Leopardus garleppi

Nascimento, Fabio Oliveira Do, Cheng, Jilong & Feijó, Anderson, 2021, Taxonomic revision of the pampas cat Leopardus colocola complex (Carnivora: Felidae): an integrative approach, Zoological Journal of the Linnean Society 191, pp. 575-611 : 598-602

LEOPARDUS GARLEPPI ( MATSCHIE, 1912), COMB. NOV.

( FIG. 13)

Garlepp’s pampas cat or northern pampas cat.

Felis pajeros: Matschie, 1894: 60 ; part; not Felis pajeros Desmarest.

Felis (Lynchailurus) pajeros garleppi Matschie, 1912: 259 ; type locality: ‘von Cuzco in Südost-Peru, im Gebiet des Apurimac, der durch den Ucayali zum oberen Amazonas abwässert’ .

Felis pajeros thomasi Lönnberg, 1913: 7 ; type locality: ‘near Quito, Ecuador’.

Felis pajeros garleppi: Lönnberg, 1913: 8 ; name combination.

Lynchailurus pajeros thomasi: Allen, 1919: 376 ; name combination.

Lynchailurus pajeros garleppi: Allen, 1919: 376 ; name combination.

Felis (Lynchailurus) colocolo garleppi: Pearson, 1951: 130 , 137; name combination.

Lynchaylurus pajeros pajeros: Yepes, 1936: 36 ; incorrect spelling; not Felis pajeros Desmarest.

L [ynchailurus]. colocolus garleppi: Cabrera, 1940: 12 ; name combination; unjustified emendation of colocola Molina.

L [ynchailurus]. colocolus thomasi: Cabrera, 1940: 12; name combination; unjustified emendation of colocola Molina.

Lynchailurus pajeros budini Pocock, 1941: 263 ; type locality: ‘ Mount Sola, 2500 m., in Salta, northern Argentine’ .

Lynchailurus pajeros steinbachi Pocock, 1941: 264 ; type locality: ‘ Tiraque , Cochabamba, western Bolivia, 4000 m. alt .’.

Lynchailurus pajeros garleppi: Pocock, 1941: 266 ; part ( Felis pajeros thomasi Lönnberg treated as a synonym).

Lynchailurus (Lynchailurus) garleppi: Schwangart, 1941: 33 ; part ( Felis pajeros thomasi Lönnberg treated as a synonym); name combination.

Felis (Lynchailurus) colocolo crespoi Cabrera, 1957: 71 ; type locality: ‘ Aguaray , província de Salta [ Argentina] ’.

Felis [( Lynchailurus )] colocolo budini: Cabrera, 1958: 276; name combination.

Felis [( Lynchailurus )] colocolo crespoi: Cabrera, 1958: 277; name combination.

Felis [( Lynchailurus )] colocolo garleppi: Cabrera, 1958: 277 ; part ( Lynchailurus pajeros steinbachi Pocock treated as a synonym); name combination.

Felis [( Lynchailurus )] colocolo thomasi: Cabrera, 1958: 278; name combination.

F [elis]. colocola budini: Ximénez, 1961: 6; name combination.

F [elis]. colocola crespoi: Ximénez, 1961: 6; name combination.

F [elis]. colocola garleppi: Ximénez, 1961: 6 ; name combination.

F [elis]. colocola thomasi: Ximénez, 1961: 6; name combination.

Lynchailurus colocolo wolffsohni García-Perea, 1994: 31 ; type locality: ‘ río Camarones , provincia Tarapacá, between 2000 and 4000 m, Chile’ .

Lynchailurus pajeros: García-Perea, 1994: 31 ; part; name combination.

L [ynchailurus]. p [ajeros]. budini: García-Perea, 1994: 31; name combination.

L [ynchailurus]. p [ajeros]. crespoi: García-Perea, 1994: 32; name combination.

L [ynchailurus]. p [ajeros]. garleppi: García-Perea, 1994: 32 ; name combination.

L [ynchailurus]. p [ajeros]. steinbachi: García-Perea, 1994: 32; name combination.

L [ynchailurus]. p [ajeros]. thomasi: García-Perea, 1994: 32; name combination.

[ Leopardus colocolo ] wolffsohni: Wozencraft, 2005: 538; name combination.

Leopardus pajeros: Wozencraft, 2005: 538 ; part; name combination.

[ Leopardus pajeros ] budini: Wozencraft, 2005: 539; name combination.

[ Leopardus pajeros ] garleppi: Wozencraft, 2005: 539 ; name combination.

[ Leopardus pajeros ] steinbachi: Wozencraft, 2005: 539; name combination.

