamento de Zoologia, CB, Universidade Federal de Pernambuco Recife, PE, Brazil. 3 Laboratório de Ecologia, Departamento de Ciências Biológicas, Universidade Federal dos Vales do Jequitinhonha e Mucuri, Diamantina, MG, Brazil. 4 Centro de Pesquisas e Estudo em Desastres, Departamento de Geologia Aplicada, FGEL, Universidade do Estado do Rio de Janeiro, Rio de Janeiro, RJ, Brazil. 5 Laboratório de Diversidade de Morcegos, Departamento de Biologia , Universidade Federal do Rio de Janeiro, Seropédica, RJ. Brazil.

ABSTRACT. Here we present an extensive survey for non-volant mammals along the margins of the Jequitinhonha River, which represents a contact area between the Cerrado and Atlantic Forest biomes. Our data were ob- tained after 13 trapping expeditions (from 2005 to 2012) on both banks of the river, from its sourcetoits mouth, enriched by data obtained from a literature review and voucher specimens deposited in six collections. All collected specimens were identified through their karyotypes and/or morphology. We obtained data for 75 localities (34 at the rigth margin, 42 at the left margin; 30 in the Cerrado and 45 in the Atlantic Forest). We obtained voucher records for 91 , ca. 17% of all known Brazilian terrestrial non-volant mam- mals. Our results include range extensions for six species (Monodelphis scalops, Thylamys velutinus, Calomys mattevii, Wiedomys pyrrhorhinos, Galea spixii, Kerodon rupestris), and karyotype information for 26 species (11 marsupials and 15 ), with four new karyotypes. While most species have wide geographical range in both biomes, 26 were recorded only in the Cerrado and 19 were recorded only in the Atlantic Forest. Recorded species represent 28.7% (6.3% endemic) of the species known from the Cerrado and 21.5% (22.2% endemic) of those from the Atlantic Forest. The richness of the fauna recorded in this region supports its importance for the understanding of mammal biogeography, diversity and evolution.

RESUMO. Mamíferos terrestres da bacia do Rio Jequitinhonha, Brasil: uma área de transição entre a Flo- resta Atlântica e o Cerrado. Aqui apresentamos uma extensa pesquisa de mamíferos não-voadores ao longo das margens do Rio Jequitinhonha, região biogeográfica interessante que representa uma área de contato entre dois biomas brasileiros (Cerrado e Mata Atlântica), ainda pouco conhecidos para a fauna de mamíferos. Nossos dados foram obtidos após 13 expedições de captura (de 2005 a 2012), em ambas as margens do rio, da nascente até à foz, enriquecida por dados obtidos a partir de amostras de material depositado em seis coleções de mamíferos. Todos os espécimes coletados foram identificados através de seus cariótipos ou morfologia. Foram obtidos dados

Recibido 23 marzo 2016. Aceptado 23 febrero 2017. Editor asociado: G D’Elía 96 Mastozoología Neotropical, 24(1):95-119, Mendoza, 2017 L Geise et al. http://www.sarem.org.ar - http://www.sbmz.com.br de 75 localidades (34 na margem direita, 41 à esquerda; 30 no Cerrado e 45 na Floresta Atlântica). Obtivemos registros de 91 espécies de mamíferos não-voadores, ca. de 17% de todos os mamíferos terrestres brasileiros conhecidos. Nossos resultados incluem extensões de distribuição de seis espécies (Monodelphis scalops, Thylamys velutinus, Calomys mattevii, Wiedomys pyrrhorhinos, Galea spixii, Kerodon rupestris), e informações de cariótipo de 26 espécies (11 marsupiais e 15 de roedores), com quatro novos cariótipos. Enquanto a maioria das espécies têm ampla distribuição geográfica, 26 ocorreram apenas no Cerrado e 19 apenas na Mata Atlântica. Foram re- gistradas de 28.7% (6.3% endêmicas) de todas as espécies conhecidas para o Cerrado e 21.5% (22.2% endêmicas) da Floresta Atlântica. A riqueza da fauna registrada nesta região apoia a sua importância da compreensão na diversidade, biogeografia e evolução dos mamíferos.

Key words: Barriers. Didelphimorphia. Karyotypes. Rodentia. Species richness.

Palavras chave: Barreiras. Cariótipos. Didelphimorphia. Riqueza de espécies. Rodentia.

INTRODUCTION The Jequitinhonha River Basin appears to be a particularly interesting region because it Brazil is the largest country in the Neotropics runs into both Atlantic Forest and Cerrado. and it has the largest mammal diversity in The vegetation along the Jequitinhonha River the world, with over 700 known species, a Basin changes along the river path. The region number that has been steadily increasing in is formed by a mosaic of vegetal physiognomies the past decades (Costa et al., 2005; Paglia et presented in two biomes (Cerrado and Atlantic al., 2012). Estimates from nearly 20 years ago Forest), with the northwest part also close to the had already predicted that the actual richness Caatinga, a uniquely Brazilian biome. Thus, such of Brazilian mammal species, given the exten- an environmental heterogeneity is likely to reveal sion of the country, was by far underestimated a peculiar mammalian fauna composition, as it is (Vivo, 1997). These facts only strengthen the surrounded by regions that harbor each a distinct importance of comprehensive and exhaustive mammal composition (Lessa and Paula, 2014). surveys, particularly in those regions that have Additionally, this sequence of vegetation also been poorly sampled, in order to accurately includes contact zones, where several mammal evaluate their species composition (Patterson, species from more than one of these biomes can 2002; Costa et al., 2005; Lessa et al., 2008). be found in sympatry (Geise and Astúa, 2009). Not only is the Brazilian mammal fauna However, the actual knowledge of mammals in highly diverse, but also the country various the Jequitinhonha River Basin is still limited biomes lead to varying levels of endemism, (Lessa et al., 2008). Thus, its mammal fauna with different levels of threat to their mammals. needs to be properly diagnosed (Drumond et Approximately 11% of the Brazilian mammal al., 2005) to provide a robust biogeographic data species are listed by the IUCN as globally to further studies and more knowledge about threatened (IUCN, 2015) and 15% as nationally the biodiversity of that region. threatened by the Brazilian National Red List In this paper we report an extensive survey (MMA, 2014). Levels of endemism in Brazil are for non-volant mammals in both riverbanks high, with ca. 30% of the recorded mammal of the Jequitinhonha River Basin, representing species being endemic (Paglia et al., 2012). As formations from two biomes (Cerrado and At- a consequence of their high endemism and hu- lantic Forest), along with karyotypic analyses. man pressure, two important Brazilian biomes, We also indicate how the Jequitinhonha River Atlantic Forest and Cerrado, were designated Basin region can be considered as an important as global hotspots (Myers et al., 2000). ecotone discontinuity area. TERRESTRIAL MAMMALS OF THE JEQUITINHONHA RIVER BASIN, BRAZIL 97

MATERIALS AND METHODS interventions based on economic reorientation, focusing on technology and competitiveness, induce Study area environmental degradation and generate economic, social and cultural negative consequences (Ferreira According to the document “Região Hidrográfica and Saadi, 2009). do Atlântico Leste”, organized by the Brazilian Min- As a whole, 75 localities were considered istry of Environment (MMA, 2006), the Jequitin- in this study; of these, 30 are in the Cerrado, honha and Pardo rivers include four sub-basins: while 45 are in the Atlantic Forest (Fig. 1). Jequitinhonha 1, Jequitinhonha 2, Jequitinhonha 3 Eleven phytophysiognomies are found along the and Pardo. Here we consider the area including three Jequitinhonha River Basin (IBGE, 2012) (Appen- sub-basins (Jequitinhonha 1, 2 and 3) following the dix I); two of them are non-natives as a conse- denomination provided by Instituto Brasileiro de quence of the anthropic impact. Agrarian Activities Geografia e Estatística (IBGE, 1997), Ferreira (2011), (originally Savanna, Steppic Savanna/Seasonal For- Ferreira and Saadi (2011), Silva and Ferreira (2011) est or Savanna/Broadleaf Forest) are present in 12 and Ferreira and Silva (2012), that includes part of localities, and Secondary Vegetation and Agrarian the northeastern portion of the state Activities (originally Seasonal Deciduous Forest, and a small portion of the southeast part of Seasonal Semideciduous Forest or Broadleaf Forest) state. The basin is located between 16o - 18oS and occur in 23 localities (IBGE, 2012). For original 39o - 44oW, for a total area of 70 315 km2. Of these, phytophysiognomy of each of these localities see 66 319 km2 are in Minas Gerais (MG) and 3996 km2 Appendix I. The phytophysiognomies, Seasonal in Bahia (BA) state (IBGE, 1997). In the North, the Decidual Submontante Forest, Moist Broadleaf Jequitinhonha River Basin borders the sub-basin of Forest of Lowlands, Mangroves, Upper Montane Pardo River, and in the South, the sub-basins of the Vegetational Refuges, Arboreous Steppic Savanna Mucuri River and the South Bahia Coast (MMA, and Savanna/Broadleaf Forest are found at a single 2006). The source of the Jequitinhonha River lies locality each. Four localities were at Arboreous near City (MG), at an elevation of 1300 m Savanna, five Grassy-Woody Savanna, 25 Savanna/ and drains into the Atlantic Ocean at Belmonte Seasonal Forest (IBGE, 2012) (Appendix I). City (BA). Some of the sampled localities are lo- cated outside the limits of the Jequitinhonha River Field work drainage basin because for data collection and field work we included all municipalities that according From those 75 localities where mammals were to the Brazilian Institute of Geography and Statistics recorded in this study (see above), small mammals (IBGE, 2016) are completely or partially inside the were trapped in 13. These localities were chosen to basin limits. cover both left (localities 2, 5, 44, 45, and 73) and Along the 920 km path of the Jequitinhonha River, right (localities 6, 7, 8, 9, 43, 49, 53 and 75) banks of 760 are in the Minas Gerais state and 160 in the the Jequitinhonha River (Fig. 1, Appendix I), were state of Bahia (Ferreira, 2009). From the river source vegetation was more preserved or where field activi- in the Cerrado down to its mouth in the Atlantic ties could be carried on without security problems for Forest, the climatic characteristics are variable, from researchers. Traps were set differently according to humid to semiarid climates, and different vegeta- topography and presence of trees (arboreal traps set tional formation are recorded (IBGE, 1997). Rains at up to 3 m high), mostly placed in lines ca. 15 to are usually concentrated from October to March, 20 m apart one from another. Three live traps models with 50% of all the rain occurring from December were used (Sherman®, Tomahawk® and pitfall) with to February (Gonçalves, 1997). According to Silva banana and manioc pieces, peanut butter and bacon and Ferreira (2011), in three localities, precipitation bits as bait in live-traps in areas of Atlantic Forest, occurs in a very irregular pattern, Araçuaí, in the and peanut butter, orange or pineapple pieces and middle region of the basin, with an annual average cotton wool soaked in cod liver oil in dryer areas of 766 mm, in , closer to the source in Cerrado. In addition to the collected specimens of the river, with 1050 mm, and in , of small mammals (marsupials and rodents), we closer to the river mouth, with 860 mm. The social also collected road-killed , obtained skulls and environmental scenario of Jequitinhonha River from farmers, searched for tracks and made visu- Basin presents a dispersed population; the State alization in the field. Table 1 indicates the name, 98 Mastozoología Neotropical, 24(1):95-119, Mendoza, 2017 L Geise et al. http://www.sarem.org.ar - http://www.sbmz.com.br

