The correct name of the Uruguayan Colocolo
7
ANARTIA
Publicación del Museo de Biología de la Universidad del Zulia
ISSN 1315-642X (impresa) / ISSN 2665-0347 (digital)
https://doi.org/10.5281/zenodo.16500289 / Anartia, 40 (junio 2025): 7-11
The correct name of the Uruguayan Colocolo is
Leopardus munoai (Ximénez, 1961) (Mammalia, Carnivora,
Felidae), not L. fasciatus (Larrañaga, 1923), and the designation
of a neotype for the latter is invalid
El nombre correcto del Colocolo Uruguayo es Leopardus munoai (Ximénez, 1961)
(Mammalia, Carnivora, Felidae), no L. fasciatus (Larrañaga, 1923), y la designación
de un neotipo para este último es inválida
Jesús Molinari
Departamento de Biología, Facultad de Ciencias, Universidad de Los Andes, Mérida 5101, Venezuela.
orcid.org/0000-0002-9393-5483
Correspondence: jmvault@gmail.com
(Received: 01-03-2025 / Accepted: 15-04-2025 / On line: 31-07-2025)
ABSTRACT
Members of the Pampas Cat species group are small felids specialized for life in open habitats. eir taxonomic history is
complex, and they were divided into ve species recently. In the case of one of the species, the Uruguayan Colocolo, there
is a recent controversy between authors who either argue that its valid name is Leopardus munoai (Ximénez, 1961), or that
it is L. fasciatus (Larrañaga, 1923). A revision and reinterpretation of the original sources for these names, namely Azara
(1801, 1802), Desmarest (1816), and Larrañaga (1923) conrms the correct name of the Uruguayan Colocolo to be L.
munoai. e neotype of L. fasciatus is invalid because: 1) as a nominal species, L. fasciatus is not a senior synonym of L.
munoai, as it was assumed by those who designated the neotype, but instead it is a junior synonym of the Southern Pampas
Cat, L. pajeros (Desmarest, 1816); and 2) the designation of the neotype involved errors typied as causes of invalidity in
Article 75 of the International Code of Zoological Nomenclature.
Key words: Argentina, Brazil, Félix de Azara, Pajero Cat, pampas, taxonomy, Uruguay, zoological nomenclature.
RESUMEN
Los miembros del grupo de especies Gato de las Pampas son pequeños félidos especializados para la vida en ambientes
abiertos. Su historia taxonómica es compleja y recientemente fueron divididos en cinco especies. En el caso de una de las es-
pecies, el Colocolo Uruguayo, hay una controversia reciente entre autores que sostienen que su nombre válido es Leopardus
munoai (Ximénez, 1961), o que es L. fasciatus (Larrañaga, 1923). Una revisión y reinterpretación de las fuentes originales
de estos nombres, a saber, Azara (1801, 1802), Desmarest (1816) y Larrañaga (1923), conrma que el nombre correcto del
Colocolo Uruguayo es L. munoai. El neotipo de L. fasciatus es inválido porque: 1) como especie nominal, L. fasciatus no
es un sinónimo más antiguo de L. munoai, como supusieron quienes designaron el neotipo, sino un sinónimo más reciente
del Gato de las Pampas del Sur, L. pajeros (Desmarest, 1816); y 2) la designación del neotipo involucró errores tipicados
como causales de no validez en el Artículo 75 del Código Internacional de Nomenclatura Zoológica.
Palabras clave: Argentina, Brasil, Félix de Azara, Gato Pajero, nomenclatura zoológica, pampas, taxonomía, Uruguay.