[ Leopardus pajeros ] thomasi: Wozencraft, 2005: 539; name combination.

Leopardus colocola: Kitchener, Breitenmoser-Würsten, Eizirik, Gentry, Werdelin, Wilting, Yamaguchi, Abramov, Christiansen, Driscoll, Duckworth, Johnson, Luo, Meijaard, O’Donoghue, Sanderson, Seymour, Bruford, Groves, Hoffmann, Nowell, Timmons & Tobe, 2017: 51 ; part; name combination.

Leopardus colocola wolffsohni: Kitchener, Breitenmoser-Würsten, Eizirik, Gentry, Werdelin, Wilting, Yamaguchi, Abramov, Christiansen, Driscoll, Duckworth, Johnson, Luo, Meijaard, O’Donoghue, Sanderson, Seymour, Bruford, Groves, Hoffmann, Nowell, Timmons & Tobe, 2017: 53 ; part; name combination.

Leopardus colocola budini: Kitchener , Breitenmoser- Würsten, Eizirik, Gentry, Werdelin, Wilting, Yamaguchi, Abramov, Christiansen, Driscoll, Duckworth, Johnson, Luo, Meijaard, O’Donoghue, Sanderson, Seymour, Bruford, Groves, Hoffmann, Nowell, Timmons & Tobe, 2017: 53; part ( Lynchailurus pajeros steinbachi Pocock treated as a synonym); name combination.

Leopardus colocola garleppi: Kitchener , Breitenmoser- Würsten, Eizirik, Gentry, Werdelin, Wilting, Yamaguchi, Abramov, Christiansen, Driscoll, Duckworth, Johnson, Luo, Meijaard, O’Donoghue, Sanderson, Seymour, Bruford, Groves, Hoffmann, Nowell, Timmons & Tobe, 2017: 53; part ( Felis (Lynchailurus) pajeros garleppi Matschie treated as a synonym); name combination.

Type locality: Cuzco, Peru: ‘von Cuzco in Südost-Peru, im Gebiet des Apurimac, der durch den Ucayali zum oberen Amazonas abwässert’ [‘from Cuzco in southeastern Peru, in the region of Apurímac, which drains through the Ucayali to the upper Amazon’] ( Matschie, 1912: 259) .

Type material: ZMB _MAM_21244, skin and skull of unknown sex adult specimen collected by Otto Garlepp.

Diagnosis: Leopardus garleppi is distinguished by the brownish grey forehead, crown and nape speckled with orange; black, dark brown, yellowish brown or dark yellowish brown transversal gular stripes, with at least one of them markedly wider than others; spinal crest dark brownish grey with some orangish hairs; background colour on the sides of the body pale brownish grey in most specimens or pale yellowish brown; well-marked rosettes with reddish brown border and orangish brown interior forming small oblique bands on the sides of body; tail with reddish brown rings present from the base to tip.

Geographical distribution: Along and on both slopes of the Andes from Ecuador to north-western Argentina (Provinces of Catamarca and Córdoba) and northern Chile (Tarapacá Region) ( Fig. 9; Supporting Information, Appendix S3, Fig. S8). Ruíz-García et al. (2013) suggested that this species possibly occurs marginally in Colombia, based on a specimen collected in the Parque Natural Nacional Volcano Galeras, Department of Nariño, south-western Colombia, in August 1989 (IAVH5857). However, the specimen identified by them as a pampas cat is, in fact, a northern tigrina (Le. tigrinus ) (see: Nascimento & Feijó, 2017). Nevertheless, it is likely that Le. garleppi also occurs in the Andean region of Colombia, as predicted in our niche model ( Fig. 7B). Future studies in this areas may confirm this hypothesis.

Taxonomic notes: Depending on the author, the Garlepp’s pampas cat was regarded as either a subspecies of L. colocola ( Cabrera, 1940, 1958; Pearson, 1951; Ximénez, 1961; Pecon-Slattery et al., 1994, 2000, 2004; Masuda et al., 1996; Johnson & O’Brien, 1997; Eizirik et al., 1998; Pecon-Slattery & O’Brien, 1998; Johnson et al., 1999; Cossios et al., 2007; Napolitano et al., 2008; Cossíos et al., 2009; Sunquist & Sunquist, 2009; Li et al., 2016; Kitchener et al., 2017) or of L. pajeros ( Matschie, 1912; Lönnberg, 1913; Allen, 1919; Pocock, 1941; García-Perea, 1994; Wozencraft, 2005).