Fig. 1. Mammal recording localities in the Cerrado and Atlantic Forest within and surrounding areas in the Jequitinhonha River Basin. Biome boundaries are according to IBGE (2016). White triangles indicate localities where trapping effort was carried out during the present study. For number of localities see Appendix I.

coordinates and trapping effort for each locality. interspersed with forest patches in the southern part Tissue (liver) samples, fixed in ethanol, were col- of the PARNASV. lected from all specimens, as well as ectoparasites. Locality #6 - Fazenda Santa Cruz (Fig. 2B). The All specimens were prepared as skins, skulls, and vegetation is Grassy-Woody Savanna, in the Cerrado partial skeletons. Specimens were collected under (IBGE, 2012). Traps were set in open grasslands sample permits from ICMBio granted to L. Geise with rocky outcrops, in gallery forests and open (IBAMA 0201.009374/02-86 and SISBIO 598633), vegetation formations. Some trapping lines were to L.G. Lessa (SISBIO 19790-1 and 42892-1) and along characteristic Cerrado formation, with many to C.E.L. Esberárd (SISBIO 10356-1). pequi trees (Caryocar brasiliense, Caryocaraceae). Below we provide a short description of those Locality #7 – Parque Estadual do Rio Preto (PERP) localities we surveyed. Localities are numbered ac- (Fig. 2C). The vegetation is Grassy-Woody Savanna, cording to Appendix I. in the Cerrado (IBGE, 2012). The PERP is located in Locality #2 - Pousada Rural Recanto do Vale the southern area of the Espinhaço Mountain Range (Fig. 2A). The vegetation is Grassy-Woody Savanna, with a mosaic of vegetation physiognomies. Traps in the Cerrado (IBGE, 2012). Traps were set in lines were settled in areas of riparian savanna, Cerrado along a small creek, on sandy ground and near a stricto sensu and open grasslands. small waterfall, on the rocks (“lajeiro”). Locality #8 - Fazenda Sumidouro (Fig. 2D). The Locality #5 – Parque Nacional das Sempre Vivas vegetation is Savanna/Seasonal Forest, in the Cer- (PARNASV). The vegetation is Upper Highlands rado (IBGE, 2012). Traps were set in fragments of Vegetational Refuges, in the Cerrado (IBGE, 2012). forest formation surrounded by pasture, some lines Traps were set in lines along a rupestrian fields area along the Araçuaí River. TERRESTRIAL MAMMALS OF THE JEQUITINHONHA RIVER BASIN, BRAZIL 99 620 360 720 105 724 683 200 878 1200 1125 9216 1325 1250 1370 (trap-nights) Trapping Effort Effort Trapping March 2011 March 2012 March 2005 March 2005 March 2012 March 2005 March 2008 March 2008 March September 2007 September 2007 September 2007 September March-April 2011 March-April Period of fieldwork of Period February to December to February 2014 November 2009 to January 2014 January to 2009 November 90 875 278 796 150 1200 1105 823-920 756-785 788-802 487-529 771-923 250– 263 250– Altitude (m) Altitude ), coordinates, altitude or altitude range where traps were settled, period were traps where range altitude or altitude ), coordinates, Appendix I Appendix Table1 Coordinates 17°55'02''S, 43°47'11''W 17°55'02''S, 43°20'25''W 18°05'20''S, 15°51'28''S, 39°24'38.2''W 15°51'28''S, 16°39'42.6''S, 41°52'5.1''W 16°39'42.6''S, 41°14'57.6''W 16°43'7.6''S, 18°23'43.2''S, 43°32'25.5''W 18°23'43.2''S, 43°23'18.8''W 18°16'16.2''S, 43°14'35.2''W 18°11'34.4''S, 43°10'26.2''W 18°05'32.5''S, 41°49'35.5''W 16°36'46.1''S, 41°47'23.4''W 16°34'56.4''S, 41°36'48.6''W 17°07'18.6''S, 39°06'49.6''W 16°21'11.6''S, Locality Name / Number (#) LocalityNumber / Name #2 / Vale do Recanto Rural Pousada #5 / (PARNASV) Vivas Sempre das Nacional Parque Cruz #6 / Santa Fazenda #7 / (PERP) RioEstadualdo Preto Parque #8 / Sumidouro Fazenda #9 / Quente Água Pousada Ilha#43 / Fazenda #44 / Marília Dona Fazenda Galiléia #45 / Fazenda #49 / Palmares Fazenda #53 / Pehy Anga Fazenda #73 / Futurosa Fazenda Estação Veracel Natural ReservaPatrimônio do Particular #75 / Veracel) (RPPN Trapping localities of the present study, including locality name, number (see number locality name, including study, the localitiespresent of Trapping effort. and trapping the fieldwork of 100 Mastozoología Neotropical, 24(1):95-119, Mendoza, 2017 L Geise et al. http://www.sarem.org.ar - http://www.sbmz.com.br

Fig. 2. Views of some surveyed localities: A) Pousada Rural Recanto do Vale (Locality 2) – Grassy-Woody Savanna; B) Fa- zenda Santa Cruz (Locality 6) – Grassy-Woody Savanna, where a Campo Limpo can be observed; C) Parque Estadual do Rio Preto (Locality 7) – Grassy-Woody Savanna with an overview of Cerrado lato sensu; D) Fazenda Sumidouro (Locality 8) – Savanna/Seasonal Forest, where a rocky margin of a small creak with arbustive vegetation can be observed; E) Pousada Água Quente (Locality 9) – Savanna/Seasonal Forest, depicting characteristic seasonal forest; F) Fazenda Ilha (Locality 43) – Savanna/Seasonal Forest, showing an arbustive and thorny vegetation; G) Fazenda Dona Marília (Local- ity 44) – Savanna/Seasonal Forest, showing an arbustive and thorny vegetation with some cactus; H) Fazenda Galiléia (Locality 45) – Savanna/Seasonal Forest – with the same vegetation observed in locality 44; I) Fazenda Palmares (Locality 49) – Secondary Vegetation and Agrarian Activities (originally Seasonal Semideciduous Forest), showing a pasture, with forest on the top of the hill; J) Fazenda Anga-Pehy (Locality 53) – Secondary Vegetation and Agrarian Activities (originally Seasonal Semideciduous Forest) = with the same vegetation observed in locality 49. All photos taken by Lena Geise, except that of locality 7, taken by Leonardo G. Lessa. TERRESTRIAL MAMMALS OF THE JEQUITINHONHA RIVER BASIN, BRAZIL 101

Locality #9 - Pousada Água Quente (Fig. 2E). The Locality #75 - Reserva Particular do Patrimônio vegetation is Savanna/Seasonal Forest, in the Cerrado Natural (RPPN), Estação Veracel. The vegetation is (IBGE, 2012). Trapping was carried out in Gallery Moist Broadleaf Forest of Lowlands, in the Atlan- Forest, with traps along the river, over rocks or tic Forest. All trapping occurred in the preserved sand and some in the river. Other lines were set in forest area. a Forest, at the slope of a small hill, a very humid area. Grasses covered the beginning of the line. The Small mammal species identification “campo sujo” was at another slope, with rocky and (karyotypes and morphology) sandy soil, some areas recovering from fire. Karyotypes were prepared in the field for all ro- Locality #43 - Fazenda Ilha (Fig. 2F). The vegetation dent specimens and at least one specimen of each is Savanna/Seasonal Forest, in the Atlantic Forest collected species of marsupial. Metaphases were according to the map (IBGE, 2012), but local ob- obtained with in vitro bone marrow culture accord- servation showed that it is clearly in the Cerrado. ing to Geise (2014). Conventional coloration with Lines were set in typical arboreal vegetation (close Giemsa 5% was used to observe diploid (2n) and thorny arbustive vegetation, with small sparse trees), fundamental (FN, excluding sexual chromosomes) with sandy soil. numbers and chromosome morphology. Rodents Locality #44 - Fazenda Dona Marília (Fig. 2G). The were identified to species level by comparison with vegetation is Savanna/Seasonal Forest, in the Atlantic voucher specimens deposited in collections, species Forest according to the map (IBGE, 2012), but local descriptions, and karyological analysis. Morphologi- observation showed that it is clearly in the Cerrado. cal characters (skin and skull) were considered for Trapping lines were in open arboreal vegetation with species identification in comparison to previous an open bush formation. Trapping lines were along descriptions for Didelphimorphia. Nomenclature the Jequitinhonha River. follows Wilson and Reeder (2005), Gardner (2008), Locality #45 - Fazenda Galiléia (Fig. 2H). The veg- Melo and Sponchiado (2012), Gurgel-Filho et al. etation is Savanna/Seasonal Forest, in the Atlantic (2015) and Patton et al. (2015). Forest according to the map (IBGE, 2012), but local observation showed that it is clearly in the Cerrado. Museum collections and literature review General vegetation where lines were mounted is ar- Medium and large mammal (Cingulata, Pilosa, Pri- boreal vegetation (close thorny arbustive vegetation, mates, Lagomorpha, Carnivora, Perissodactyla, and with sparse low trees), with sandy soil. Trapping was Cetartiodactyla) records, in addition to other small carried out during the rainy season, so vegetation mammal species, were obtained from museum col- was green. lections (Museu Nacional do Rio de Janeiro, Museu Locality #49 - Fazenda Palmares (Fig. 2I). The veg- de Zoologia da USP, Coleção do Laboratório de etation is Secondary Vegetation and Agrarian Activi- Mastozoologia e Manejo da Fauna do Departamento ties (originally Seasonal Semideciduous Forest), in de Zoologia, Universidade Federal de Minas Gerais, the Atlantic Forest (IBGE, 2012). Traps were set in Museu de Ciências Naturais da PUC Minas, Museu disturbed forest fragments located on mountaintops de Zoologia da Universidade Federal de Viçosa and surrounded by pastures. Coleção de Mastozoologia da Universidade Federal Locality #53 - Fazenda Anga Pehy (Fig. 2J). The dos Vales do Jequitinhonha e Mucuri). vegetation is Secondary Vegetation and Agrarian Literature review was carried out to complete all Activities (originally Seasonal Deciduous Forest), in mammal species occurrence. We do not consider the Atlantic Forest (IBGE, 2012). Traps were set in specimens of cursor that were not identified disturbed forest fragments located on mountaintops through genetic techniques, as identification based surrounded by pastures, one along a creek, another solely on skull and skin morphology is not reliable around a swampy area covered by graminae. Another for this species (Geise, 2012). vegetation was a dry altered forest, locally called “Mata da Chapada”. Localities characterization, species Locality #73 - Fazenda Futurosa. The vegetation is geographic pattern distribution Agrarian Activities (original vegetation was Savanna/ and classification of endangered species Broadleaf Forest), in the Atlantic Forest (IBGE, 2012). The sampled area is a forest fragment sur- Coordinates and altitude for each locality were rounded by eucalyptus plantations and the “Cabruca”, obtained at each capture location and gazetteers a cacao trees plantation grown under the Atlantic online (www.geonames.org, www.falingrain.com/ Forest canopy, also surrounded by pasture. world andwww.splink.cria.org.br/geoloc), using SAD 102 Mastozoología Neotropical, 24(1):95-119, Mendoza, 2017 L Geise et al. http://www.sarem.org.ar - http://www.sbmz.com.br

