Leopardus geoffroyi (d’Orbigny & Gervais, 1844)

Nascimento, Fabio Oliveira do, 2014, On the morphological variation and taxonomy of the Geoffroy's cat Leopardus geoffroyi (d'Orbigny & amp; Gervais, 1844) (Carnivora, Felidae), Papéis Avulsos de Zoologia (São Paulo) 54 (11), pp. 129-160 : 141-151

Leopardus geoffroyi (d’Orbigny & Gervais, 1844)

Geoffroy’s cat

Leopardus himalayanus Gray, 1843: 44 (non Jardine, 1834) (nomem nudum).

Felis geoffroyi d’Orbigny & Gervais, 1844a: 40 ; 1844b: plate LVII; 1847: 21, plate XIII, plate XIV. Type locality: “des rives Du Rio Negro, Patagonie”.

Felis (Oncifelis) geoffroyi : Severtzov, 1858: 386 (name combination).

Pardalina warnickii : Gray, 1867a: 267; plate 24.

Felis pardoides Gray, 1867b: 403 (non Felis pardoides Owen, 1846 ) (preoccupied name).

Felis guigna : Mivart, 1881: 410 (non Felis guigna Molina, 1782 ) (preoccupied name).

Felis (Oncoides) geoffroyi : Lahille, 1899: 178 (name combination).

Felis salinarum Thomas, 1903: 239 .

Oncoides geoffroyi : Allen, 1905: 180 (name combination).

Felis melas Bertoni, 1914 (non Cuvier, 1809) (preoccupied name).

Herpailurus geoffroyi : Pocock, 1917: 347 (name combination).

Oncifelis geoffroyi Allen, 1919: 366 , figs. 5a, 7b, 13a and 23.

Oncifelis salinarum Allen, 1919: 367 (name combination).

Felis geoffroyi mac-donaldi Marelli, 1932: 38 .

Oncifelis geoffroyi paraguae Pocock, 1940: 351 .

Oncifelis geoffroyi leucobaptus Pocock, 1940: 351 .

Oncifelis geoffroyi euxanthus Pocock, 1940: 352 .

Oncifelis geoffroyi argentea Schwangart, 1941: 16 .

Oncifelis geoffroyi flava Schwangart, 1941: 16 .

Felis (Leopardus) geoffroyi geoffroyi : Cabrera, 1958: 280 (name combination).

Felis (Leopardus) geoffroyi leucobapta : Cabrera, 1958: 280 (name combination).

Felis (Leopardus) geoffroyi paraguae : Cabrera, 1958: 280 (name combination).

Felis (Leopardus) geoffroyi salinarum : Cabrera, 1958: 281 (name combination).

Felis (Leopardus) geoffroyi geoffroyi : Ximénez, 1975: 1 (name combination).

Felis (Leopardus) geoffroyi salinarum : Ximénez, 1975: 1 (name combination).

Felis (Leopardus) geoffroyi paraguae : Ximénez, 1975: 1 (name combination).

Felis (Leopardus) geoffroyi euxantha : Ximénez, 1975: 1 (name combination).

Felis colocolo geoffroyi : Hershkovitz, 1987: 73, fig. 19 (name combination; probable misspelling).

Leopardus geoffroyi geoffroyi : Wozencraft, 2005: 538 (name combination).

Leopardus geoffroyi euxanthus : Wozencraft, 2005: 538 (name combination).

Leopardus geoffroyi leucobaptus : Wozencraft, 2005: 538 (name combination).

Leopardus geoffroyi paraguae : Wozencraft, 2005: 538 (name combination).

Leopardus geoffroyi salinarum : Wozencraft, 2005: 538 (name combination).

Type locality: “des rives Du Rio Negro, Patagonie” (= riverbanks of Rio Negro, Buenos Aires Province, Argentina) ( d’Orbigny & Gervais, 1844a). “Elle habite les Pampas de Buenos-Ayres jusqu’au 44°. degree de latitude sud” ( d’Orbigny, 1847).

