applyparastyle “fig//caption/p[1]” parastyle “FigCapt” Zoological Journal of the Linnean Society, 2021, 191, 575–611. With 14 figures. Taxonomic revision of the pampas cat Leopardus colocola complex (Carnivora: Felidae): an integrative approach FABIO OLIVEIRA DO NASCIMENTO1,*†, , JILONG CHENG2, , and ANDERSON FEIJÓ2,*†, Downloaded from https://academic.oup.com/zoolinnean/article/191/2/575/5848237 by guest on 28 September 2021 1Mastozoologia, Museu de Zoologia da Universidade de São Paulo, São Paulo, SP, Brazil 2Key Laboratory of Zoological Systematics and Evolution, Institute of Zoology, Chinese Academy of Sciences, Beichen West Road, Chaoyang District, Beijing 100101, China Received 31 December 2019; Revised 31 March 2020; accepted for publication 31 March 2020 The pampas cat Leopardus colocola has been subject to conflicting classifications over the years. Currently, one polytypic species with seven subspecies is recognized, but integrative taxonomic study for this debated group has never been done. Here, we combine the broadest morphological coverage of the pampas cat to date with molecular data and ecological niche models to clarify its species composition and test the validity of recently proposed subspecies. The multiple lines of evidence derived from morphology, molecular, biogeography and climatic niche datasets converged on the recognition of five monotypic species: L. braccatus, L. colocola, L. garleppi (including thomasi, budini, steinbachi, crespoi and wolffsohni as synonyms), L. munoai and L. pajeros (including crucina as synonym). These five species are morphologically diagnosable based on skin and skull traits, have evolved in distinct climatic niche spaces and were recovered in molecular species delimitation. Contrary to previous taxonomic arrangements, we do not recognize subspecies in pampas cats. To objectively define the two most controversial species, we designate neotypes for L. colocola and L. pajeros. The diversification of pampas cats is associated with Middle Pleistocene glaciations, but additional genetic samples from the central Andean region are still needed to conclusively reconstruct its evolutionary history. ADDITIONAL KEYWORDS: Andes – ocelot – open-dwellers – Pleistocene glaciations – species delimitation – subspecies. INTRODUCTION from sea level up to 5000 m (Redford & Eisenberg, 1992; García-Perea, 1994; Nowak, 1999; Sunquist & The pampas cat, Leopardus colocola (Molina, 1782), Sunquist, 2002, 2009). Phylogenetic studies show that is a South American small-sized felid (± 3 kg) pampas cats are part of the group informally known characterized by long hairs on the body, an erectile as the ‘ocelot lineage’, which corresponds to small- and spinal crest slightly darker than ground colour, medium-sized Neotropical spotted cats of the genus transverse dark stripes on the throat, markings on Leopardus Gray, 1842 (Johnson et al., 1999, 2006; the flanks, legs with transverse dark stripes in the Mattern & McLennan, 2000; Li et al., 2016; Kitchener proximal portion, ears more pointed and tail relatively et al., 2017). Pampas cats are closely related to the shorter than other South American felids and a Andean mountain cat, Leopardus jacobita (Cornalia, lingual cavity between lower canine teeth (Pocock, 1865), the oncilla or tigrina, Leopardus tigrinus 1941; Cabrera, 1958, 1961; Guggisberg, 1975; Redford (von Schreber, 1775) species group, Geoffroy’s cat, & Eisenberg, 1992; Salles, 1992; García-Perea, 1994; Leopardus geoffroyi (d’Orbigny & Gervais, 1844) and Eisenberg & Redford, 1999; Sunquist & Sunquist, the kodkod, Leopardus guigna (Molina, 1782) (Johnson 2002, 2009; Prevosti, 2006). Pampas cats are typical of & O’Brien, 1997; Johnson et al., 1999, 2006; Mattern & open areas and show a broad elevational distribution McLennan, 2000; Li et al., 2016). The taxonomic history of pampas cats started *Corresponding authors. E-mail: [email protected]; with Molina (1782) describing Felis colocola, based [email protected] on a cat from the forests of Chile (‘Boschi del Chili’). †The authors contributed equally to the present study. Since then, numerous related species and subspecies © 2020 The Linnean Society of London, Zoological Journal of the Linnean Society, 2021, 191, 575–611 575 576 F. O. DO NASCIMENTO ET AL. have been described, usually based on a single or a when Felis jacobita Cornalia, 1865 (= Leopardus few specimens: Felis pajeros Desmarest, 1816 from jacobita) was erroneously associated with the name Argentina, Felis braccata Cope, 1889 from central colocola (Philippi, 1869, 1870, 1873; Burmeister, 1879; Brazil, Felis pajeros crucina Thomas, 1901 from Lydekker 1896; Allen, 1919; Yepes, 1929; Pocock, 1941; southern Argentina, Lynchailurus pajeros garleppi Schwangart, 1941). Matschie, 