Mastchie (1912) described a specimen from Peru as Felis (Lynchailurus) pajeros garleppi and, a year later, Lönnberg (1913) described in detail another individual from Ecuador with similar pelage characters and named it Felis pajeros thomasi . Lönnberg also examined the type of garleppi and albeit indicated a close relationship between both taxa, he pointed out some characters that would be sufficient to differentiate these two species. However, Pocock (1941), examining the material from Ecuador and Peru available at the Museum of Natural History in London, did not detect significant differences between the populations of these countries that would justify the recognition of two taxa and, therefore, treated thomasi as a junior synonym of garleppi . In addition, Pocock described two new subspecies, Lynchailurus pajeros budini (Salta, NW Argentina) and Ly. p. steinbachi (W Bolivia), which showed characters similar to garleppi . In the same year, Schwangart (1941) had independently reached the same conclusion in treating garleppi and thomasi as synonyms.

Cabrera (1957) described a new subspecies from the province of Salta, north-western Argentina, named by him as Felis colocolo crespoi , which shared similarities with thomasi in the pattern of coat markings, but they differed from each other in background colour, and crespoi mainly differed from budini and pajeros by longer fur during the summer. Cabrera (1958), in his taxonomic catalogue of South American mammals, recognized garleppi and thomasi as distinct subspecies and considered steinbachi as the junior synonym of the former, and recognized crespoi and budini as valid subspecies. Later, Cabrera (1961) commented that budini would be an intermediate geographic form between garleppi from southern Peru and south-western Bolivia and colocola from Chile, resembling the former more than the latter, and the difference between budini and garleppi did not seem so evident and it is possible that a comparative study with good sampling of specimens could prove budini to be a junior synonym of garleppi .

Decades later, García-Perea (1994) recognized garleppi , thomasi, steinbachi, crespoi and budini as subspecies of L. pajeros . She examined the populations from Ecuador (thomasi) (N = 2) and Peru ( garleppi ) (N = 7) and realized that the skull and pelage characters, as well as habitat preferences (highelevation steppes), are similar, but they only differ in body size, with garleppi being larger than thomasi. García-Perea also affirmed that budini and crespoi showed the same colour pattern, but they differ from each other in the colour shades of the markings, and they inhabit a transitional zone between the higher steppes and dry forests. Based on the similarities in the pelage characters and habitat preferences, García- Perea suggested that crespoi should be a synonym of budini, but due to the small sample size and the poor condition of the type specimen (confirmed by one of us, FON, who examined the specimen), she provisionally treated them as distinct subspecies.

Moreover, García-Perea (1994) recognized the populations in northern Chile as a new subspecies and named these Lynchailurus colocolo wolffsohni . According to García-Perea’s description, the skull characters are close to those found in Ly. c. colocolo , but her Ly. c. wolffsohni differed from the nominal subspecies mainly by the pelage pattern, which in turn was similar to the pattern of the Peruvian and Bolivian populations. However, our results indicate that wolffsohni is a junior synonym of Le. garleppi , in agreement with previous authors. Osgood (1943) noted that the northern Chilean specimen could not ‘be identified satisfactorily as to subspecies, but perhaps will prove to be nearer to garleppi than colocolo ’. Furthermore, Osgood pointed out that the position of the central Chilean populations (= colocola ) in relation to the northern ‘varieties’ ( garleppi , thomasi, budini and steinbachi) was uncertain. In addition, Pine et al. (1979), comparing specimens of colocola from Central Chile and garleppi from Peru with the specimen from Tarapacá, north Chile, concluded ‘that the form in northern Chile is F. c. garleppi ’. Consonantly, molecular data suggested that samples from northern Chile, Bolivia and Peru form a well-supported group ( Johnson, 1999; Napolitano et al., 2008; Cossíos et al., 2009; Ruiz et al., 2013; Santos et al., 2018). Therefore, contradicting García-Perea (1994), the pampas cats from the western Andean slope of Peru and northern Chile belong to the same taxon.

Based on our results and on previous studies, we conclude that all forms – budini, crespoi, garleppi , steinbachi, thomasi and wolffsohni – are synonyms and should be united in a single species, Le. garleppi . The colour and spot patterns are similar and geographically consistent. The variation in colour shades, fur length and the skull dimensions, pointed out by previous authors are, in fact, of an individual nature.

Remarks: Most of the specimens showed the overall ground colour of body pale brownish grey, but some were pale yellowish brown. We did not find melanistic specimens in museum specimens, but Giordano et al. (2012) reported a melanistic individual in south-west of Parque Nacional Yanachaga-Chemillén, Pusapno, Peru (10°43’26”S, 75°27’43”W).