Type material: d’Orbigny & Gervais (1844a) described the species based on three specimens collected by Alcides d’Orbigny in the lower Río Negro, at the southern end of the Province of Buenos Aires, that are deposited in the collection of the Museum National d’Histoire Naturelle, Paris, France (“Trois exemplaires recueillis par M. d’Orbigny sont depuis longtemps exposés dans les galleries du Muséum”), under the numbers MNHN-ZM-MO-2001-298 (= “number 90”) (mounted skin, skull in the skin; “Rives du Rio Negro, Patagonie, Argentine ”; February, 1831), MNHN-ZM-MO-2001-299 (= “number 92”) (mounted skin, skull in the skin; “Rives du Rio Negro, Patagonie, Argentine ”; February, 1831; individual featured in “l’Atlas mammalogique du voyage de M. d’Orbigny, plate XIV”) and MNHN-ZM-MO-2001-300 (= “number 91”) (mounted skin, skull extracted; “ Buenos Aires, Argentine ”; July, 1829) ( Figs. 1b, 1d and 2 ). However, d’Orbigny & Gervais (1844a, b) did not indicate which specimens was indeed the holotype ( Schwangart, 1941; Cabrera, 1961; Ximénez, 1971, 1975). From information provided to Schwangart (1941, page 15) by Dr. P. Rhodes ( MNHN), this author stated that specimen number 92 was the holotype and both this one and specimen number 90 were from Patagonia, each showing yellowish white ground col- or, while specimen number 91 came from [the Province of] Buenos Aires and had a deep yellow ground color. Cabrera (1961) commented, based on the information from Dr. Jean Dorst ( MNHN), that one of the three specimens of the type series is labeled “un des types”, and he concluded that this specimen should then be the lectotype, but Cabrera did not indicate which one. However, as can be seen in Fig. 1 , present on the labels of all three specimens (which, by the way, are very old), is the term “type” contrasting with the information provided by Dorst. Later, Dr. Francis Petter ( MNHN) reported in Ximénez (1971, 1975) that the holotype would be specimen number 91 which has “Republique Argentine ” written on its label, while specimens numbers 90 and 92 had “Patagonie” on the respective labels. Furthermore, in the publication of Ximénez (1971), there is a picture of one of these specimens (page 68, fig. 1) with the word “type” written on its label. Ximénez informed that this specimen was set for exhibition, and it would be impossible to determine whether the original color was indeed clear or the lighter coloration was due to prolonged exposure to light. This specimen represented in the Ximénez article is the same showed here in Fig. 1c . In comparison to Fig. 1a , it can be concluded that this specimen – MNHN-ZM-MO-2001-298 (= “number 90”) – was used as the basis for this plate, which was originally published in 1844 (plate 57; in d’Orbigny & Gervais, 1844b). Furthermore, this plate is the first formal representation of the species. In view of this, and contrary to what previous authors have suggested, I conclude that this specimen – MNHN-ZM-MO-2001-298 – is actually the lectotype of L. geoffroyi . Consequently, the other two specimens (MNHN-ZM-MO-2001-299 and MNHN-ZM-MO-2001-300) are the paralectotypes of the species.

Classification Resultsb,c Subspecies Original Count geoffroyi paraguae euxanthus salinarum leucobaptus % geoffroyi paraguae euxanthus salinarum leucobaptus Cross-validated a Count geoffroyi paraguae euxanthus salinarum leucobaptus % geoffroyi paraguae euxanthus salinarum leucobaptus