Molinari
8
INTRODUCTION
Within the genus Leopardus Gray, 1842 (Mammalia,
Carnivora, Felidae), the members of the Pampas Cat spe-
cies group are the most specialized for life in open habitats
(García-Perea 1994, Nascimento et al. 2021). Before the
mid 1990’s, they were deemed to represent a single species
(Kitchener et al. 2017). ey were divided into three spe-
cies by García-Perea (1994), and into ve by Nascimento
et al. (2021). Based on the latter authors, the current classi-
cation of the species group divides them into: 1) the Cen-
tral Chilean Colocolo, Leopardus colocola (Molina, 1782),
endemic to Chile; 2) the Southern Pampas Cat, or South-
ern Colocolo, L. pajeros (Desmarest, 1816), occurring in
Argentina (the northern limit is the Catamarca Province),
and southern Chile; 3) the Uruguayan Colocolo, L. mu-
noai (Ximénez, 1961), occurring in Uruguay, Argentina
(Corrientes Province), and Brazil (Rio Grande do Sul);
4) the Pantanal Cat, L. braccatus (Cope, 1889), occur-
ring in northern Argentina (Formosa Province), Paraguay,
Bolivia (Beni Department), and Brazil (Mato Grosso and
Piauí States); and 5) the Northern Colocolo, L. garleppi
(Matschie, 1912), occurring in northern Argentina (Cata-
marca and Córdoba Provinces), northern Chile (Tarapacá
Region), Bolivia, Peru, Ecuador, and southwestern Co-
lombia (Nascimento et al. 2021, Astorquiza et al. 2023,
Distel et al. 2023, ASM 2024, Cabrera-Ojeda & Meléndez
2024). e species group is divided into two clades that
diverged 0.54 Myr ago: one of them includes L. colocola, L.
pajeros and L. garleppi; the other includes L. munoai and
L. braccatus (Nascimento et al. 2021).
e oldest binomial, Felis pajeros, applied to the South-
ern Pampas Cat (Desmarest 1816) was based on the earlier
morphological description under the vernacular names
‘Chat pampa’ (Azara 1801: 179–184) and ‘Gato pajero’
(Number XVIII; Azara 1802: 160–167). In the last de-
cades, the species has been referred to as Lynchailurus pa-
jeros (García-Perea 1994) and as Leopardus colocola pajeros
(Kitchener et al. 2017); though it is currently known as
Leopardus pajeros (for a detailed synonymy, see Nasci-
mento et al. 2021). Desmarest’s (1816) description was as
follows: ‘Twenty-third Species.— e P of d’Azara;
Felis pajeros, Nob., is also an animal om Paraguay (not yet
gured), the size of the wild cat, whose hair is long, so and
gray-brown above, with reddish transverse bands under the
throat and belly, and dark rings on the legs. e Pampa cat,
by the same author, does not seem to dier om it’1. Because
no holotype was ever designated for the species, and be-
cause a name-bearing type was deemed necessary to dene
the nominal taxon objectively, Nascimento et al. (2021)
selected a neotype for L. pajeros.
e Uruguayan Colocolo, Leopardus munoai, originally
deemed identical to the ‘Gato pajero’ (Azara 1801, 1802),
was described as Felis colocola muñoai. Its subspecic name
was emended to F. c. munoai by Ximénez (1970). It was
transferred to the genus Leopardus by Wozencra (2005),
and elevated to species by Nascimento et al. (2021).
RECOGNITION AND NEOTYPIFICATION OF
LEOPARDUS FASCIATUS
Dámaso Antonio Larrañaga (Montevideo, 1771–
1848) was contemporaneous with Félix de Azara (Bar-
buñales, 1742–1821) and Anselme Gaëtan Desmarest
(Paris, 1784–1838). Some of his most important writings
(Larrañaga 1922, 1923, 1924) were rst published by the
Instituto Histórico y Geográco del Uruguay (IHGU).