69 23 K Datum. Vegetation classification of each increases the species distribution eastwards in locality is according to the IBGE (2012). 190 km from , MG (Gurgel-Filho For geographic range increase of species, we et al., 2015) to Fazenda Dona Marília-, searched each species known geographic distribution MG (locality 44). Galea spixii distribution in- provided by Gardner (2008), Geise and Astúa (2009), creased southwards in 178 km from Campos Cáceres (2012), Gurgel-Filho et al. (2015) and Patton Gerais de São Felipe, BA (Dunnum, 2015) to et al. (2015). For biome endemism, we used Paglia et al. (2012). Classification of endangered species Araçuaí municipality (locality 34). The record of (Vulnerable, Endangered and Critically Endangered) Kerodon rupestris in the Parque Estadual do Rio is according to IUCN (2015) and MMA (2014). Preto, São Gonçalo do Rio Preto municipality Near Threatened (NT) and Data Deficient (DD) (locality 7) increases the species distribution are according to ICMBio (2014) and IUCN (2015). to the southeast in 330 km from Riacho da Cruz, MG (Dunnum, 2015). The record of RESULTS angouya in Distrito Carne Seca, Fazenda Corredor, MG (locality 12) increases Our collecting effort in 13 localities and with the species distribution to the northwest in the inclusion of museum specimens and lit- 450 km from Rio São José, ES (Percequillo, erature information resulted in a list of 91 2015b). Thylamys velutinus was recorded in Rio mammal species in 75 localities along the do Pardo de Minas municipality, MG (locality Jequitinhonha River Basin (Appendices I and 48), which increases the species distribution II); 15 species belong to the Order Carnivora, to the east in 150 km from Lagoa Santa, MG four to Cetartiodactyla, six to Cingulata, 14 to (Creighton and Gardner, 2008). The record of Didelphimorphia, one to Lagomorpha, one to Monodelphis scalops at the Parque Nacional Perissodactyla, three to Pilosa, 11 to Primates, das Sempre Vivas, Diamantina municipality, and 36 species to Rodentia. We trapped 33 MG (locality 5) increases the species distribu- species of small mammals (13 marsupials tion to the northwest in 400 km from Santa and 20 rodents), three species were visualized Teresa, ES (Pine and Handley, 2008). Wiedomys (Cabassous unicinctus, Callithrix geoffroyi and pyrrhorhinos had a distribution increase to the Guerlinguetus brasiliensis), one was recorded south in 145 km from Juramento, BA (Bonvi- from footprints (Procyon cancrivorus) and cino, 2015) to Pousada Água Quente, Felício two species of medium sized-mammal were dos Santos municipality, MG (locality 9). recovered as road-kills (Cerdocyon thous and We obtained karyotype for 26 species (11 Sylvilagus minensis) (Table 2). Didelphimorphia and 15 Rodentia; Table 3). According to the IUCN (2015) 20 out of the Eight species presented karyotype 91 species recorded for the Jequitinhonha River variation—Akodon cursor, scotti, Basin are threatened, while according to the , , MMA (2014) and ICMBio (2014) 27 species squamipes, Rhipidomys mastacalis, are threatened. For IUCN (2015), the most Trinomys albispinus and Wiedomys pyrrhorhinos endangered species are Brachyteles hypoxanthus (Table 3). Undescribed chromosomal variation and Sapajus xanthosternos (with the status of was found in C. scotti, C. subflavus, T. albispinus Critically Endangered), and Leontopithecus and W. pyrrhorhinos. speci- chrysomelas and Sapajus robustus (with the mens, collected in Fazenda Santa Cruz (locality status of Endangered). According to the MMA 6) presented two distinct fundamental numbers, (2014) and ICMBio (2014) the worst cases are 68 (MN82753, Fig. 3) and 72 (MN82752). The B. hypoxanthus as Critically Endangered, and karyotype with FN=68 is composed by six pairs L. chrysomelas, S. robustus, S. xanthosternos, of biarmed and 22 acrocentric chromosomes. Leopardus guttulus, Thalpomys lasiotis and The karyotype of C. subflavus(MN82759, col- Trinomys mirapitanga with the status of En- lected also in locality 6) presented variation in dangered (Table 2). the sexual pair, in which the X chromosome We increased the known distribution for is a large submetacentric and the Y is a small eight species. The record ofCalomys mattevii acrocentric (Fig. 4). Trinomys albispinus was TERRESTRIAL MAMMALS OF THE JEQUITINHONHA RIVER BASIN, BRAZIL 103 Savanna/ = MMA ICMBio MMA Least Concern, = SeasonalDecidual = ConservationStatus IUCN Secondary Vegetation and Agrarian Agrarian and Secondary Vegetation b = c, e c, e,k a, b a, Type a, c, e c, a, e c, a, a, c, e, k e, c, a, a, c, d, e d, c, a, Vegetation Vegetation a, b, c, d, e d, c, b, a, e d, c, b, a, a, c, d, e, k e, d, c, a, a, b, c, d, e, f e, d, c, b, a, Arboreous Steppic Savanna; h Savanna; Steppic Arboreous Upper Montane Vegetational Refuges; c Vegetational Montane Upper CE CE MA = = Biome CE/AF CE/AF CE/AF CE/AF CE/AF CE/AF CE/AF CE/AF CE/AF voucher specimens. Biome according to IBGE to (2012): according Biome specimens. voucher = ), type of record, biome, vegetation and conservation status. Type Type status. conservation and vegetation biome, ), typerecord, of VS VS VS VS VS R(1), VS R(1), VS R(5), VS R(1), VS R(1), VS R(1), VS R(1), VS R(4), VS visualization; Arboreous Savanna; g Savanna; Arboreous = Referencenumber Type of Record and and Record of Type = Appendix I Appendix Grassy-Woody Savanna; b Savanna; Grassy-Woody = Table2 Vulnerable, according to International Union for Conservation of Nature (IUCN), Ministério (IUCN), Ministério Nature Conservation of for Union International to according Vulnerable, LC Conservation Status: for Legend Lowlands. of Forest Broadleaf Moist = = Road killed V animals; 5 7, 5 7, 9, 49 9, = Localities 7, 8, 57, 64 57, 8, 7, 2, 6, 7, 9, 60 9, 7, 6, 2, 49, 61, 68, 75 68, 61, 49, 7, 17, 54, 64, 75 64, 54, 17, 7, Mangroves; k Mangroves; 5, 6, 7, 9, 30, 35, 45, 46 45, 35, 30, 9, 7, 6, 5, 7, 8, 9, 17, 30, 46, 64, 75 64, 46, 30, 17, 9, 8, 7, = 30, 31, 33, 45, 49, 53, 60, 65 60, 53, 49, 45, 33, 31, 30, 2, 3, 7, 9, 12, 31, 37, 43, 45, 47 45, 43, 37, 31, 12, 9, 7, 3, 2, 2, 5, 7, 9, 33, 38, 43, 45, 46, 49, 53, 56 53, 49, 46, 45, 43, 38, 33, 9, 7, 5, 2, 2, 3, 5, 6, 7, 8, 9, 11, 12, 14, 17, 27, 28, 27, 17, 14, 12, 11, 9, 8, 7, 6, 5, 3, 2, Critically Endangered, V Critically Endangered, = ) References: 1. Chiarello et al. (2006), 2. Falcão et al. (2012), 3. Leal et al. (2008), 4. Lessa et al. (2008), 5. Lessa and Paula (2014), (2012), 3. Leal et al. (2006), 2. Falcão et al. (2008), 4. Lessa et al. Paula 1. Chiarello (2008), 5. Lessa et al. and References: (É. Geoffroy) (É. Endangered, CE Endangered, = Müller = Savanna/ Broadleaf Forest; j Forest; Broadleaf Savanna/ Atlantic Forest. Vegetation according to IBGE to (2012): a according Vegetation Forest. Atlantic Linnaeus = Lund = (Lund) (Burmeister Agrarian Activities (originally Savanna, Steppic Savanna/Seasonal Forest or Savanna/ Broadleaf Forest); e Forest); Broadleaf Savanna/ or Forest Savanna/Seasonal Steppic Savanna, (originally Activities Agrarian Linnaeus (Tate) = (Wagner) (Wagner) footprint; R footprint; Thomas (Wied-Neuwied) = Near Threatened, E Threatened, Near Cerrado AF and Caluromys philander philander Caluromys Didelphisalbiventris aurita D. Gracilinanusagilis microtarsus G. Marmosamurina M.paraguayana Marmosopsincanus nudicaudatus Metachirus americana Monodelphis domestica M. scalops M. = = Melo (2005), 7. Melo et al. (2004), 8. Oliveira and Gonçalves (2015), 9. Oliveira et al. (2013), 10. Pardiñas et al. (2014), 11. Pessôa et al. (2015), 12. Pinto and and (2015), 12. Pinto et al. (2014), 11. Pessôa et al. (2013), 10. Pardiñas et al. (2015), 9. Oliveira Gonçalves and (2004), 8. Oliveira et al. (2005), 7. Melo Melo DIDELPHIMORPHIA Gill DIDELPHIMORPHIA Registered mammal species, with their locality numbers (for localities name see localities name (for species, their locality with mammal numbers Registered F record: of 6. (2015); RK (1988), 14. Voss et al. (1997), 13. Rylands Rylands CE d SeasonalForest; f Forests); Broadleaf Moist Seasonal (originally or Decidual, Semideciduous Activities i Forest; Submontante NT Conservaçãode (ICMBio). Chico Mendes da Biodiversidade Instituto (MMA), Ambiente Meio do 104 Mastozoología Neotropical, 24(1):95-119, Mendoza, 2017 L Geise et al. http://www.sarem.org.ar - http://www.sbmz.com.br Vu Vu Vu Vu CE NT NT DD (Table 2 cont.) 2 (Table MMA ICMBio MMA

