Leopardus tigrinus ( Schreber, 1775 )

Nascimento, Fabio Oliveira do & Feijó, Anderson, 2017, Taxonomic revision of the tigrina Leopardus tigrinus (Schreber, 1775) species group (Carnivora, Felidae), Papéis Avulsos de Zoologia 57 (19), pp. 231-264 : 243-247

Leopardus tigrinus ( Schreber, 1775)

Northern tigrina

Felis tigrina Schreber, 1775 : plate CVI [based on the plate XXXVII, “Le Margay ”, Buffon (1765)]; 1777:396 (text). Type locality “südlichen Amerika” (= ”South America”), subsequently restricted to “Cayenne” by J.A. Allen (1919:356).

Felis margay Müller, 1776:29 . Based on “Le Margay ” Buffon (1765).

Felis (Oncoides) tigrina : Severtzov, 1858:386 (name combination).

Felis pardinoides Gray, 1867a:400 . Type locality “ India ”, subsequently redefined to “Bogotá” by Gray (1874:475).

Felis geoffroyi : Elliot, 1872:203 (part) (non d’Orbigny & Gervais, 1844).

Felis tigrina : Thomas, 1880:396 (non Felis tigrina Schreber, 1775 ).

Felis pardinoides andina Thomas, 1903:238 . Type locality “ Jima , Province of Azuay, Ecuador ”.

Felis pardinoides emerita Thomas, 1912:44 . Type locality “Montes de la Cutala, Merida, Venezuela ”.

Margay tigrina emerita : Allen, 1915:631 (name combination).

Margay tigrina elenae : Allen, 1915:631 (name combination).

Margay caucensis Allen, 1915:631 . Type locality “ Las Pavas , Colombia ”.

Margay tigrina wiedi : Allen, 1916:233 (part) (non Felis wiedii Schinz, 1821 ).

Margay tigrina andina : Allen, 1916:581 (name combination).

Margay tigrina tigrina : Allen, 1919:350 (name combination).

Oncilla pardinoides emerita: Allen, 1919:359 (name combination).

Oncilla pardinoides elenae: Allen, 1919:360 (name combination).

Oncilla caucencis: Allen, 1919:360 (name combination).

Leopardus tigrinus : Pocock, 1941a:237 (name combination, first use of current name combination).

Felis (Leopardus) tigrina pardinoides : Cabrera, 1958:286 (name combination).

Felis (Leopardus) tigrina tigrina : Cabrera, 1958:287 (part) (name combination).

Oncifelis tigrina : Wozencraft, 1993:290 (part).

Leopardus tigrinus tigrinus : Wozencraft, 2005:539 (part) (name combination).

Leopardus tigrinus pardinoides : Wozencraft, 2005:539 (name combination).

Type locality: “südlichen Amerika” [= “South America”], restricted by J.A. Allen (1919:356) to “Cayenne” [ French Guiana]: “Based on Buffon’s description and figure of a specimen from Cayenne” [“Le Margay (pl. XXXVII) qui a servi de sujet pour cette description, ayant été tué à Cayenne (…)” ( Buffon, 1765:252)] ( Husson, 1978; Wozencraft, 2005).

Type material: Schreber (1775, 1777) based his Felis tigrina on “Le Margay ” of Buffon (1765) (plate XXXVII) ( Fig. 7A ). The plate of Schreber’s F. tigrina ( Fig. 7B ) was published in 1775 [as plate CVI (= 106)] and the text with its description in 1777. Husson (1978) stated that the animal figured in the Schreber’s plate is the holotype of the species, but Thomas (1903:235) had previously designated the Buffon’s as the type. Thus, following the opinion of Thomas (1903) and in accordance with the ICZN Code (articles 74.4 and 74.6) (1999), we recognize the Buffon’s plate as the lectotype of L. tigrinus .

Diagnosis: Small sized; fur relatively harsh; ground color dark brown and orangish brown to yellowish brown and grayish brown, becoming lighter on the sides of the body; venter white or light gray; medium-sized rosettes on the sides of the body form small and/ or medium-sized oblique bands arranged in scapular-inguinal direction.

Body measurements: See Morphogroup I in Table 2.

Geographical distribution: The map of the Fig. 8 shows the points where the museum specimens were collected, which include the northern Brazil (State of Amapá; left bank of the Amazon River), Guyanas, Venezuela, Colombia, Ecuador, Peru, northwestern Argentina and Costa Rica. Leopardus tigrinus probably occurs in Bolivia, but there is no museum record (see Anderson, 1997:334). A specimen ( MNK 4595; not MNK 3730) from Puerto Limón, Santa Cruz, Bolivia, was identified as a L. tigrinus by Huáscar Azurduy (2005), but it is in fact a specimen of L. wiedii (it exhibits directed backwards hair on the nape and overall softer fur). There is a gap in the geographic distribution between Central American and northern South American population ( Figs. 2 , 4 and 8 ), but it is unknown whether the absence in this gap is natural or a sampling artifact. The distribution of L. tigrinus is mainly associated to lowland, premontane and montane forests ( Eisenberg, 1989; Garrido & González-Maya, 2011), from sea level up to 3,000 -3,200 m ( Mondolfi, 1986; Sunquist & Sunquist, 2002, 2009), but some individuals have been recorded as high as 4,500 -4,800 m ( Melquist, 1984; Cuervo et al., 1986; Nowell & Jackson, 1996; Macdonald et al., 2010). Apparently it has a marginal distribution in the open areas of the Llanos ( Fig. 4 : Morphogroup I, Fig. 8 ).