Larrañaga summarily described hundreds of new species
of plants and animals. In 1818, Larrañaga (1922: 85) was
aware of Desmarest’s description of Felis pajeros, to which
he referred to as ‘Felis Pajeros ( facciatus ) Dicc.’ [Dicc. =
Diccionarie, a misspelling of Dictionnaire]. Neverthe-
less, also based on Azara’s (1802) Number XVIII cat, he
devised his own binomial, which through the agency of
the IHGU was promulgated as Felis fasciatus Larrañaga,
1923. Ximénez et al. (1972) deemed F. fasciatus a junior
synonym of F. colocola pajeros. However, the Uruguayan
form was referred to as Felis colocola fasciatus by Klap-
penbach (1997), as Lynchailurus braccatus fasciatus by
González (2001), and as Leopardus braccatus fasciatus by
González & Martínez-Lanfranco (2010). Nascimento et
al. (2021) treated Felis fasciatus as a junior synonym of
Leopardus pajeros, while noting that González & Mar-
tínez-Lanfranco (2010) used fasciatus in place of munoai
without justication and ignoring the discussion of Xi-
ménez et al. (1970).
Based on their interpretation of Azara’s (1801, 1802)
and Larrañaga’s (1923) writings (the relevant passages are
quoted below), Martínez-Lanfranco & González (2022)
doubled down on their argument that the correct name
for the Uruguayan Colocolo is Leopardus fasciatus instead
of L. munoai. ey even designated a neotype for L. fas-
1 ‘Vingt-troisième Espèce.— Le P de d’Azara; Felis pajeros, Nob., est aussi un animal du Paraguay (non encore guré), de la taille du chat sauvage,
dont le poil est long, doux et gris-brun en dessus, avec desbandes transverses roussâtres sous la gorge et le ventre, et des anneaux obscurs sur les pattes. Le
chat Pampa, du même auteur, ne paroît pas en diérer’ (Desmarest 1816).
The correct name of the Uruguayan Colocolo
9
ciatus, with a type locality near the Río Negro of Uruguay,
~100 km to the northeast of that of L. munoai. Following
Martínez-Lanfranco & González (2022), the latest ver-
sion of the Mammal Diversity Database of the American
Society of Mammalogists (ASM 2024) accepts L. fasciatus
as the valid name of the Uruguayan Colocolo.
Azara (1801: 179) stated that: ‘I do not know, and I
have not heard that it lives in Paraguay. It is found in the
Pampas, south of Buenos Aires, in the grasslands between the
55th and 36th degrees of latitude’ 2. Azara (1802: 160) ex-
panded these remarks as follows: ‘NÚM. XVIII. ON THE
PAJERO. ey call it Pajero cat because it lives in the elds,
hiding in the grasslands without entering the forests and
bushes, where the previous ones live. I do not know, nor have
I heard that it exists in Paraguay; and it could be, because
having a sucient population and fewer elds, they might
have been able to exterminate it. I caught four in the Pampas
of Buenos Aires between the 35 and 36 degrees, and another
three where the Black [donde al Negro]’ 3 [bold mine].
Azara (1802: 161) also stated that the species was found
‘on both sides of the La Plata River with identity of shapes,
colors and habits’ 4, hence he clearly was collectively refer-
ring to populations at present classied (Nascimento et
al. 2021) as Leopardus pajeros (Argentina) and L. munoai
(Uruguay).
Larrañaga (1923: 345) described L. fasciatus as follows:
‘Sp. 5. F(elis). fasciatus—tail elongate immaculate, woolly,
body above dilutely brown, below white with cinnamon-
banded feet. Sp. n. HABITAT more common than the previ-
ous one, does not enter forests: length 34⅓, tail 11¾. Azara
Number XVIII Pajero’ 5. Larrañaga (2023) made no other
mention or comment about L. fasciatus other than listing
the name in the indexes of the book, and in two tables
(inserted between pages 340 and 341) entitled ‘Classica-
tion of the mammals of the La Plata River, particularly its
Eastern Bank according to the system of Cuvier, with their
characters adapted to the country’ 6 and ‘Classication of the
mammals of this country according to the system of Cuvier’ 7.