ConservationStatus Vu CE NT IUCN c c e e e b e,k a, b a, b a, b a, d,e Type a, b, e b, a, a, b, k b, a, a, b, e, k e, b, a, a, b, c, k c, b, a, Vegetation Vegetation a, b, c, d, e d, c, b, a, a, c, d, e, k e, d, c, a, b, c, d, e, f, j f, e, d, c, b, CE CE CE CE CE MA MA MA MA MA MA Biome CE/AF CE/AF CE/AF CE/AF CE/AF CE/AF CE/AF VS VS R(3) R(7) R(1) R(3,9) R(3, 4) R(3, 2) R(1, 9) R(3, R(1, 13) R(1, R(1, 3, 4) 3, R(1, R(2, 3), VS 3), R(2, R(1, 2, 3, 4) 3, 2, R(1, R(3, 13), VS 13), R(3, R(2, 3, 4), V 4), 3, R(2, R(1, 3, 4), VS 4), 3, R(1, R(1, 6, 13), VS 13), 6, R(1, R(1, 2, 4, 13), V 13), 4, 2, R(1, Referencenumber Type of Record and and Record of Type 9 5 48 65 5, 7 5, 7 5, 7 5, 55, 64 55, 5, 7, 65 7, 5, Localities 3, 5, 7, 75 7, 5, 3, 5, 7, 45, 75 45, 7, 5, 75 65, 7, 5, 64, 65, 68, 75 68, 65, 64, 65 63, 61, 57, 5, 7, 15, 16, 46, 65 46, 16, 15, 7, 5, 33, 41, 42, 52, 55, 57, 65 57, 55, 52, 42, 41, 33, 7, 9, 29, 39, 40, 52, 58, 64, 75 64, 58, 52, 40, 39, 29, 9, 7, 5, 21, 22, 25, 26, 32, 33, 41, 46, 59, 74 59, 46, 41, 33, 32, 26, 25, 22, 21, 5, us e

Linnaeus (Kuhl) (Linnaeus) Linnaeus (Kerr) Schinz (Wagner) (Humboldt) (Desmarest) (Olfers) Linnaeus (Coimbra-Filho) Humboldt (Wied-Neuwied) (É. Geoffroy) (É. (Linnaeus) Humboldt Philander frenatus frenatus Philander velutinus Thylamys Cabassous tatouay unicinctus C. Dasypusnovemcinctus septemcinctus D. Linna sexcinctus Euphractus maximus Priodontes variegatus Bradypus tridactyla Myrmecophaga tetradactyla Tamandua caraya Alouatta guariba A. Brachyteleshypoxanthus Callicebuskuhlii melanochir C. geoffroyi Callithrix penicillata C. CINGULATA Illiger CINGULATA Flower PILOSA Linnaeus PRIMATES TERRESTRIAL MAMMALS OF THE JEQUITINHONHA RIVER BASIN, BRAZIL 105 En En En En Vu Vu Vu Vu Vu Vu Vu NT (Table 2 cont.) 2 (Table MMA ICMBio MMA

ConservationStatus En En Vu Vu CE NT NT NT NT IUCN e e e e a b b a, b a, b a, b a, b a, b,k Type a, b, e b, a, b, g, k b, a, b, c, e c, b, a, a, b, e, k e, b, a, a, b, e, h e, b, a, Vegetation Vegetation a, b, c, e, g e, c, b, a, a, b, c, d, e d, c, b, a, b, c, d, e, k e, d, c, b, CE CE CE CE CE CE CE MA MA MA MA Biome CE/AF CE/AF CE/AF CE/AF CE/AF CE/AF CE/AF CE/AF CE/AF R(2) R(4) R(3) R(1) R(3) R(13) R(12) R(3-4) R(1, 6) R(1, 4) R(3, 4) R(3, 4) R(3, 3) R(2, R(1, 3, 4) 3, R(1, F, R(1, 3, 9) 3, R(1, F, R(1, 2, 3, 4) 3, 2, R(1, R(1, 3, 4), VS 4), 3, R(1, R(1, 3, 4), RK 4), 3, R(1, RK 4), 3, R(1, R(1, 2, 3, 13), VS 13), 3, 2, R(1, Referencenumber Type of Record and and Record of Type 7 5 5 20 64 5, 7 5, 7 5, 7 5, 7 5, 5, 75 5, 5, 7, 64 7, 5, 5, 50, 75 50, 5, 69, 70, 71 70, 69, Localities 55, 57, 65, 68 65, 57, 55, 5, 7, 43, 50, 65 50, 43, 7, 5, 75 61, 57, 7, 5, 5, 7, 9, 48, 51, 57 51, 48, 9, 7, 5, 66 64, 57, 9, 7, 5, 5, 7, 19, 36, 46, 57 46, 36, 19, 7, 5, 5, 25, 33, 64, 74, 75 74, 64, 33, 25, 5, Kuhl Illiger (Boddaert) (Wied-Neuwied) G. [Baron] Cuvier [Baron] G. (Lund) (Thomas) Olfers (Linnaeus) É. Geoffroy Saint-Hilaire Geoffroy É. Linnaeus Linnaeus Geoffroy Linnaeus Linnaeus Schreber (Kuhl) Hensel (Schinz) C. personatus personatus C. chrysomelas Leontopithecus xanthosternos Sapajus robustus S. minensis Sylvilagus Cerdocyon thous brachyurus Chrysocyon Conepatussemistriatus EiraBarbara Leoparduspardalis Linnaeus guttulus L. wiedi L. longicaudis Lontra nasua Nasua onca Panthera Potosflavus Pumaconcolor yagouaroundi P. cancrivorus Procyon vetulus Pseudalopex LAGOMORPHA Brandt LAGOMORPHA Bowdich CARNIVORA 106 Mastozoología Neotropical, 24(1):95-119, Mendoza, 2017 L Geise et al. http://www.sarem.org.ar - http://www.sbmz.com.br Vu Vu Vu DD (Table 2 cont.) 2 (Table MMA ICMBio MMA ConservationStatus Vu Vu NT DD DD IUCN e e b b b c, e c, e c, a, b a, b a, b a, c, d c, Type a, b, k b, a, a, b, e, k e, b, a, k e, b, a, k e, b, a, a, b, d, e, f e, d, b, a, Vegetation Vegetation a, b, c, d, e, f, g f, e, d, c, b, a, CE CE CE CE CE CE MA MA MA MA Biome CE/AF CE/AF CE/AF CE/AF CE/AF CE/AF CE/AF VS VS VS VS R(3) R(3) R(10) R(14) R(3, 4) R(3, R(1, 14) R(1, R(1), VS R(1), VS R(4), VS R(1), R(2, 3, 4) 3, R(2, 4) 2, R(1, R(1, 2, 3, 4) 3, 2, R(1, 4) 3, 2, R(1, Referencenumber Type of Record and and Record of Type 5 5 5 63 5, 7 5, 7 5, 37, 44 37, 54 45, 62 61, 5, 6, 7 6, 5, 5, 7, 75 7, 5, 17, 49, 53 49, 17, Localities 5, 7, 57, 75 57, 7, 5, 48, 50, 53, 61 53, 50, 48, 5, 7, 57, 64, 75 64, 57, 7, 5, 5, 7, 57, 64, 65, 75 65, 64, 57, 7, 5, 3, 5, 12, 27, 34, 46, 47, 48, 65 48, 47, 46, 34, 27, 12, 5, 3, 3, 5, 6, 7, 9, 11, 12, 22, 27, 35, 45, 46, 45, 35, 27, 22, 12, 11, 9, 7, 6, 5, 3,

Olfers Linnaeus Erxleben Montgelard, Catzeflis and Douzery and Catzeflis Montgelard, Pardiñas, Lessa, Teta, Pardiñas, Olfers Lichtenstein (Langguth (Langguth (Lund) Linnaeus G. Fischer [von Waldeheim] [von Fischer G. (Linnaeus) (Winge)