Variation: L. tigrinus does not show sexual dimorphism for external characters. The overall ground color of the head and body varies from dark brown to light yellowish brown orange, with the sides of the body usually lighter (buff colored) towards the venter ( Fig. 9 ). The rosettes and the small and medium-sized oblique bands on the sides of body have black or very dark brown rims with most specimens the color inside the rosettes and bands is similar to that show in the dorsum, which may be brownish orange or dark brown. These patterns of ground color and markings in L. tigrinus are very similar those found in L. wiedii ( Fig. 10 ). It likely may lead to a misidentification of these species in the field, especially in a quick observation. However, these two species can be distinguished from each other by a series of characters, in special the direction of the hairs on nape, which are facing back in L. tigrinus and forward in L. wiedii . Furthermore, some individuals of L. tigrinus (for example: AMNH 14187, from Pavas, Valle del Cauca, Colombia; AMNH 149319, from El Tambo, Cauca, Colombia; and USNM 362126, from Guyana) have a darker color inside the rosettes and bands, almost similar to the rims that surround them, a pattern very similar to that observed in some specimens of L. wiedii (Nascimento, pers. obs.). Furthermore, melanistic specimens are present in L. tigrinus (for example, USNM 371278, from Caracas, Venezuela).

Taxonomic notes: Despite some authors ( Erxleben, 1777; Gmelin, 1788; F. Cuvier, 1826; Temminck, 1827; Fischer, 1829; Jardine, 1834; Gray, 1867a, 1867b, 1869) in 18 th and 19 th century followed Schreber’s F. tigrina , other authors associated this name with the forms related to wiedii Schinz, 1821 (and in a lesser extent to pardalis Linnaeus, 1758 and guigna Molina, 1782 ) due to their morphologic similarities among the specimens studied by them, creating doubts regarding the exact identity of the animals that bear these names (see Elliot, 1877, 1883; Allen, 1919). Besides tigrina Schreber, 1775 , the names pardinoides Gray, 1867a , guttula Hensel, 1872 , pardinoides Thomas, 1903 and oncilla Allen, 1904 are commonly considered as subspecies of L. tigrinus ( Pocock, 1941a; Cabrera, 1958; Wozencraft, 2005). According to our results, the putative subspecies L. t. pardinoides ( Gray, 1867a) does not differ significantly from L. t. tigrinus , and so we recognize the former as the junior synonym of the latter, rather than its subspecies or a distinct taxon. In addition, specimens from the northeastern Brazil are traditionally considered to belong to tigrinus Schreber, 1775 (e.g., Cabrera, 1958), but our results indicate that they belong to a distinct taxon, L. emiliae ( Thomas, 1914) .

Regarding to the taxonomy of Central American population, Thomas (1903:237) described a specimen from Volcan de Irazu, Costa Rica, as Felis pardinoides oncilla , which showed size and general characters as in F. pardinoides Gray, 1867a , but with ground color of pelage “much richer and deeper”, and the lateral rosettes little elongated. According to Gardner (1971), the specimens from Central America resemble the type of Margay caucensis J.A. Allen, 1915 (from Las Pavas, Rio Cauca, Colombia) [which was synonymized in Felis (Leopardus) tigrina pardinoides by Cabrera (1958)] in the intensity of markings and richness of color. Nevertheless, due the apparent absence of the tigrina between southern Panama and northern South America ( Figs. 2 , 4 and 8 ), Gardner recognized the Central American population as distinct from the northern South American population (= O. t. pardinoides ). According to the molecular data, the Central American population of tigrinas is highly genetically divergent from the southern South American tigrina, a divergence higher than that between L. pardalis and L. wiedii and O. geoffroyi and O. guigna ( Johnson et al., 1999) . However, samples from other populations, mainly from northwestern South America (for example, from Colombia and Venezuela), were not included in the molecular study.

Figures

FIGURE 2: Distribution of the specimens of tigrinas (L. tigrinus group) studied. Black dots correspond to the specimens from scientific collections.

FIGURE 4: Distribution of morphogroups of tigrinas (L. tigrinus group) over the biomes of South America and part of Central America (data obtained from WWF – World Wide Fund for Nature). For color figure, see online version.

FIGURE 7: Reproduction of the original plates: (A): “Le Margay” of Buffon (1765) (obtained from Gallica – Bibliothèque Nationale de France), the lectotype of the species; and (B): Felis tigrina of Schreber (1775). For color figure, see online version.

FIGURE 8: Geographic distribution of the specimens of L. tigrinus studied. Numbers correspond to collection localities listed in the gazetteer (see Appendix II) and the star refers to the type locality. For color figure, see online version.

FIGURE 9: Pattern of coloration and markings in L. tigrinus: (A): EBRG9609 (Raul Leoni, Bolivar, Venezuela); (B): USNM374861 (Los Patos, El Manteco, Bolivar, Venezuela); AMNH69116 (Rio Chili, Manizales, Loreto, Colombia); (C): NMW-B455 (Carrillo, S des Llano de Santa Clara, San Jose, Costa Rica). Bar = 100 mm. For color figure, see online version.

FIGURE 10: Comparison of the patterns of ground color and markings in (A): northern tigrina (L. tigrinus) (USNM374861, Los Patos, El Manteco, Bolivar, Venezuela) and (B): margay (L. wiedii) (NMNH388255, Rio Cunucunuma, Belen, Amazonas, Venezuela). Bar = 100 mm. For color figure, see online version.