Journal of South American Earth Sciences 82 (2018) 76e90 Contents lists available at ScienceDirect Journal of South American Earth Sciences journal homepage: www.elsevier.com/locate/jsames The southernmost record of a large erethizontid rodent (Hystricomorpha: Erethizontoidea) in the Pleistocene of South America: Biogeographic and paleoenvironmental implications * Raúl I. Vezzosi a, , Leonardo Kerber b a Laboratorio de Paleontología de Vertebrados, Centro de Investigaciones Científicas y Transferencia de Tecnología a la Produccion, Consejo Nacional de Investigaciones Científicas y Tecnicas, Materi y Espana,~ E3105BWA, Diamante, Argentina b CAPPA - Centro de Apoio a Pesquisa Paleontologica da Quarta Colonia,^ Universidade Federal de Santa Maria, Sao~ Joao~ do Pol^esine, Rua Maximiliano Vizzotto, 598, CEP 97230-000, Brazil article info abstract Article history: The South American porcupines (Erethizontidae) are included in two genera: Chaetomys and Coendou. Received 19 October 2017 The latter is a very speciose taxon, with about 13 living species. During at least the late Plioceneeearly Received in revised form Pleistocene, erethizontids immigrated to Central and North America during the Great American Biotic 23 December 2017 Interchange. Although some Pleistocene fossils have been reported, the Quaternary history of this clade Accepted 24 December 2017 is still understudied. The only known extinct species is Coendou magnus. In this work, a fossil of a Available online 30 December 2017 porcupine is reported from an Upper Pleistocene fluvial sedimentary sequence cropping out in the Northern Pampa geomorphological region, Santa Fe Province, Argentina. Despite this group having Keywords: Fossil record different living forms widely distributed in South American Neotropical woodland habitats, the Pleis- Quaternary tocene occurrences of Erethizontidae are scarce and limited to Upper Pleistocene deposits from Bolivia, Regional extinction Brazil, and Uruguay. Currently, the specimen here reported represents the only Pleistocene porcupine Distribution displacement from Argentina with a stratigraphical context. The morphological characters as well as the dimensions Environmental changes indicate that it is close to the Pleistocene erethizontid Coendou magnus. In this context, the presence of this erethizontid in such a southern locality, together with other taxa recorded from this site and the associated geological and paleoenvironmental evidence, indicates subtropical conditions, compared with the current conditions, which may have allowed a southern displacement of taxa more related to woodlands and xeric subtropical environments. © 2017 Elsevier Ltd. All rights reserved. 1. Introduction Patterson, 2015). Commonly known as porcupines, or ‘puercoes- pines’ in the Spanish language, this group is restricted to the Among Hystricognathi rodents, the Erethizontidae (the New Americas. The living representatives of this clade include three World porcupines) is a clade morphologically disparate from other living genera allocated to two ‘subfamilies’ that collectively range caviomorphs, which has led some authors to suppose that they from boreal North America (Alaska and northern Canada) to sub- were the first lineage to differentiate from other forms during the tropical South America (Uruguay and northern Argentina): Coendou early evolutionary history of Caviomorpha (Bugge, 1971; Woods, Lacep ede, 1799, Erethizon Cuvier, 1822 and Chaetomys Gray, 1843 1972; Bryant and McKenna, 1995; Candela, 1999). However, with (see Tate, 1935; Woods, 1973; Alberico et al., 1999; Bonvicino et al., the increase in the resolution of time-calibrated phylogenies based 2002; Voss, 2011; Voss et al., 2013). From a taxonomical approach, on molecular data, today they are considered the sister-group to at least 13 living Neotropical species of Coendou inhabit Central to Cavioidea, with a simultaneous period of diversification of the South America in tropical and subtropical moist and dry forests other higher-level clades within Caviomorpha (Upham and from sea level to about 3500 m (Voss and Angermann, 1997; Voss and da Silva, 2001; Voss, 2015). The semi-arboreal Erethizon dor- satum (Linnaeus, 1758) is the only monotypic extant species in * Corresponding author. temperate North America, from northern Mexico to Alaska and E-mail addresses: [email protected] (R.I. Vezzosi), leonardokerber@gmail. Canada (Roze, 1989). The southernmost distribution of Coendou is com (L. Kerber). https://doi.org/10.1016/j.jsames.2017.12.015 0895-9811/© 2017 Elsevier Ltd. All rights reserved. R.I. Vezzosi, L. Kerber / Journal of South American Earth Sciences 82 (2018) 76e90 77 recorded in northern Argentina and Uruguay (Anderson, 1997; Here, we expand the fossil record of South American