Erxleben Linnaeus (Wagner) (Winge) Percequillo, Hinst-Zaher Hinst-Zaher Percequillo, Tapirus terrestris Tapirus americana Mazama gouazoubira M. bezoarticus Ozotocerus tajacu Pecari cursor Akodon apicalis Calassomys Câmara and Salazar-Bravo mattevii Calomys tener C. aperea Cavia scotti Cerradomys Bonvicino) and subflavus C. vivoi C. Bonvicino and Chaetomyssubspinosus Coendouinsidiosus paca Cuniculus azarae Dasyprocta PERISSODACTYLA Owen PERISSODACTYLA CETARTIODACTYLA Bowdich RODENTIA TERRESTRIAL MAMMALS OF THE JEQUITINHONHA RIVER BASIN, BRAZIL 107 En DD (Table 2 cont.) 2 (Table MMA ICMBio MMA ConservationStatus NT DD IUCN b b d e,i e,i e,i b,e c, k c, a, b a, c, d c, d c, a, d a, Type a, b, c b, a, c b, a, b, c, d, f d, c, b, Vegetation Vegetation a, b, c, e, k e, c, b, a, a, b, c, d, e d, c, b, a, e d, c, b, a, a, c, d, e, k e, d, c, a, a, b, c, d, e, f e, d, c, b, a, CE CE CE CE CE CE CE MA MA MA Biome CE/AF CE/AF CE/AF CE/AF CE/AF CE/AF CE/AF CE/AF CE/AF CE/AF VS VS VS VS VS VS VS VS VS VS VS R(3) R(2), VS R(2), VS R(1), VS R(1), VS R(8), VS R(3), R(3, 4), VS 4), R(3, VS 4), R(3, R(1, 3, 4), V, VS V, 4), 3, R(1, Referencenumber Type of Record and and Record of Type 5 5 12 5, 6 5, 18, 75 18, 48 34, 72 64, 72 54, 33 17, 47 45, 5, 7, 13 7, 5, 13 7, 5, 54 5, 1, 6, 12, 31 12, 6, 53, 54, 72 54, 53, Localities 5, 27, 30, 46 30, 27, 5, 5, 7, 9, 12, 31, 33, 47 33, 31, 12, 9, 7, 5, 5, 7, 8, 9, 24, 53, 57, 64, 75 64, 57, 53, 24, 9, 8, 7, 5, 5, 7, 9, 49, 57, 61, 64, 65, 68, 75 68, 65, 64, 61, 57, 49, 9, 7, 5, 3, 4, 5, 6, 7, 9, 17, 33, 45, 46, 49, 73 49, 46, 45, 33, 17, 9, 7, 6, 5, 4, 3, 2, 5, 6, 7, 8, 9, 12, 15, 24, 27, 28, 31, 44, 31, 28, 27, 24, 15, 12, 9, 8, 7, 6, 5, 2, (G. Fischer) (G. (Linnaeus) (Gmelin) (Schinz) (Winge) (Lund) (Gervais) (Olfers) (Brants) (Weksler, Geise (Weksler, (Thomas) Thomas (Fischer) (Thomas) (Lund) (Wied-Neuwied) (Lund) (Wagler) (Linnaeus) D. leporina leporina D. (Wagner) russatus spinosus Euryzygomatomys spixii Galea brasiliensis Guerlinguetus hydrochaeris Hydrochoerus seuanezi Kerodonrupestris lasiurus Necromys Nectomyssquamipes Oecomyscatherinae nigripes dasytrichus lamarum Phyllomys simplex macrurus Rhipidomys R.mastacalis angoya Sooretamys lasiotis Thalpomys apereoides Thrichomys and Cerqueira) and 108 Mastozoología Neotropical, 24(1):95-119, Mendoza, 2017 L Geise et al. http://www.sarem.org.ar - http://www.sbmz.com.br

collected only in locality 45 (MN82935, from Fazenda Galiléia), the fundamental number (FN) 108, with 25 pairs of biarmed and four En pairs of acrocentric chromosomes (Fig. 5). (Table 2 cont.) 2 (Table

MMA ICMBio MMA Wiedomys pyrrhorhinos presented two different fundamental numbers, FN=98 (MN82944, from Fazenda Ilha, locality 43; MN82938 and 82939 ConservationStatus DD

IUCN from Fazenda Galiléia, locality 45; MN82940, 82941, 82942 from Pousada Água Quente, locality 9), with 19 pairs of biarmed and 11 pairs of acrocentric chromosomes (Fig. 6A) e

Type and FN=99 (MN82943, from Fazenda Ilha, a, c, d, f d, c, a, a, c, d, e d, c, a, b, c, e, f, h f, e, c, b, Vegetation Vegetation locality 43) composed by 19 pairs of biarmed and 10 pairs of acrocentric chromosomes, and a heteromorphic pair of small chromosomes

MA (Fig. 6B). Biome CE/AF CE/AF CE/AF DISCUSSION

Our trapping efforts, coupled with all the

VS information obtained from museums and the R(1)

R(11), VS R(11), literature, provided a comprehensive view of R(1, 11), VS 11), R(1, the non-volant mammal diversity along the Referencenumber Type of Record and and Record of Type Jequitinhonha River Basin. Terrestrial mam- mal species richness recorded in Jequitinhonha River Basin represents 17% of all known ter- restrial species in Brazil, not considering Calassomys apicalis, described after Paglia et al. (2012). Most recorded species (91) have a wide geographical distribution range and

61, 68 61, may occur in more than one biome in South Localities America (Bonvicino et al., 2002; Geise and Astúa, 2009; Pereira and Geise, 2009; Car-

7, 9, 11, 12, 28, 34, 43, 45 43, 34, 28, 12, 11, 9, 7, mignotto et al., 2012; Costa and Leite, 2012;

9, 12, 17, 23, 53, 54, 57, 58, 65, 68 65, 58, 57, 54, 53, 23, 17, 12, 9, Paglia et al., 2012); however, we collected 19 5, 10, 12, 22, 23, 27, 28, 45, 54, 65, 66 65, 54, 45, 28, 27, 23, 22, 12, 10, 5, species only in Atlantic Forest localities and 26 in Cerrado localities. Currently, 32 of 251 (not including C. apicalis) and 90 of 298 species of mammals occur/are endemic and registered in the Cerrado and Atlantic Forest, respectively (Carmignotto et al., 2012; Costa and Leite, 2012; Paglia et (Wied-Neuwied) (Wied-Neuwied) al., 2012). We trapped a considerable por- tion of this diversity, and recorded 28.7% Lara, Patton Lara,Patton (6.3% endemic) of all Cerrado and 21.5%

(Desmarest) (22.2% endemic) of all Atlantic Forest spe- cies, thus showing the biological importance of the surveyed area. We recorded species

Trinomys albispinus (I. Geoffroy St.-Hilaire) Geoffroy (I. albispinus Trinomys mirapitanga T. Hingst-Zaher and setosus T. pyrrhorhinos Wiedomys considered to be restricted to the Cerrado and Caatinga in the Atlantic Forest (Calomys TERRESTRIAL MAMMALS OF THE JEQUITINHONHA RIVER BASIN, BRAZIL 109

Table 3 Species karyotyped with locality number (for name of locality see Appendix I). Diploid number (2n) and fundamental number (FN). * Indicates those species with new karyotype variation.

Species Locality number 2n FN

Didelphimorphia Caluromys philander 8 14 24 Didelphis albiventris 9 22 20 Gracilinanus agilis 9, 43, 45 14 20 G. microtarsus 2, 6, 9 14 20 Marmosa murina 75 14 20 M. paraguayana 8, 9 14 20 Marmosops incanus 2, 6, 8, 9, 45, 49, 53 14 24 Metachirus nudicaudatus 75 14 20 Monodelphis americana 9, 49 18 22 M. domestica 6, 9, 45 18 22 Philander frenatus 9 22 20 Rodentia Akodon cursor 17, 49, 53 14/16 18/19/20/21/26 Calomys mattevii 44 66 68 Cerradomys scotti* 6 58 68/72 C. subflavus* 6, 53 54 62 C. vivoi 45 50 64/66 53 80 84 Guerlinguetus brasiliensis 49 40 76 8, 9, 49, 53, 75 56/57 56/57/58/60 9 62 82 Rhipidomys macrurus 6 44 48 R. mastacalis 6, 9, 49, 73 44 72/74 Thrichomys apereoides 2, 6, 8, 9, 44, 45 28 50 Trinomys albispinus* 45 60 108 T. setosus 9, 53 56 108 Wiedomys pyrrhorhinos* 9, 43, 45 62 98/99

mattevii, Didelphis albiventris, Gracilinanus Sapajus robustus, Sooretamys angoya and agilis, Rhipidomys macrurus, Thrichomys Trinomys setosus) (Table 2). Such records show apereoides and Wiedomys pyrrhorhinos), as well the need of carefully reassess the scheme of bi- as species considered restricted to the Atlantic ome endemicity provided by Paglia et al. (2012). Forest in Cerrado areas (Callithrix geoffroyi, Habitat and vegetation characteristics were Dasyprocta leporina, Gracilinanus microtarsus, observed during field work, differing from Guerlinguelus brasiliensis, Monodolphis scalops, biome and vegetation maps (IBGE 2012), Oxymycterus dasytrichus, Phyllomys lamarum, showing that the Cerrado and Atlantic Forest 110 Mastozoología Neotropical, 24(1):95-119, Mendoza, 2017 L Geise et al. http://www.sarem.org.ar - http://www.sbmz.com.br

Fig. 3. Karyotype in conventional staining of a female Cerradomys scotti (MN82753), collected at Fazenda Santa Cruz (locality 6), showing 2n=58, FN=68, with six pairs of biarmed and 22 pairs of acrocentric chromosomes. X chromosome as a medium biarmed chromosome.