Predicted Group Membership geoffroyi paraguae euxanthus salinarum leucobaptus Total 2 0 0 0 0 2 2 13 0 0 6 21 1 0 0 1 0 2 0 0 0 1 0 1 0 0 0 0 1 1 100.0 0.0 0.0 0.0 0.0 100.0 9.5 61.9 0.0 0.0 28.6 100.0 50.0 0.0 0.0 50.0 0.0 100.0 0.0 0.0 0.0 100.0 0.0 100.0 0.0 0.0 0.0 0.0 100.0 100.0 2 0 0 0 0 2 2 12 0 0 7 21 1 0 0 1 0 2 0 0 1 0 0 1 0 1 0 0 0 1 100.0 0.0 0.0 0.0 0.0 100.0 9.5 57.1 0.0 0.0 33.3 100.0 50.0 0.0 0.0 50.0 0.0 100.0 0.0 0.0 100.0 0.0 0.0 100.0 0.0 100.0 0.0 0.0 0.0 100.0 the analysis. In cross validation, each case is classified by the functions derived from all. classified. females obtained by discriminant function analysis concerning the probabilities of one of the five subspecific taxa. Classification Resultsb,c Predicted Group Membership

Subspecies geoffroyi paraguae euxanthus salinarum leucobaptus Total Original Count geoffroyi 0 3 4 0 0 7 paraguae 3 18 5 2 2 30 euxanthus 0 0 1 0 0 1 salinarum 0 0 0 1 1 2 leucobaptus 0 0 0 0 1 1 % geoffroyi 0.0 42.9 57.1 0.0 0.0 100.0 paraguae 10.0 60.0 16.7 6.7 6.7 100.0 euxanthus 0.0 0.0 100.0 0.0 0.0 100.0 salinarum 0.0 0.0 0.0 50.0 50.0 100.0 leucobaptus 0.0 0.0 0.0 0.0 100.0 100.0 Cross-validated a Count geoffroyi 0 3 4 0 0 7 paraguae 4 15 6 2 3 30 euxanthus 1 0 0 0 0 1 salinarum 0 0 0 1 1 2 leucobaptus 0 0 0 1 0 1 % geoffroyi 0.0 42.9 57.1 0.0 0.0 100.0 paraguae 13.3 50.0 20.0 6.7 10.0 100.0 euxanthus 100.0 0.0 0.0 0.0 0.0 100.0 salinarum 0.0 0.0 0.0 50.0 50.0 100.0 leucobaptus 0.0 0.0 0.0 100.0 0.0 100.0

Diagnosis: Small (2-5 kg); ground color of pelage varying from light yellowish brown to smoky gray; pattern of markings on the body predominantly composed of medium and/or small solid black spots; melanistic specimens may be present; long ringed tail; short and robust skull with well-developed zygomatic arches; profile of skull slightly convex in the frontal region; short and high sagittal crest; notch in the posterior margin of the palate may be present, sometimes with two small lateral projections.

Description: Small sized cat, length of head and body between 498 and 750 mm (601.58 ± 59.91 mm; n = 12) in males and between 390 and 515 mm (456.67± 62.92 mm;n= 3)in females,tail length between 300 and 410 mm (348.83 ± 27.36 mm; n = 12) in males and between 265 and 275 mm (271.67 ± 5.77 mm; n = 3) in females, hind foot length is between 115 and 150 mm (132.27 ± 10.56 mm; n = 11) in males and between 100 and 216 mm (140.33 ± 65.58 mm; n = 3) in females, and the body mass between 3590 and 5900 g (5205 ± 949.08 g; n = 6) in males and between 2000 and 3000 g (2500 ± 707.11 g; n = 2) in females. Rounded ears measuring between 45 and 62 mm (53.73 ± 4.67 mm; n = 11) in males and between 47 and 57 mm (51.33 ± 5.13 mm; n = 3) in females.