Quaternary Emmons, 1997; Barquez et al., 2006; Gonzalez and Martínez erethizontids and report the first undoubted occurrence for the Lanfranco, 2010). These forms are strictly adapted to an arboreal Pleistocene of Argentina, recovered from fluvial sedimentary de- life, thereby restricting their home range. Nevertheless, there are posits of the Upper Pleistocene sequence in the Carcaran~a River also species distributed in open and transitional areas such as the fossiliferous area. Moreover, we discuss the relevance of Erethi- Cerrado and Caatinga in Brazil, the Venezuelan plains in Venezuela, zontidae in the austral temperate plains. and the austral Chaco in Bolivia and northern Argentina (Roze, This area is an interesting fossiliferous site that has yielded a 1989; Anderson, 1997; Parera, 2002; Candela and Morrone, 2003; large number of megamammal specimens for many years Barquez et al., 2006). In this sense, the particular adaptation of (Frenguelli, 1928; Castellanos, 1943; Bordas, 1942; Brandoni and the Neotropical erethizontids to a particular environment such as McDonald, 2015; Vezzosi, 2015). However, many of them are woodlands (Candela and Morrone, 2003) could suggest that their without an accurate stratigraphical context (see discussion in evolution and geographical distribution were driven by this life- Vezzosi, 2015; Vezzosi et al., 2017). The specimen here described is style, which is strongly associated with these environments. possibly related to the only extinct species of Quaternary erethi- Fossil records are known from the late Oligocene to middle zontid from South America. This taxon was erected in the nine- Miocene of Patagonia, the Oligocene of Bolivia, and from the middle teenth century based on a few records from Brazilian Holocene Miocene of La Venta in Colombia (Ameghino, 1887; Simpson, 1950; caves and is the largest extinct South American porcupine that lived Wood and Patterson, 1959; Hoffstetter and Lavocat, 1970; Walton, in PleistoceneeHolocene times (Lund, 1839; Winge, 1887). 1997; Candela, 1999, 2002). During the late Miocene (Huay- querian SALMA or Tortonian Global Stage/Age; Bossi and Muruaga, 2. Materials and methods 2009) the taxonomic richness and distribution of the group decreased drastically in southern South America; only the giant In order to identify and define anatomical and proportional porcupines Neosteiromys bombifrons Rovereto, 1914 and Neo- differences in the fossil from Santa Fe, we studied numerous den- steiromys pattoni Candela, 2004 survived in northwestern taries and teeth series of fossil and living erethizontids, which have Argentina, as well as the monospecific Paradoxomys cancrivorus sufficient anatomical complexity to permit the recognition of Ameghino, 1885 from northeastern Argentina (‘Conglomerado consistent differences between the genera and species. Osífero’, lower member of the Ituzaingo Fm., Huayquerian Stage/ Age; Frenguelli, 1920; Vucetich and Candela, 2001; Brunetto et al., 2.1. Anatomical terminology 2013). PlioePleistocene (Stage/Age late Blancaneearly Irvingtonian) The anatomical nomenclature follows Woods and Howland erethizontids from North America belong only to the genus Ere- (1979), Frazier (1981), Perez (2010), and Sussman (2011) for the thizon (Frazier, 1981; Simpson, 1950; Webb, 1976; Morgan and dentary, and Marivaux et al. (2004) for the tetralophodont molars: Hulbert, 1995; Hulbert, 1997). However, many Erethizon species Afd, anterofossetid; chin, posterior joint of the symphysis; cp, have been assigned to the South American genus Coendou with coronoid process; cop, condylar process; hc, horizontal crest; hg, some new species (White, 1968, 1970; Sussman et al., 2016); hystricognathous groove, Hfd, hypoflexid; Hld, hypolophid; HD, although, the assignments of extinct species to this genus (versus height of dentary; i, incisor; ln, lunar notch; LS, length of the dental Erethizon) are sometimes controversial (Sussman, 2011). series; LLW, labio-lingual width; MDW, mesio-distal width; m; Erethizon dorsatum is well recorded in the loweremiddle molar; mc, masseteric crest; maf, mandibular foramen; mf, men- Pleistocene (Stage/Age middle Irvingtonian) and Holocene of North tonian foramen; mn, mandibular notch; Med I, metalophid I; Med America (Frazier, 1981; Hulbert, 1997). In contrast, the reliable fossil II, metalophid II; Mfd, metaflexid; Msd, mesoflexid; pma, premolar record of South American porcupines is very scarce towards the alveolus; ptc, pterygoid crest; Psd, posterolophid; rf, retromolar southern portion of the continent, encompassing the extinct spe- fossa; sys, symphysis. cies Coendou magnus Lund, 1839 from the middleelate Pleistocene of Bolivia (Tarija Valley; Hoffstetter, 1963), late Pleistoceneeearly 2.2. Institutional abbreviations