1912 from Peru, Felis pajeros thomasi Throughout the 20th and early 21st centuries, a Lönnberg, 1913 from Ecuador, Lynchailurus pajeros series of taxonomic studies involving pampas cats budini Pocock, 1941 from north-western Argentina, was carried out, resulting in contrasting taxonomic Lynchailurus pajeros steinbachi Pocock, 1941 schemes (Table 1). In the most recent revision of from Bolivia, Felis (Lynchailurus) colocolo crespoi the group, García-Perea (1994), based on skull and Downloaded from https://academic.oup.com/zoolinnean/article/191/2/575/5848237 by guest on 28 September 2021 Cabrera, 1957 from north-western Argentina, Felis external morphological characters of 86 specimens, colocola muñoai Ximénez, 1961 from Uruguay and recognized three allopatric polytypic species: Lynchailurus colocolo wolffsohni García-Perea, 1994 Lynchailurus colocolo (including Ly. c. colocolo and from northern Chile. Ly. c. wolffsohni) distributed on the western slope of The use of the specific name colocola and its the southern Andes (northern and central Chile), derivations (colocolo, colorolla, colicollo, etc.) (Molina, Ly. pajeros (including Ly. p. pajeros, Ly. p. crucinus, 1782, 1786, 1788, 1789, 1808, 1809, 1810; Bechstein, Ly. p. budini, Ly. p. steinbachi, Ly. p. crespoi, 1800; Shaw, 1800; Fischer, 1829; Wagner, 1841) has Ly. p. garleppi and Ly. p. thomasi) distributed along the been historically debated since the late 18th century Andes, from Ecuador to southern Chile and Argentina (Cabrera, 1940, 1958, 1961; Osgood, 1943; Kitchener and Ly. braccatus (including Ly. b. braccatus and et al., 2017). The main problem started when Griffith Ly. b. munoai) found in Brazil, Paraguay and Uruguay. (1821) and Griffith et al. (1827) associated Molina’s This arrangement was largely followed by Wozencraft F. colocola with an unidentified specimen from (2005), but he placed the group in the genus Leopardus Guyana based on information and plates provided by and considered crespoi Cabrera, 1957 and crucina Charles Hamilton Smith, an opinion largely followed Thomas, 1901 as junior synonyms of L. p. budini in subsequent works (e.g. Fischer, 1829; Jardine, 1834; (Pocock, 1941). Later, Nascimento (2010), in his Reichenbach, 1834; Gay, 1847; Giebel, 1855; Gray, 1867; taxonomic revision of genus Leopardus, recognized six Fitzinger, 1869; Mivart, 1881; Trousseart, 1885, 1897, species for the pampas cat species complex: Le. colocolo, 1904; Elliot, 1883; Matschie, 1894, 1895), although Le. pajeros, Le. braccatus, Le. munoai, Le. garleppi rejected by others (e.g. Swainson, 1838; Wagner, and Le. budini. Recently, Kitchener et al. (2017), 1841; Gray, 1874). The confusion became even greater evaluating previous studies, recognized only one species Table 1. Main taxonomic arrangements of L. colocola since the beginning of the 20th century Authors Taxa Allen (1919) Lynchailurus pajeros pajeros, Ly. p. crucina, Ly. p. thomasi, Ly. p. thomasi, Ly. p. braccatus Cabrera (1940) Lynchailurus colocolus colocolus, Ly. c. pajeros, Ly. c. braccatus, Ly. c. crucinus, Ly. c. garleppi, Ly. c. thomasi Pocock (1941) Lynchailurus pajeros pajeros, Ly. p. huina, Ly. p. budini, Ly. p. steinbachi, Ly. p. garleppi, Ly. p. braccatus Schwangart (1941) Lynchailurus (Lynchailurus) pajeros pajeros, Ly. (Ly.) p. braccatus, Ly. (Ly.) p. garleppi Cabrera (1958) Felis (Lynchailurus) colocolo braccatus, F. (Ly.) c. budini, F. (Ly.) c. colocolo, F. (Ly.) c. crespoi, F. (Ly.) c. garleppi, F. (Ly.) c. pajeros, F. (Ly.) c. thomasi Ximénez (1961) Felis colocola colocola, F. c. braccata, F. c. budini, F. c. crespoi, F. c. garleppi, F. c. pajeros, F. c. thomasi, F. c. muñoai García-Perea (1994) Lynchailurus colocolo: Ly. c. colocolo, Ly. c. wolffsohni Lynchailurus pajeros: Ly. p. budini, Ly. p. crespoi, Ly. p. crucinus, Ly. p. garleppi, Ly. p. pajeros, Ly. p. steinbachi, Ly. p. thomasi Lynchailurus braccatus: Ly. b. braccatus, Ly. b. munoai Wozencraft (2005) Leopardus colocolo: Le. c. colocolo, Le. c. wolffsohni Leopardus pajeros: Le. p. budini, Le. p. garleppi, Le. p. pajeros, Le. p. steinbachi, Le. p. thomasi Leopardus braccatus: Le. b. braccatus, Le. b. munoai Kitchener et al. (2017) Leopardus colocola colocola, Le. c. wolffsohni, Le. c. pajeros, Le. c. budini, Le. c. garleppi, Le. c. braccatus, Le. c. munoai Present study Leopardus colocola, Le. pajeros, Le. garleppi, Le. braccatus, Le. munoai © 2020 The Linnean Society of London, Zoological Journal of the Linnean Society, 2021, 191, 575–611 TAXONOMY OF LEOPARDUS COLOCOLA 577 with seven subspecies: Leopardus colocola colocola, Coleção de Mamíferos, Universidade de Brasília, Le. c. wolffsohni, Le. c. pajeros (including crucina), Brasília, Brazil (UnB); Estación Biológica de Doñana, Le. c. budini (including steinbachi), Le. c. garleppi