Overall coloration of the head varies from light yellowish brown to smoky gray, except, chin, cheeks, throat, and around the lips and eyes, which are white or very light gray. Two genal black or very dark brown stripes cross in parallel the cheeks in longitudinal direction. In the region of the posterior end of these stripes there is a stripe like an incomplete transversal necklace (“anterior transverse stripe or hyoid stripe”). The upper genal stripe connects with the black stripe around the eyes, the dark orbital stripe. In the mystacial region, there are four or five rows of spots that may or may not coalesce, and with the exception of the top two rows, the remaining ones extend to philtrum. The supraorbital spots or narrow stripes are arranged longitudinally on each side of the head and can connect to form frontal-parietal strips. Between these two stripes there are numerous, small, rounded or elliptical spots. Usually five longitudinal stripes run along the nape and the lateral region of the neck to the anterior part of the dorsum in the interscapular region. The hairs on head and neck are short and slightly harsh, and on nape they are pointing backward. Ears rounded with dorsal surface almost entirely black except at the base, which has the same color of the rest of the head, and a white spot is centrally disposed on the dark surface of the dorsal surface of the ear. The body shows the same overall coloration found in the head and neck, especially in the dorsum and the dorsal surface of the fore and hind limbs. In the dorsum, rounded and/or elliptical spots may be separated or interconnected forming more or less longitudinal rows. The spot pattern in the interscapular region shows high individual variation. On the side of the body, small and/or medium-sized solid rounded and elliptical black spots are present, generally not forming rosettes or oblique bands, but a few individuals may exhibit open and incomplete rosettes. The venter is white or very light gray and has small and medium-sized rounded very dark spots. The hairs on the body are short and slightly harsh, but slightly longer than the head, and in the inguinal region the hairs are longer than the rest of the body. Melanistic specimens are also present. The dorsal surface of limbs has the same color pattern of the dorsum of the body, while the ventral surface has the same pattern of the venter. On the dorsal surface of the fore and hind limbs are present round and/or elliptical spots of medium size in the proximal region, while only spots of smaller size are present in the distal region on the feet. On the ventral surface of the fore and hind limbs are present medium and small sized spots.The hairs on the feet are short and slightly harsh. The tail is relatively long, corresponding to between 52 and 63% of the length of the head and body, and displays black or very dark brown rings alternating with rings of the same color of the dorsum, and the tip is dark colored.