MISINTERPRETATION OF AZARA (1801, 1802)
AND LARRAÑAGA (1923)
Martínez-Lanfranco & González (2022) did two
things. First, they argued Leopardus fasciatus (Larrañaga,
1923), and not L. munoai (Ximénez, 1961), to be the
oldest name available for the Uruguayan Colocolo. In
support of this claim, they stated that Larrañaga (1923)
‘clearly distinguished’ between Uruguayan and Argentin-
ian specimens (Premise 1), and that he ‘was explicit’ in that
his description of F. fasciatus was based solely on Uruguay-
an specimens (Premise 2). Second, ‘to anchor Larrañaga’s
fasciatus to Uruguayan pampas cats’, they designated a
neotype for this taxon; to justify this action, they inferred
that, because Azara (1802) mentioned the Pajero cat to in-
habit both sides of the La Plata River, his ‘donde al Negro’
[‘where the Black’] was intended to mean that this felid oc-
curs in the vicinity of the Río Negro of Uruguay (Premise
3); accordingly, for the neotype, they chose a type locality
just to the south of this river. Unfortunately, Martínez-
Lanfranco & González (2022) went beyond the mean-
ing and scope of the statements (transcribed in full in the
preceding section) of Azara (1801, 1802) and Larrañaga
(1923) on the Pajero cat, as the examination that follows
of what I am referring to as their premises reveal.
Premise 1) Larrañaga (1923) ‘clearly distinguished’
between Uruguayan and Argentinian specimens.—It is
hard to see how by means of a description containing only
14 words5, and without a diagnosis or comparisons, two
closely related, variable and similar felids could be ‘clearly
distinguished’; and there is no reason to assume that the
use of such words5 indicates that Larrañaga knew and in-
tended to say that the two felids represent dierent species
or subspecies. Martínez-Lanfranco & González (2022)
claimed that ‘in the context of Larrañaga’s diagnosis of the
new species, fasciatus refers to the specimens as having cin-
namon-colored banded feet. Azara (1802:162), in contrast,
mentioned that the “Pajero” had lightly cinnamon-colored
2 ‘JE ne sache point, et je n’ai pas ouï dire qu’il habite le Paraguay. On le rencontre dans les Pampas, au Sud de Buenos-Ayres, lieu où, entre les pajonals du
55.e et du 36.e degré de latitude’ (Azara 1801).
3 ‘NÚM. XVIII. DEL PAJERO. Le llaman Gato pajero, porque habita los campos, escondiéndose en los pajonales sin entrar en los bosques y matorrales,
donde habitan los precedentes. No sé, ni he oído que exista en el Paragüay ; y podrá ser, porque teniendo bastante poblacion y ménos campos, le habrán
podido exterminar. Yo pillé quatro en las Pampas de Buenos Ayres entre los 35 y 36 grados, y otros tres donde al Negro’ (Azara 1802).
4 ‘en ámbas bandas del Rio de la Plata con identidad de formas, colores y costumbres’ (Azara 1802).
5 ‘Sp. 5.a. F. fasciatus—cauda elongata inmaculata, lanosa, corpore supra dilute fusco, ina albido cum pedibus cinnamomo-fasciato. Sp. n. H com-
munior precedenti, nemora non ingreditur : longitudine 34⅓, cauda 11¾. Azara N.° XVIII Pajero’ (Larrañaga1923).
6 ‘Clasicación de los mamilares del Río de la Plata, particularmente de su Banda Oriental según el sistema de Cuvier, con sus caracteres acomodados al
país’ (Larrañaga 1923).
7 ‘Clasicación de los mamilares de este país según el sistema de Cuvier’ (Larrañaga 1923).
Molinari
10
limbs without bands’. However, this contraposition is not
supported by the authors’ words: Larrañaga (1923) wrote
‘with cinnamon-banded feet’5; whereas Azara (1802: 162)
wrote ‘e forelimbs and hindlimbs are whitish on the in-
side, and cinnamon-white on the outside; but they also have
dark transverse stripes or rings across them that are dark and
very evident’ 8.