Fig. 4. Karyotype in conventional staining of male Cerradomys subflavus (MN82759), collected at Fazenda Santa Cruz (locality 6), showing 2n=54, FN=62, with five pairs of biarmed and 21 pairs of acrocentric chromosomes. X chromosome as a medium biarmed, and Y chromosome as a small acrocentric. TERRESTRIAL MAMMALS OF THE JEQUITINHONHA RIVER BASIN, BRAZIL 111

Fig. 5. Karyotype in conventional staining of Trinomys albispinus (MN82935) collected at Fazenda Galiléia (locality 45) showing 2n=60, FN=108, with 25 pairs of biarmed and four pairs of acrocentric chromosomes; male individual, X chro- mosome as a large biarmed, and Y a small acrocentric chromosome. borders are not precisely delimited, either due [REBIO] Mata Escura), and one in Bahia State to the low resolution of IBGE data or acceler- (Atlantic Forest Biome: Reserva Particular de ated rates of deforestation. This could be seen in Proteção Natural [RPPN] Estação Veracel). In localities 43, 44 and 45, which are not Atlantic those protected areas, according to the infor- Forest, but dry vegetation of Cerrado (for a mation provided by environmental government better conclusion, see Figs. 2F, 2G and 2H) agencies (ICMBio, 2016 and IEF/MG, 2016 or Fazenda Palmares (locality 49), according accessed August/2016) 14 endangered taxa may to Fig. 1 in the Atlantic Forest (Secondary occur (Alouatta spp., Brachyteles hypoxanthus, Vegetation and Agrarian Activities, originally Callicebus spp., Callithrix geoffroyi, Chaetomys Seasonal Semidecidous Forest; IBGE, 2012), subspinosus, Chrysocyon brachyurus, Leopardus but according to fieldwork, a transition area pardalis, Lontra longicaudis, Myrmecophaga between both biomes. tridactyla, Panthera onca, Priodontes maximus, The importance of the species survey in- Puma concolor, Sapajus robustus and Tapirus creases when the conservation status is consid- terrestris), all medium or large mammals. ered. Eight major protected areas are located However, we have in our list three species along the basin, three of them surveyed by of small mammals under conservation threat us. Seven protected areas are in Minas Gerais (Thalpomys lasiotis, Thylamys velutinus and state (six in Cerrado biome: Parque Nacional Trinomys mirapitanga), with only few oc- Sempre Vivas, Parque Estadual do Rio Preto, currences in protected areas. The sum of all Parque Estadual de Biribiri, Parque Estadual those eight protected area is 2570 km2, which do Pico do Itambé, Parque Estadual da Serra represents only a small fraction (3.7%) of the Negra, Parque Estadual de Grão Mogol; and whole basin (70 315 km2). Most of the pro- one in Atlantic Forest biome: Reserva Biológica tected areas are in the Cerrado, even when 112 Mastozoología Neotropical, 24(1):95-119, Mendoza, 2017 L Geise et al. http://www.sarem.org.ar - http://www.sbmz.com.br

A

B

Fig. 6. Karyotypes in conventional staining of Wiedomys pyrrhorhinos collected at Fazenda Ilha (locality 43). A) MN82944, 2n=62, FN=98, with 19 pairs of biarmed and 11 pairs of acrocentric chromosomes; female individual, X chromosome as a medium acrocentric. B) MN82943, with 2n=62, FN=99, with 19 pairs of biarmed and 10 pairs of acrocentric chro- mosomes, and a heteromorphic pair of small chromosomes; female individual, X chromosome as a medium autosomal. a larger area of the basin lies in the Atlantic We obtained new karyotypes for Cerradomys Forest (Fig. 1). scotti, C. subflavus, Trinomys albispinusand All collected marsupial species, a group Wiedomys pyrrhorhinos. The fundamental known to have a very conservative chromo- number described in the present paper for somal composition (Svartman and Vianna- C. scotti is lower, with an increase of acrocentric Morgante, 1998; Astúa, 2015), showed diploid chromosomes, than those previously reported and autosomal numbers already described in (Bonvicino et al., 1999). The sexual pair here the literature (Yunis et al., 1973; Carvalho reported for one specimen of C. subflavus dif- et al., 2002; Paresque et al., 2004; Pereira et fers from the previously described karyotype al., 2008). (Bonvicino et al., 1999). The fundamental TERRESTRIAL MAMMALS OF THE JEQUITINHONHA RIVER BASIN, BRAZIL 113 number of T. albispinus presented here (FN trapping effort is necessary to further character- = 108) differs because of the presence of 25 ize (e.g., phylogeography) the registered species. biarmed pairs instead of 29 as described by Pessôa et al. (2015). The published karyotypes ACKNOWLEDGMENTS for W. pyrrhorhinos show that the species is We are thankful to Júlia Lins Luz, Maíra de Godoy Sant’Ana, highly variable (2n=62, FN=86, 90 and 104; Luciana de Moraes Costa, Thais Lira and Vera de Ferran for Maia and Langguth, 1987; Gonçalves et al., fieldwork help; to Heitor Bispo, Fábio Henrique Alves Bispo, 2005; Pereira and Geise, 2007). The karyo- Luiz Carlos da Silva, Rita de Cassia de Melo Tolentino and type presented here (2n=62, FN=98/99), with Georg Marksteinr (Pousada Água Quente and Fazenda Sumidouro), Sueli Vieira (Fazenda Ilha), Mr. Túlio (Fazenda another different autosomal number, increases Palmares), Mr. and Mrs. Guimarães (Fazenda Anga-Pehy) the known intraspecific variation. and all the staff of the Reserva Particular do Patrimônio Chromosome data available for Brazilian Natural, Estação Veracel, for lodging and permission to populations of are scarce (Fagundes et work within their properties. Leila Maria Pessôa helped al., 2003), with most species presenting a simi- in the identification ofTrinomys spp. Fabiano Rodrigues de Melo was of great help providing information about all lar karyotype, 2n=40, FN=74 ( aestuans Primates species; R. Moratelli kindly assessed Oxymycterus ingrami) and 2n=40, FN=76 (S. a. alphonsei dasytrichus specimens in the Smithsonian Institution, and S. a. spadiceus, described by Lima and Washington DC. Field work was carried out with funding Langguth 2002); both karyomorphs differ in provided by Conselho Nacional de Desenvolvimento Cientí- the 19th pair, with metacentric chromosomes in fico e Tecnológico (CNPq, Edital Universal # 473596/2006-7 and 472909/2009-6 andComCerrado#563134/2010-0 Edital the former and acrocentric in the second one. nº 47/2010 - Chamada 2), Fundação de Amparo à Pesquisa The specimens we collected match the descrip- do Estado do Rio de Janeiro (FAPERJ), Fundação de tion of Vivo and Carmignotto (2015) and Vivo Amparo a Pesquisa do Estado de Minas Gerais (FAPE- (personal communication) for Guerlinguelus MIG - # APQ01034/09) and Programa SISBIOTA/CNPq brasiliensis brasiliensis, which currently include (# 56314/2010.0). L. Geise receives productive grants from CNPq (# 306161/2016-8) and UERJ/Prociência, C.E.L. Es- S. a. alphonsei and S. a. spadiceus as synonyms. bérard received a CNPq productive grant (# 151029/2004-0) So, we conclude that the karyotype observed by and financial support from FAPERJ to “Young Scientist of us for our sample of G. brasiliensis is the same of Our State” (E-16/170.449/2007 and E-26/102.201/2009). D. those described by Lima and Langguth (2002). Astúa was supported by FAPERJ (E-26/150.353/2005) and The area surveyed and studied here shows by a CNPq fellowship (306647/2013- 3). Marcia Aguieiras had fellowship from FAPERJ (E-26/102.327/2011) and few particular patterns of species distribution has a doctoral scholarship from CAPES. Paulo H. Asfora occurrence. Some are already known in litera- had a fellowship from FAPERJ (E-26/100.331/2007) and ture, such as the rodent species Akodon cursor Fundação de Amparo à Ciência e Tecnologia do Estado (Geise, 2012) and Cerradomys spp., where de Pernambuco (FACEPE - DCR-0053-2.04/11, Bolsa C. subflavus occurs only on the right side of DCR e APQ-0020-2.04/12). F. Dourado receives financial support from FAPERJ to “Young Scientist of Our State” the river as opposed to C. vivoi, which occurs (E-16/203.218/2015). We are grateful to all comments made north of the left side (Bonvicino et al., 1999; by two anonymous reviewers. Weksler et al., 2006; Percequillo, 2015a). Leite et al. (2016) presented a complete explana- LITERATURE CITED tion about the Rio Doce as a discontinuity area. Further extensive collecting effort in the ASTÚA D. 2015. Family Didelphidae (Opossums). Pp. 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APPENDIX I Gazetteer with geographical and vegetation information for localities from Jequitinhonha River Basin with mammal registers, arranged from the source to the mouth region of the river. Under- lined names = municipalities. Number before locality name corresponds to localities plotted in Fig. 1. Localities with our collecting effort are in bold. nln = no locality name. When coordinates of a locality were not found we used coordinates of the municipality. After coordinates, vegetation type (IBGE 2012): Agrarian Activities (originally Savanna, Steppic Savanna/Seasonal Forest or Savanna/ Broadleaf Forest) = AA; Arboreous Savanna = AS; Arboreous Steppic Savanna = ASS; Grassy-Woody Savanna = GWS; Mangroves = MA; Moist Broadleaf Forest of Lowlands = MBFL; Savanna/ Broadleaf Forest = S/BF; Savanna/Seasonal Forest = S/SF; Seasonal Decidual Submontante Forest = SDSF; Secondary Vegetation and Agrarian Activities (originally Seasonal Decidual, Semi- deciduous or Moist Broadleaf Forests) = SV/AA; Upper Montane Vegetational Refuges = UMVR. MINAS GERAIS STATE: Serro, 1 – nln (18°36’10”S, 43°22’45”W, SV/AA, originally SSF); Diamantina, 2- Pousada Rural Recanto do Vale (18°23’43.2”S, 43°32’25.5”W, GWS), 3 - Lavrinha, Mineração Tejucana (18°14’58”S, 43°36’01”W, GWS), 4 - Conselheiro Mata (18°14’44”S, 43°54’02”W, GWS), 5 – Parque Nacional das Sempre Vivas (17°55’02”S; 43°47’11”W, UMVR), 6 - Fazenda Santa Cruz (18°16’16.2”S, 43°23’18.8”W, GWS); São Gonçalo do Rio Preto, 7 - Parque Estadual do Rio Preto (18°05’20”S, 43°20’25”W, GWS); Felício dos Santos, 8 - Fazenda Sumidouro (18°11’34.4”S, 43°14’35.2”W, S/ SF), 9 - Pousada Água Quente (18°05’32.5”S, 43°10’26.2”W, S/SF); Bocaiúva, 10 - nln (17°06’59”S, 43°49”58”W, S/SF), 11 - Mineração SADA (17°06’54”S, 43°49’16”W, S/SF), 12 - Distrito Carne Seca, Fazenda Corredor (17°02’14”S, 43°52’16”W, AA, originally S); , 13 - Veredas de Botumirim, 32 km NW Caçaratiba (17°07’S, 43°02’W, S/SF); Turmalina, 14 - nln (17°17’30”S, 42°43’05”W, S/SF), 15 - Road close to Córrego Divisão, 1 km SW Peixe Cru and Alambique do Joel, 800 m Peixe Cru (17°16’56”S, 42°44’07”W, S/SF), 16 - Usina Hidrelétrica de Santa Rita (17°17’08”S, 42°43’48”W, S/ SF), 17- , Estação Ecológica de Acauã, 17 km N from Turmalina (17°08’S, 42°46’W, S/SF), 18 - Rodovia MG 367, km 405 (17°13’07”S, 42°35’25”W, S/SF); , 19 - nln (17°12’05”S, 42°36’07”W, S/SF), 20 - Córrego da Chácara (17°45’S, 42°30’W, SV/AA, originally SSF); Cristália, 21 - nln (16°49’S, 42°41’W, S/SF), 22 - Córrego Contendas (16°45’S, 42°52’W, AS), 23 - Left margin of Jequitinhonha River (16°43’S, 42°37’W, S/SF); , 24 - Buriti, margin of Jequitinhonha River (16°49’S, 42°41’W, S/SF), 25 - Buriti, Fazenda Irmãos Atachi (16°51’S, 42°38’W, S/SF); Grão Mogol, 26 - Fazenda Giro (16°34’S, 43°02’W, AS), 27 - Usina Hidrelétrica Kubitschek (UHE Irapé) (16°33’26”S, 42°53’38”W, AS), 28 - Fazenda Curral Velho, 28 km E from Grão Mogol, Fazenda Maria das Neves and Fazenda do Matão (16°33’34”S, 42°53’23”W, AS); , 29 - Rio Capivari (17°06’S, 42°32’W, S/SF); Francisco Badaró, 30 (16°58’52”S, 42°20’45”W, S/SF); , 31- Ponte do Colatino, left margin of Jequitinhonha River (16°36’S, 42°12’W, AA, originally SS/SF); , 32 - Fazenda Mandacarú (16°42’S, 42°13’W, S/SF), 33 - Fazenda Paiol (16°50’S, 42°13’W, AA, originally SS/SF); Araçuaí, 34 - nln (16°51’20”S, 42°03’25”W, AA, originally SS/SF), 35- , 50 km from Araçuaí (17°04’39”S, 42°15’25”W, SV/AA, originally SSF), 36 - Baixa Quente (16°57’00”S, 42°04’00”W, AA, originally SS/SF), 37 - Chapada do Lagoão, Serra do Tombo and Calhaquizinho (16°51’00”S, 42°04’13”W, AA, originally SS/ SF), 38 - Trilha do Rio Piauí (16°49’15”S, 41°52’47”W, AA, originally SS/SF), 39 - Fazenda Arqueana, Piauí river (16°43”S, 41°53’W, S/SF); Itinga, 40 – Taquaral (16°43’S, 41°52’W, S/SF), 41 - Fazenda Santana (16°40’S, 41°59’W, AA, originally SS/ SF), 42 - Fazenda Santa Maria (16°36’S, 41°56’W, AA, originally SS/SF), 43 - Fazenda Ilha (16°39’42.6”S, 41°52’5.1”W, S/ SF), 44 - Fazenda Dona Marília (16°36’46.1”S, 41°49’35.5”W, S/SF), 45 - Fazenda Galiléia (16°34’56.4”S, 41°47’23.4”W, S/SF); Salinas, 46 - Fazenda Bamburral, Fazenda Umburana and Ribeirão Bananal (16°10’15”S, 42°17’26”W, AA, originally SS/SF); , 47 – nln (15°46’06”S, 42°14’33”W, SV/AA, originally SDF); , 48 – nln (15°36’02”S, 42°32’22”W, S/SF); Padre Paraíso, 49 - Fazenda Palmares 17°07’18.6”S, 41°36’48.6”W, SV/AA, originally SSF); , 50 - Galery forest from the Piauí river, BR367 road (16°33’42”S, 41°30’12”W, ASS); Joaíma, 51 - Joaíma-Santana de Araçuaí road (16°45’24.6”S, 41°26’47.5”W, SSF), 52 - Fazenda Anta Podre (16°40’S, 41°59’W, AA, originally SS/SF), 53 - Fazenda Anga Pehy (16°43’7.6”S, 41°14’57.6”W, SV/AA, originally SDF); Jequitinhonha, 54 - Mata da Torre da Telemig (16°21’S, 41°05’W, SV/AA, originally SSF), 55 - Reserva Biológica Federal da Mata Escura (16°24’28”S, 41°02’05”W SV/AA, originally SDF), 56 – nln (16°25’56”S, 41°00’06”W, SV/AA, originally SSF); Almenara, 57 - Fazenda Limoeiro (16°02’57”S, 40°51’02”W, SV/AA, originally SSF), 58 - nln (16°03’31”S, 40°39’26”W, SV/AA, originally SSF), 59 - Fazenda Estancia Betania (16°01’S, 40°51’W, SV/AA, originally SSF), 60 – Copasa (15°51’19”S, 40°38’55”W, SV/AA, originally SSF); Bandeira, 61 - Fazenda Serra Azul (15°48’64.4”S, 40°30’86.1”W, SV/AA, originally SSF), 62 - nln (15°52’59”S, 40°34’00”W, SV/AA, originally SSF); Jordania, 63 - nln (15°54’00”S, 40°10’59”W, SV/AA, originally SDF); , 64 - Fazenda Duas Barras (16°14’03”S, 40°08’42”W, SV/AA, originally SSF), , 65 - Fazenda Santana (15°57’S, 40°05’W, SV/AA, originally SDF), 66 – nln (16°00’57”S, 39°56’53”W, SDSF). BAHIA STATE; Itarantim, 67 - Fazenda Boa Vista (15°53’S, 40°09’W, SV/AA, originally SDF), 68 - Fazenda Bom Jardim (15°39’15”S, 40°03’48”W, SV/AA, originally SDF), 69 - Fazenda Alsácea (15°30’S, 40°04’W, SV/AA , originally); Itapebi, 70 - Fazenda Santa Inês (15°46’S, 39°40’W, SV/AA, originally SSF), 71 - nln (15°57’03”S, 39°32’02”W, SV/AA, originally MBF); Mascote, 72 - Fazenda São José (15°34’05”S, 118 Mastozoología Neotropical, 24(1):95-119, Mendoza, 2017 L Geise et al. http://www.sarem.org.ar - http://www.sbmz.com.br