Robust skull with broad and short rostrum, corresponding to between 32.29 and 37.99% ( RL /GLS: 35.09 ± 1.41%; n = 35) of the greatest length of skull in males and between 29.71 and 37.06% ( RL /GLS: 34.49 ± 1.49%; n = 49) in females. The nasals are broad distally, and narrow shortly thereafter to converge at the posterior end, where they articulate with the frontal; there may or may not be a depression in this region. The anterior margin of the nasals is predominantly curved or moderately curved ( Table 4). The anterior ends of the pre-maxillae are not projected and thus, in side view, they are aligned with the anterior end of the nasals. When the skull is in dorsal view, the nasals fully cover the incisive foramina. The orbits are large, rounded and forward-faced, being positioned entirely in the anterior half of the skull. The anteriormost margin of the orbit is aligned at the P3 parastyle, while the posteriormost point of the margin of the orbit coincides with the alignment of the end of the post-orbital process of the jugal. The upper and lower postorbital processes are not connected and, therefore, they do not form a complete and fused postorbital bar. The zygomatic plate, which is part of the maxilla, is well developed and forms the floor of the orbital region. The frontal is well developed and extends from the maxilla-frontal suture and nasal-frontal to the anterior portion of the braincase, articulating with the parietal. In lateral view, the skull has a slightly convex profile in the frontal region, providing a less evident curvature. The interorbital region is narrow and its width in proportion to the greatest length of skull shows values between 15.47 and 24.20% ( IOB /GLS: 18.52 ± 1.69; n = 35) ( Table 3) in males and between 15.78 and 20.33% ( IOB /GLS: 18.26 ± 1.05; n = 50) ( Table 3) in females. The braincase is large and oval, with the proportion of its width in relation to greatest length of skull varying between 39.44 and 47.51% ( GBB /GLS: 43.01 ± 2.47; n = 34) ( Table 3) in males and between 41.69 and 50.01% ( GBB /GLS: 46.21 ± 1.83; n = 50) ( Table 3) in females. The sagittal crest is present in most specimens, with most males showing type 2 (poorly developed and restricted to the region interparietal) (50.00%; n = 17) and type 3 (moderately developed) (35.29%, n = 12) ( Table 4), while most females have type 2 (82.35%; n = 42) ( Table 4). Only a small fraction of sampled specimens have the sagittal crest well developed; and they are all males (8.83%, n = 3) ( Table 4) and one specimen of unknown sex. Overall, the sagittal crests are well developed and moderately high. Temporal lines are present and from the lyriform type. The lambdoidal crest may be present and poorly, moderately or well developed. The palate is relatively large and its width between P3 varies from 32.84 to 42.53 mm ( GPB: 39.34 ± 2.07; n = 36) in males and 30.69 to 41.27 mm ( GPB: 36.64 ± 1.92; n = 54) in females ( Table 3). In most of cases, the length of palate is equal or less than the width of palate [GPL/ GPB: 101.24 ± 5.12 (n = 35) in males; 99.07 ± 4.57 (n = 53) in females ( Table 3)]. The notch of the postpalatine vein is broad and comparatively shallow in most specimens (males: 85.29%, n = 29; females: 75.00%, n = 39) ( Table 4) and the posterior margin of the palate (or anterior margin of mesopterygoid fossa) has a U-shaped edge. It may or may not have a medial notch, which can be shallow or deep ( Table 4). Furthermore, the posterior margin of the palate in L. geoffroyi distinguishes from other small and medium-sized Neotropical felids in two ways. Firstly, it is related to the shape of the margin when the notch is absent, in other words, this area of postpalatine may have a regular and uniform margin or may have a projection towards the mesopterygoid fossa, in the same line of the left and right palatine suture. Another point is when the notch is present. In addition to the aforementioned typical shape (deep or shallow), another shape can be found, which is the presence of two lateral projections to the central notch, one on each side, and it is represented in 23.58% of the overall sample (29 in 123 individuals), independent of the sex or geographic group. The presphenoid, centrally located in the mesopterygoid fossa, is narrow, very elongated and arranged longitudinally, showing lateral expansions in the median area. The basioccipital, located between auditory bullae, is usually narrow. The mastoid processes are arranged in a posterolateral position in relation to the auditory bullae and they are anteriorly articulated to the paraoccipital processes. The mastoid processes shape in 96.97% of males (n = 32) and 83.76% of females (n = 41) ( Table 4) is posteriorly poorly developed, separated from paraoccipital processes by a notch, enabling the visualization of the surface of the auditory bulla. The zygomatic arches are more expanded laterally, and the average width of the braincase relative to the zygomatic width ( GBB /ZB) is around 65% in males (n = 34) and 69% in females (n = 51) ( Table 3). The occipital condyle is elongated, robust and spirally curved and encloses the foramen magnum, which is well developed. The auditory bulla is relatively large and oval, with ectotympanic smaller than entotympanic in 93.94% of males (n = 31) and 86.0% of females (n = 43) ( Table 4). The mandible is well developed, the horizontal ramus is high and curved, especially in the anterior region, and the masseteric fossa is deep and broad, extending almost the entire ascending ramus. The ascending ramus is high and extends from the angular process to the outermost end of the coronoid process. The coronoid process is well developed, can be broad or narrow, rounded and curved, resembling a hook in lateral view. The condyloid process is robust, bar shaped, aligned transversely to the ascending ramus, and on the same occlusional plane of the lower tooth row. The angular process is relatively large and rounded, which can be aligned to or positioned a little posteriorly to the condyloid process. The C-M1 length varies from 28.66 to 34.97 mm ( CM1 L: 31.96 ± 1.71 mm; n = 37) in males and from 25.86 to 33.78 mm ( CM1 L: 29.37 ± 1.38 mm; n = 53) in females ( Table 3), while the p3-m1 length varies from 20.48 to 25.04 mm (p 3m 1L: 23.22 ± 1.19 mm; n = 35) in males and from 19.08 to 24.64 mm (p 3m 1L: 21.62 ± 1.11 mm; n = 31) in females ( Table 3). The shape of P3 paracone may be narrow and long [males: 75.86% (n = 22); females: 63.83% (n = 30)] or short and wide [males: 24.14% (n = 7); females: 36.17% (n = 17)] ( Table 4), and P3 parastyle is absent in all specimens, except for three unknown sexed specimens ( Table 4). The P 4 paracone is present in almost all studied specimens [males: 100% (n = 31); females: 96.15% (n = 50) ( Table 4)]. Traces of a talonid on m1 are absent in most of the sample [males: 83.87% (n = 26); females: 91.67% (n = 44) ( Table 4)] .