Premise 2) Larrañaga (1923) ‘was explicit’ in that his
description of F. fasciatus was based solely on Uruguay-
an specimens.—e argument of Martínez-Lanfranco
& González (2022) is as follows: ‘Larrañaga (1923), in
turn, was explicit in that the taxa he was enumerating and
describing, e. g., Felis fasciatus, were om Uruguay and not
generically om the Río de la Plata (which could have also
included Argentina; Larrañaga 1923:341–342)’. At least
in my copy of Larrañaga’s (1923) book, there is nothing
between pages 340 and 342, other than two tables6,7 listing
mammals occurring in Uruguay, including native, intro-
duced and domestic species. erefore, not only it is not
explicit, but it is not even implicit that any of these mam-
mals occurs in Uruguay and not in Argentina. Most im-
portantly, both Desmarest (1816) and Larrañaga (1923)
described their species based on Azara’s Number XVIII cat
(said to inhabit both sides of the La Plata River) as a whole
and not divided into parts, thus their binomials originally
were applied to both Argentinian and Uruguayan popula-
tions.
Premise 3) rough the expression ‘donde al Negro’, A z-
ara (1802: 160) intended to mean that his ‘Gato pajero’ oc-
curs in the vicinity of the Río Negro of Uruguay.—In his
works, when referring to rivers, Azara typically used the
word ‘Rio’ before de name of rivers, e.g., Rio de la Plata, rio
Negro, rio Paragüay, rio Paraná, rio Uruguay. In Spanish,
the expression ‘donde el’ (of which Azara’s ‘donde al’ is a
rare variant, or perhaps a typographical error) followed by
the name of a living being (usually a person, but also an an-
imal or a plant) is oen used to refer to a place belonging
to such being, or where such being is. For these reasons, it
is likely that when Azara wrote ‘donde al Negro’ what he
intended to say is that he saw his Pajero cat in the same
place where he also saw his ‘Gato Negro’ (almost certainly
melanistic individuals of another species of Leopardus), to
which he also referred as ‘(d)el Negro’ (‘NÚM XV. DEL
NEGRO’; Azara 1802: 154). Based on Azara’s account, he
saw his ‘el Negro’ in Rio Grande do Sul, 450–500 km to
the east of the type locality that Martínez-Lanfranco &
González’s (2022) chose for their neotype.
CONCLUSIONS
Martínez-Lanfranco & González (2022) incorrectly as-
sumed that, when describing L. fasciatus, Larrañaga (1923)
distinguished between Uruguayan and Argentinian speci-
mens, and that he based this taxon solely on Uruguayan
specimens. In fact, like Desmarest’s (1816) description of
L. pajeros, that of L. fasciatus was based on Azara’s (1801,
1802) account of the ‘Chat pampa’ or ‘Gato pajero’, which
the latter author conceptualized as occurring on both sides
of the La Plata River. erefore, contrary to the claims of
Martínez-Lanfranco & González (2022), and as previous-
ly concluded by Ximénez et al. (1972) and Nascimento et
al. (2021), L. fasciatus (Larrañaga, 1923) should be con-
sidered a junior synonym of L. pajeros (Desmarest, 1816),
and not a senior synonym of L. munoai (Ximénez, 1961).
Whereas synonymizing and reviving synonyms in the
manner regulated by the International Code of Zoological
Nomenclature (ICZN 1999) (herein Code) are common-
place in taxonomy, neotypication is not (ICZN 1999:
Article 75, Yanega et al. 2018). is is so because the des-
ignation of a neotype can create redundancies with other
type material, mismatches between the nominal taxa and
the natural populations represented by the neotype, and
erroneous type localities. is is exemplied by the neo-
type of L. fasciatus: it is redundant with the holotype of
L. munoai; it stands for a nominal species occurring in Ar-
gentina and for natural populations occurring in Uruguay;
and it has a type locality based on a misinterpretation of
Azara’s (1802) and Larrañaga’s (1923) writings.