39°17’07”W, S/BF); Belmonte, 73 - Fazenda Futurosa, Santa Maria Eterna (15°51’28”S, 39°24’38.2”W, AA, originally S/ BF); Passuí, 74 – nln (15°51’S, 38°54’W, MA); Santa Cruz de Cabrália, 75 – Reserva Particular do Patrimônio Natural Estação Veracel (16°21’11.6”S, 39°06’49.6”W, MBFL).

APPENDIX II Mammal species from the Jequitinhonha River Basin. Species are listed in alphabetic order. Speci- mens captured by us are indicated by a star (*), specimens recovered from roads (road kill) are indicated by (RK) and karyotyped are indicated by a letter K (K). Localities are numbered as in Appendix I and Fig. 1, and specimens are ordered by sex. F = females, M = males, U shows a specimen of undetermined sex. Acronyms are: Museu Nacional do Rio de Janeiro (MN), Museu de Zoologia da USP (MZUSP), Coleção do Laboratório de Mastozoologia e Manejo da Fauna do Departamento de Zoologia, Universidade Federal de Minas Gerais (UFMG and CC), Museu de Ciências Naturais da PUC Minas (MCN-M), Museu de Zoologia da Universidade Federal de Viçosa (MZUFV) and Coleção de Mastozoologia da Universidade Federal dos Vales do Jequitinhonha e Mucuri (MDIA)). Field number acronyms are LG (L. Geise). After first citation of an acronym, only the number is given.

Akodon cursor -17 (F - UFMG2668, 2669K); 49 (M - MN82750*K); 53 (F - MN82738*K, 82741*K, 82742*K, 82743*K, 82744*K, 82745*K, 82746*K, 82748*K; M - MN82739*K, 82740*K, 82747*K, 82749*K). Alouatta guariba - 33 - (U - UFMG1473). Callithrix penicillata - 46 (M - MCN-M38); 74 (M - MN23794). Calomys mattevii - 37 (U - MN42843); 44 (F - MN81105*K, 81106*K, 81107*K, 81109*K, 81110*K, 81113*, 81114*, 81115*; M - MN81108*K, 81111*K; U – MN81112*). Calomys tener - 3 (M - UFMG1608); 12 (F - UFMG2454; M - UFMG2452, 2453); 27 (F - MCN-M1105); 34 (F - MCN- M1089, 1106; M - MCN-M1094); 46 (M - MN42840); 47 (F - MCN-M732); 48 (F - MCN-M728, U- MCN-M727). Caluromys philander - 7 (M - MDIA43*); 8 (M - MN82751*K); Cavia aperea - 5 (F - MCN-M1046; M - MDIA52*). Cerdocyon thous - 43 (F - LG 1167RK); 50 (U – UFMG 3078). Cerradomys scotti - 5 (M - MDIA24*); 6 (F - MN82753*K; M - MN82752*K); 7 (M - MDIA49*). Cerradomys subflavus - 3 (U - UFMG1609, 1610, 1611); 5 (M - MDIA51*, 57*); 6 (F - MN82759*K); 7 (F - MDIA16*; M - UFMG2852, 2853; MDIA23*); 9 (F - MN82758*); 11 (M - MCN-M1); 12 (M - UFMG2457, 2458); 22 (F - UFMG1459); 27 (F - MCN-M1075); 35 (M - MZUFV1877, 1923); 45 (M - UFMG1453, 1454); 46 (F - MN42841, MCN-M332; M - MN42844); 48 (M - MCN-M725); 50 (F - UFMG907); 53 (F - MN82754*K, 82757*K; M - MN82755*K, 82756*K). Cerradomys vivoi - 45 (F - MN82762*, 82763*K, 82764*K, 82765*K; M - MN82760*K, 82761*K, 82766*K); 54 (M - UFMG1458). Conepatus semistriatus - 19 (M - MCN-M99); 36 (U - MN43924); 46 (U - MN42855). Dasypus novemcinctus - 3 (U - UFMG3085, 3086); Dasyprocta leporina -18 (M - UFMG1565) Didelphis albiventris - 2 (F - MN82769*); 5 (M - MDIA50*); 9 (F - MN82770*K); 33 (F - UFMG1471; M - UFMG1472); 38 (U: UFMG1566); 43 (M - MN82771*); 45 (F - MN82767*); 46 (M - MN42834); 49 (F - MN82772*); 53 (F - MN82768*); 56 (F - UFMG905). Didelphis aurita - 49 (F - MN82777*); 75 (F - MN82774*, 82775*; M - MN82773*, 82776*). Euphractus sexcinctus - 15 (U - UFMG1105); 16 (U - UFMG3093); 46 (U - MN42849, 42851, 42852). Euryoryzomys russatus - 53 (M - MN82778*K); 54 (M - UFMG1457); 72 (M - CC37). Euryzygomatomys spinosus - 5 (M - MDIA78*; 84*). Gracilinanus agilis - 2 (F - MN72692*); 3 (U – MCN-M315); 7 (M – MDIA06*; 14*, 19*; F - MDIA15*, UFMG2493, 2494); 9 (M - MN72693*K); 12 (F - UFMG2433, 2434, 2435); 31 (F - UFMG2495); 37 (M - MN42842; U - MN42845); 43 (F - MN82782*K, 82783*K, 82784*K, 82785*K, 82786*K; M - MN82779*K, 82780*K, 82781*K); 45 (F - MN72687*, 72690*, 72691*; M - MN72686*K, 72688*, 72689*; U - UFMG1464); 47 (F - MCN-M734). Gracilinanus microtarsus - 2 (F - MN72702*K); 6 (F - MN82788*K); 7 (F - MDIA44*); 9 (F - MN82787*K; M - MN72701*K); 60 (U - UFMG1465). Galea spixii - 34 (U - MCN-M989); 48 (F - MCN-M735, M - MCN-M761). Guerlinguetus brasiliensis - 49 (F – MN82791*K; M - MN82790*K, 82792*K); 75 (F – MN82789*). Hydrochoerus hydrochaeris - 71 (U - UFMG1048). Hylaeamys seuanezi - 72 (M - CC35). Kerodon rupestris - 13 (U - UFMG1101). Leopardus wiedi - 50 (U - UFMG3076). TERRESTRIAL MAMMALS OF THE JEQUITINHONHA RIVER BASIN, BRAZIL 119