Geographical distribution: Leopardus geoffroyi is found in a wide variety of temperate and subtropical habitat types from sea level to up to 3,300 m of altitude, associated to open vegetational formations, which includes the pampas grasslands, dry Chaco shrub and woodlands from southern Bolivia and northwestern Argentina, Paraguay, southern Brazil, Uruguay and south through Argentina and bordering areas of southern Chile to Patagonia and the Strait of Magellan ( Figs. 4 and 5 ) ( Ximénez, 1975; Redford & Eisenberg, 1992; Nowell & Jackson, 1996; Sunquist & Sunquist, 2002, 2009; Lucherini et al. 2006).

Geographic and local variation: Leopardus geoffroyi does not show sexual dimorphism on external morphology, but does for some craniodental characters. The size, shape and arrangement of spots, as well as the pattern of ground color, varies greatly between specimens from the same locality, resulting in a large individual variation in pelage. Some individuals show spots well separated and distinct from each other, while other specimens have solid spots very close together, making them almost indistinguishable and generating a speckled pattern. In some cases, the aggregation of spots generally occurs along the axis of the back, leaving some individuals with a pattern similar to a stripe. All these patterns are individual and are found widely in different geographic groups.

The ground color, which is the main criterion for defining the putative subspecies, shows high individual variation ( Fig. 13 ). In a same population – for example in Uruguay – it is possible to find different specimens exhibiting color characters that define all subspecies (e.g., ochraceous ground color and dark gray ground color). Ximénez (1971) reported that individuals from different locations (and by extension from different subspecies) showed similar ground color patterns. This author noted that specimens of Salta ( MACN36.619 and MACN36.229 ) and southern Brazil are no different from those of Uruguay, a pattern that I also observed. Some southern individuals (e.g., from Neuquén, Rio Negro, La Pampa) are slightly lighter than their northern counterparts, with a slight tendency to gray, but in La Pampa there is a specimen with the “typical” color ( MACN51.168 ) and other more light grayish ( MACN22025 ). In Pozo de Maza , Formosa, in the northern Argentina, a specimen ( MACN47.404 ) shows a grayish color. Specimens from Jujuy are usually also lighter, very subtly, but one specimen ( MACN34.556 ) from Villa Unión (La Rioja) has a dark ochraceous ground color. An individual, without known locality, identified as “leucobapta” ( MACN31.248 ), has a more yellowish color than the other groups described above. Furthermore , in the six specimen series (all collected in July 1936) from Santiago de Estero , Lavalle, Argentina, some specimens ( AMNH41551 and AMNH41555 ) exhibit colors tending to gray, resembling the specimen from Cañadon de las Vacas, Corpen Aike, Santa Cruz ( AMNH16696 ); while other specimens ( AMNH41550 , 41552 , 41553 and 41554) have a coloration similar to the predominant color in the specimens of Uruguay (light yellowish brown to dark yellowish brown) .

Regarding the craniodental characters, the sagittal crest is highly variable in individual and geographical terms, but the males usually have longer and higher sagittal crests (type III, moderately developed; no specimens shows the type IV, the well-developed crest) and broader, more robust zygomatic arches than are observed in females. Furthermore, regardless of sex, L. geoffroyi shows characters of the posterior margin of the palate that differ from other Neotropical felid species. Individuals who do not possess a notch in the postpalatine may have the regular margin, or there may be a projection toward the fossa mesopterygoid at the height of the suture that articulates left and right palatine, similar to what is observed in specimens of domestic cat, Felis catus Linnaeus, 1758 . Some specimens who have the notch may also exhibit two projections lateral to it, one on each side. Aside from L. geoffroyi , I just noticed this character on one specimen of L. wiedii from Buena Vista, Bolivia ( AMNH 61790). There are no significant differences in cranial dimensions in the studied groups along the latitudinal distribution of the species.