e designation of a neotype by Martínez-Lanfranco
& González (2022) is invalid and problematic because L.
fasciatus is invalid. In addition, the Code denes certain
precise conditions under which a neotype shall not be des-
ignated. ose that applied to the neotype of L. fasciatus
are: 1) ‘A neotype is not to be designated as an end in itself,
or as a matter of curatorial routine, and any such neotype
designation is invalid’ (Article 75.2); 2) ‘A neotype is val-
idly designated when there is an exceptional need and only
when that need is stated expressly and when the designation
is published with the following particulars: 75.3.1. a state-
ment that it is designated with the express purpose of clari-
fying the taxonomic status or the type locality of a nominal
taxon; 75.3.6. evidence that the neotype came as nearly as
practicable om the original type locality [Art. 76.1]’ (Ar-
ticle 75.3). uite clearly, Martínez-Lanfranco & González
(2022) designated a neotype for L. fasciatus in the absence
8 ‘Los brazos y piernas por dentro blanquecinos, y por fuera blancos acanelados ; pero ademas tienen zunchos ó anillos al través obscuros muy reparables’
(Azara 1802).
The correct name of the Uruguayan Colocolo
11
of an exceptional need, and without proper support for
their choice of a type locality.
ACKNOWLEDGMENTS
I am grateful to Aaron M. Bauer and Ángel L. Viloria
for their comments on the manuscript.
REFERENCES
ASM [American Society of Mammalogists]. 2024. Mammal
diversity database, version v1.13. https://doi.org/10.5281/
zenodo.10595931
Astorquiza, J. M., E. A. Noguera-Urbano, C. Cabrera-Ojeda,
B. Cepeda-uilindo, J. F. González-Maya, E. Eizirik, A. Bo-
nilla-Sánchez, D. L. Buitrago, P. Pulido-Santacruz & H. E.
Ramírez-Chaves. 2023. Distribution of the northern pampas
cat, Leopardus garleppi, in northern South America, conr-
mation of its presence in Colombia and genetic analysis of a
controversial record from the country. Mammalia 87: 606–
614. https://doi.org/10.1515/mammalia-2022-01
Azara, F. de. 1801. Essais sur l’Histoire naturelle des quadrupèdes
de la Province du Paraguay. Tome Premier. Paris: Charles
Pougens, lxxx + 366 pp.
Azara, F. de. 1802. Apuntamientos para la historia natural de los
quadrúpedos del Paragüay y Rio de la Plata. Tomo Primero.
Madrid: Imprenta de la Viuda de Ibarra, xix + 318 pp.
Cabrera-Ojeda, C. & J. Meléndez. 2024. Nuevo registro del
gato de las pampas de Garlepp (Leopardus garleppi, Matschie,
1912) en el valle del Patía, Colombia. Mammalogy Notes 10:
452. https://doi.org/10.47603/mano.v10n2.452
Desmarest, A. G. 1816. CHAT, Felis, Linn., Briss., Schreb., Cuv.
pp. 73–123. In: Nouveau dictionnaire d’Histoire naturelle.
Tome VI. Paris: Deterville.
Distel, A., M. S. Di Bitetti, S. Cirignoli, Y. E. Di Blanco & J.
A. Pereira. 2023. e last stronghold of Muñoa’s Pampas
cat (Leopardus munoai) in Argentina? Journal for Na-
ture Conservation 74: 126449. https://doi.org/10.1016/j.
jnc.2023.126449
García-Perea, R. 1994. e Pampas cat group (genus Lynchailu-
rus Severtzov, 1858) (Carnivora, Felidae), a systematic and
biogeographic review. American Museum Novitates 3096:
1–36.
González, E. M. 2001. Guía de campo de los mamíferos de Uru-
guay: introducción al estudio de los mamíferos. Montevideo:
Vida Silvestre Uruguay, 340 pp.