Metachirus nudicaudatus - 7 (F - MDIA12*; M - MDIA13*); 17 (U - UFMG2654, 2655); 54 (F - UFMG1462; M - UFMG1568; U – UFMG1567); 75 (F - MN82847*; M - MN82846*K). Marmosa paraguayana - 7 (M - UFMG2648); 8 (F - MN82795*K; M - MN82794*K, 82796*K, 82797*K); 9 (M - MN82799*K, 82800*K); 17 (F - UFMG2611; M - UFMG2612, 2613); 30 (F - MCN-M36); 46 (F - MCN-M26); 75 (M - MN82798*). Marmosa murina - 75 (M - MN82793*K). Marmosops incanus - 2 (F - MN82812*K, 82814*K, 82815*K; M - MN82813*K); 3 (F - UFMG1607); 5 (M - MDIA69*, 85*); 6 (F - MN82820*K, 82821*, 82822*K, 82823*K); 7 (M - MDIA04*; UFMG2601); 8 (M - MN82808*K, 82809*K, 82810*K); 9 (F - MN82817*K, 82818*K; M - LG596*K, MN82816*K, 82819*K); 11 (F - MCN-M326); 12 (F - UFMG2436, 2440, 2441, 2442; M - UFMG2437, 2438); 14 (U - MN33840, 34426); 17 (F - UFMG2605); 27 (F - MCN-M1078, 1111); 28 (F - UFMG1102); 30 (F - MCN-M46; M - MCN-M318); 31 (U - UFMG2610); 33 (M - UFMG1468, 1469, 1470); 45 (M - MN82801*, 82802*K, 82803*); 49 (F - MN82824*K, 82828*K, 82829*K, 82830*, 82831*, 82835*, 82838*, 82839*K, 82840*K, 82842*, 82843*; M - MN82825*K, 82826*K, 82827*K, 82832*, 82833*, 82834*, 82836*, 82837*, 82841*, 82844*, 82845*); 53 (F - MN82804*; M - MN82805*, 82806*, 82807*K); 60 (M - UFMG1466, 1467). Monodelphis americana - 9 (M - MN82848*K); 49 (M - MN82849*K, 82850*K). Monodelphis domestica - 5 (M - MDIA47*; U - MCN-M1042); 6 (F - MN82866*K, 82867*K); 9 (F - MN82863*, 82864*K; M - MN82868*, 82865*K); 12 (F - UFMG2439); 30 (M - MCN-M41); 35 (M – MZUFV1925, 1926); 45 (F - MN82851*, 82852*K, 82856*, 82857*K, 82860*, 82861*, UFMG1463; M - MN82853*K, 82854*, 82855*, 82858*, 82859*, 82862*); 46 (U - MN46588, 46589; F - MCN-M24). Monodelphis scalops - 5 (M - MDIA86*). Necromys lasiurus - 5 (M - MDIA53*); 27 (M - MCN-M1079); 30 (F - MCN-M334); 46 (F - MCN-M35). Nectomys squamipes- 5 (M - MDIA75*); 7 (M - UFMG2984, 2985); 8 (F - MN82881*K; M - MN82880*K, 82882*K); 9 (M - MN82883*K, 82884*K); 24 (M - UFMG1451, 1452); 49 (F - MN82886*K); 53 (F - MN82869*K, 82872*K, 82876*, 82877*; M - MN82870*K, 82871*K, 82873*K, 82874*K, 82875*, 82878*, 82879*); 75 (F - MN82885*K; M - MN82887*K). catherinae - 54 (M - UFMG1455); 72 (M - CC36). Oligoryzomys nigripes - 5 (F - MDIA60*); 7 (M - MDIA11*); 9 (M - MN82888*K); 12 (F - UFMG2456, 2455); 31 (F - UFMG2743); 33 (F - UFMG1479); 47 (F - MCN-M733). Oxymycterus dasytrichus - 5 (M - MDIA63*); 54 (F - UFMG1476; M - UFMG1475). Philander frenatus- 9 (F - MN82889*K, 82890*, 82891*, 82892*). Phyllomys lamarum - 17 (U - UFMG3016); 33 (M - UFMG1491). Rhipidomys macrurus - 6 (M - MN82893*K); 12 (F - UFMG2460, 2461, 2462, 2463, 2464, M - UFMG2465), 31 (F - UFMG2934). Rhipidomys mastacalis - 3 (U - UFMG1605, 1606); 4 (U – MCN-M49); 5 (M - MDIA76*, 77*); 6 (M - MN82896*K); 7 (F - UFMG2926, 2927, 2928, M - UFMG2925, 2929); 9 (M - MN82895*K), 17 (F - UFMG2930); 33 (F - UFMG1450); 46 (F – MN30021, U - MN30015); 49 (F - MN82897*K, 82900*K; M - MN82898*K, 82899*K); 54 (F - UFMG1461; M - UFMG1456, 1460); 73 (F - MN82894*K). Sapajus robustus - 74 (F - MN23230, 23233, M - MN23228, 23229, 23232, 23234). Sooretamys angouya - 12 (M - MCN-M1100). Sylvilagus minensis - 48 (M - MCN-M860); 51 (M - MN82901RK). Thalpomys lasiotis - 6 (M - MDIA9*). Thrichomys apereoides - 2 (F - MN82914*K, 82915*K, 82916*K, 82917*, 82921*K, 82922*; M - MN82913*K, 82918*K, 82919*K, 82920*K); 5 (M - MDIA54*, 55*, 56*); 6 (F - MN82932*, 82934*K; M - MN82933*K); 7 (F - UFMG3005, 3006, 3007); 8 (F - MN82908*K, 82910*K, 82912*; M - MN82909*K, 82911*K); 9 (F - MN82923*K, 82927*K, 82928*K; M - MN82924*K, 82925*, 82926*K, 82929*K, 82930*K, 82931*K); 12 (F - UFMG2467, M - UFMG2468, 2469, 2472); 15 (U - UFMG1096); 24 (F - UFMG1489, 1490); 27 (U - MCN-M1065); 28 (M - UFMG1098, 1482, 1486, 1487; MCN- M1044; F - UFMG1480, 1481, 1483, 1484, 1485, 1488); 31 (U - UFMG3009); 35 (U – MZUFV1929, 1930, 1934; F - MZUFV1927; M - MZUFV1928); 44 (F - MN82903*K, 82904*K, 82905*K, 82906*K, 82907*K); 45 (F - MN82902*K); 47 (M - MCN-M762, 763). Thylamys velutinus- 48 (F - MCN-M724). Trinomys albispinus - 5 (F - MCN-M1132, M - MCN-M1033, 1131); 10 (MN 73436); 12 (F - UFMG2478, 2479, 2480, M - UFMG2473, 2474, 2475, 2476); 22 (F - UFMG1445, 1446, 1447; M - UFMG1441); 23 (F - UFMG1439, M - UFMG1442, 1448); 27 (F - MCN-M1073, 1076; M - MCN-M1050, 1067, 1068, 1074; U - MCN-M1039, MN73429); 28 (U - UFMG1103, F - MCN-M1036); 45 (M - MN82935*K); 54 (M - UFMG1440); 65 (F - MCN-M972); 66 (F - MCN-M993, 994, 996; M - MCN-M992, 995, 997). Trinomys setosus - 9 (F - MN82937*K); 12 (M - UFMG2477); 17 (F - UFMG3036, 3037, 3038); 23 (M - UFMG1436); 53 (M - MN82936*K); 54 (F - UFMG1437, 1438, M - UFMG1449); 58 (MNRJ 73409); 60 (U - UFMG1443, 1444). Wiedomys pyrrhorhinos - 7 (M - MDIA80*); 9 (F - MN82940*K, 82942*K; M - MN82941*K); 11 (F - MCN-M3); 12 (F - UFMG2466); 28 (F - UFMG1100; U - MCN-M1052); 34 (M - MCN-M1054); 43 (F - MN82943*K, 82944*K); 45 (F - MN82938*K, 82939*K, M - UFMG1474).