Comparisions: Here I present the differences that distinguish L. geoffroyi from other small and medium sized felids found throughout its geographical distribution.

Leopardus geoffroyi is distinguished from Leopardus pardalis (Linnaeus, 1758) by the smaller body size, premaxilla not projected forward and holdings its anterior end aligned to the anterior end of the nasal, hairs on the nape facing backward, small and/or medium-sized solid rounded and elliptical black spots distributed almost all over the body; while L. pardalis shows medium body size, rosettes that can coalesce and form oblique bands arranged in scapular-inguinal direction, hairs on the nape facing forward, larger and more robust skull, premaxillary projected anteriorly in relation to the anterior end of the nasal, the region of the frontal slightly convex in profile, sagittal crest well developed and often high, lambdoidal crests well developed and robust and well developed zygomatic arches.

Leopardus geoffroyi is distinguished from Leopardus wiedii (Schinz, 1821) by the larger body size of the former, premaxilla not projected forward and holding its anterior end aligned to the anterior end of the nasal, hairs on the nape facing backward, small and/or medium-sized solid rounded and elliptical black spots distributed almost all over the body, whereas L. wiedii shows small size, presence of small rosettes on the sides of the body which usually coalesce and form small oblique bands arranged in scapular-inguinal direction, the tail proportionately longer, hairs on the nape facing forward, smaller and more gracile skull, the region of the frontal slightly convex, premaxilla projected forward in relation to the anterior end of the nasal, more rounded braincase, sagittal crest usually absent and narrow zygomatic arches, delicate.

Leopardus geoffroyi is distinguished from Leopardus guttulus by a the larger body size, larger and more robust skull, small to moderately developed sagittal crest, small and/or medium-sized solid rounded and elliptical black spots distributed almost all over the body, while L. guttulus has a smaller body size, presence of small distinctive rosettes on the body, smaller and more gracile skull, sagittal crest (when present) undeveloped and restricted to the interparietal region, and narrow and delicate zygomatic arches.

Leopardus geoffroyi is distinguished from Leopardus guigna by larger body size, light yellowish brown to smoky gray ground color of body, larger and more robust skull, small or moderately developed sagittal crest; while L. guigna has a smaller body size, grayish fawn, reddish brown or dark reddish brown ground color, a quite large and conspicuous lateral rostral stripe, presence of a third less extensive stripe running roughly parallel and between the upper and lower genal stripes, an even thicher tail, smaller and less robust skull, the presence of “internal temporal lines” that are located between each parietal suture and actual temporal lines (or “external temporal lines”), parietal slightly higher in the region between the internal temporal lines.

Leopardus geoffroyi is distinguished from the pampas cat group ( L. pajeros and L. braccatus ) by presenting more rounded ears with a white spot centrally disposed on the dark surface of its dorsal surface, small and/or medium-sized solid rounded and elliptical black spots distributed almost all over the body and tail with very dark rings, while the pampas cat group shows more triangular ears with hair tuft at the tip, spinal crest little darker than ground color, and legs with transverse stripes present in the proximal portion. The Northern forms of Leopardus pajeros (Desmarest, 1816) (which includes the putative Bolivian and northern Argentinean subspecies budini Pocock, 1941, crespoi Cabrera, 1957 and steinbachi Pocock, 1941) are distinguished from L. geoffroyi by a yellowish gray or grayish brown ground color with the flanks covered by rusty “ocelotlike” rosettes forming oblique bands. The Southern forms of L. pajeros (Desmarest, 1816) (which includes the putative subspecies pajeros Desmarest, 1816 and crucina Thomas, 1901) differ from L. geoffroyi by almost uniformly grayish brown ground color with faint and almost imperceptible oblique lines on the flanks, tail not ringed. L. braccatus braccatus (Cope, 1889) is distinguished from L. geoffroyi by its uniform brown agouti ground color, the presence of fainted traces of dark brown rosettes in the flanks, the black stripes in the proximal area of the legs and the feet all black. L. braccatus munoai differs from L. geoffroyi by uniform yellowish brown agouti ground color, traces of dark brown rosettes in the flanks, the black stripes in the proximal area of the legs, the feet black only in the ventral (= palmar and plantar) surfaces, tail with discontinuous rings or not ringed.