González, E. M. & J. A. Martínez-Lanfranco. 2010. Mamíferos
de Uruguay: guía de campo e introducción a su estudio y
conservación. Montevideo: Ediciones de la Banda Oriental,
464 pp.
ICZN [International Commission on Zoological Nomencla-
ture]. 1999. International Code of Zoological Nomenclature,
Fourth Edition. London, UK: International Trust for Zoo-
logical Nomenclature, xxix + 126 pp.
Kitchener, A. C., Ch. Breitenmoser-Würsten, E. Eizirik, A. Gen-
try, L. Werdelin, A. Wilting, N. Yamaguchi, A. V. Abramov,
P. Christiansen, C. Driscoll, J. W. Duckworth, W. E. John-
son, S.-J. Luo, E. Meijaard, P. O’Donoghue, J. Sanderson, K.
Seymour, M. Bruford, C. Groves, M. Homann, K. Nowell,
Z. Timmons & S. Tobe. 2017. A revised taxonomy of the Fe-
lidae. e nal report of the Cat Classication Task Force
of the IUCN/SSC Cat Specialist Group. Cat News Special
Issue 11: 1–80.
Klappenbach, M. A. 1997. Larrañaga naturalista. Algunos
aspectos poco conocidos de su obra. Revista del Instituto
Histórico y Geográco del Uruguay 27: 287–304.
Larrañaga, D. A. 1922. Escritos de Don Dámaso Antonio Larra-
ñaga. Tomo I. Montevideo: Instituto Histórico y Geográco
del Uruguay, xxiii + 439 pp.
Larrañaga, D. A. 1923. Escritos de Don Dámaso Antonio Larra-
ñaga. Tomo II. Montevideo: Instituto Histórico y Geográco
del Uruguay, 512 pp.
Larrañaga, D. A. 1924. Escritos de Don Dámaso Antonio Lar-
rañaga. Tomo III. Montevideo: Instituto Histórico y Geográ-
co del Uruguay, xix + 306 pp.
Martínez-Lanfranco, J. A. & E. M. González. 2022. e oldest
available name for the pampas cat of the Uruguayan Savannah
ecoregion is Leopardus fasciatus (Larrañaga 1923). erya
13: 259–264. https://doi.org/10.12933/therya-22-1187
Nascimento, F. O., J. Cheng & A. Feijó. 2021. Taxonomic revi-
sion of the Pampas Cat Leopardus colocola complex (Carniv-
ora: Felidae): An integrative approach. Zoological Journal of
the Linnean Society 191: 575–611. https://doi.org/10.1093/
zoolinnean/zlaa043
Wozencra, W. C. 2005. Order Carnivora. pp. 532–628. In:
Wilson, D. E. & D. M., Reeder (eds). Mammal species of the
world, 3rd Edition. Baltimore, USA: Johns Hopkins Univer-
sity Press.
Ximénez, A. 1970. Notas sobre félidos neotropicales I: Felis
colocola braccata y sus relaciones com Felis colocola munoai y
Felis colocola pajeros. Comunicaciones Zoológicas del Museo de
Historia Natural de Montevideo 10: 1–6.
Ximénez, A., A. Langguth & R. Praderi. 1972. Lista sistemática
de los mamíferos del Uruguay. Anales del Museo Nacional de
Historia Natural de Montevideo 7: 1–49.
Yanega, D., T. Pape, F. Welter-Schultes, J.-I. Kojima, N. L. Even-
huis, F.-T. Krell, M. J. Grygier, S. T. Ahyong, A. Ballerio, P.
Bouchard, F. E. Rheindt, D. A. Dmitriev, M. S. Harvey, G. La-
mas, R. L. Pyle, B. Halliday & Z.-Q. Zhang. 2018. When zo-
ological type specimens are lost: ICZN-compliant guidelines
for when and when not to designate neotypes Version 12-
ix-2018. http://dx.doi.org/10.13140/RG.2.2.17568.51201