Leopardus geoffroyi is distinguished from Leopardus jacobita (Cornalia, 1865) by the spotted pattern of the pelage, the dorsal profile of skull being convex, and the ectotympanic chamber smaller than the entotympanic, while L. jacobita has spots that tend to be in vertical lines on the body, bushy and long ringed tail, the dorsal profile of skull is more flat and elongated, and the ectotympanic chamber is equal to or larger than the entotympanic.

Leopardus geoffroyi is distinguished from Puma yagouaroundi (É. Geoffroy, 1803) by the spotted pattern of the pelage, ringed tail, presence of dark lines across the cheeks, a fossa shallow along the posterior internasal and anterior interfrontal sagittal sutures, and broad auditory bullae, whereas P. yagouaroundi has a uniform coloration (gray, yellow, red, brown or black), not ringed tail, absence of dark lines across the cheeks, a deep fossa along the posterior internasal and anterior interfrontal sagittal sutures, and narrower auditory bullae.

Leopardus geoffroyi is distinguished from Felis catus by the spotted pattern of the pelage, the rounded ears without tufts in the tip, the posterior margin of palate with or without a medial notch, a small projection in the margin of the palate towards the mesopterygoid fossa may be present, while F. catus has a great individual variation in color pattern, but never consists of black spots on yellowish or orange background ( Anderson, 1997), a triangular ear with small tufts in the tip, the zygomatic plate proportionally larger, the upper postorbital processes are large and almost reaching the lower postorbital processes, the posterior margin of the palate without a medial notch and a small projection in the margin of palate towards the mesopterygoid fossa is always present.

Figures

FIGURE 1: Reproduction of the original plates and syntypes of Leopardus geoffroyi (d’Orbigny & Gervais, 1844b, 1847): (A) overall view of the species (plate published in 1844); (B) overall view of the species (plate published in 1847); (C) specimen MNHN-ZM-MO-2001-298; (D) specimen MNHN-ZM-MO-2001-299; and (E) specimen MNHN-ZM-MO-2001-300.Photos of specimens by Cécile Callou (MNHN).

FIGURE 2: (A) lateral and ventral views of the skull (plate published in 1847); (B) lateral view from the skull of one of the syntypes, the specimen MNHN-ZM-MO-2001-300; and (C) dorsal view from the skull of one of the syntypes, the specimen MNHN-ZM-MO-2001-300. Photo of specimens by Cécile Callou (MNHN).

FIGURE 4: Distribution of the specimens of L. geoffroyi studied. Numbers correspond to collection localities listed in the gazetteer (see Appendix 2). Gray shadow corresponds to the geographic range of the species suggested by IUCN.

FIGURE 5: Distribution of the putative subspecies of L. geoffroyi over the biomes of southern South America (data obtained from WWF – World Wide Fund for Nature). The stars represent the type localities of the putative subspecies.

FIGURE 13: Pattern of coloration and markings in L. geoffroyi: (A) MACN34.556 (unknown sex) from Villa Unión, La Rioja, Argentina; (B) AMNH39010 (male) from Pulqui, Oropeza, Chuquisaca, Bolivia; (C) MACN14590 (female) from Estación Romero, El Cuy, Río Negro, Argentina; (D) MACN47.404 (unknown sex) from Pozo de Maza, Formoza, Argentina; (E) AMNH41555 (unknown sex) from Lavalle, Santiago del Estero, Argentina; (F) AMNH41550 (unknown sex) from Lavalle, Santiago del Estero, Argentina. Bar = 100 mm.