Journal of Mammalogy, 83(1):110–124, 2002

ANDEAN MOUNTAIN CAT, OREAILURUS JACOBITA:

MORPHOLOGICAL DESCRIPTION AND COMPARISON WITH
OTHER FELINES FROM THE ALTIPLANO

ROSA GARCı́A-PEREA*

Museo Nacional de Ciencias Naturales, CL. J. Gutierrez Abascal 2, Madrid 28006, Spain

Recent field surveys searching for the rare Andean mountain cat, Oreailurus jacobita, have
had difficulty in identifying this species from sightings and skins. This is caused by the
paucity of museum specimens (only 3 skulls available to date) and the lack of criteria to
differentiate this species from other small sympatric felines (e.g., Lynchailurus pajeros). In
a study to solve these problems, 3 new skulls of O. jacobita were identified and a total of
5 skulls and 41 skins examined. Skulls of O. jacobita average 12–14% larger than skulls
of L. pajeros, showing a large anterior chamber in the bulla. Andean mountain cats have
vertical series of yellowish-brown blotches on the sides, and a long, bushy tail with wide
dark rings. Two keys to differentiate these felines, based on these and other characters, are
offered.

Key words: Andean altiplano, Andean mountain cat, feral cat, Lynchailurus pajeros, morphological
keys, Oreailurus jacobita, pampas cat, South America

Two species of small wild cat occur in
and around the Andean altiplano of Peru,
Bolivia, Chile, and Argentina: Andean
mountain cat (Oreailurus jacobita Cornalia,
1865) and pampas cat (Lynchailurus pajer-
os (Desmarest, 1816)). The 1st species is a
poorly known felid, whose morphology and
biometry have been poorly described on the
basis of 3 skulls (Kuhn 1973; Pearson
1957; Philippi 1870), 14 skins (Burmeister
1879; Cabrera 1961; Cornalia 1865; Greer
1965; Matschie 1912; Philippi 1870, 1873;
Pine et al. 1979; Pocock 1941; Scrocchi and
Halloy 1986; Yepes 1929), and photographs
of 3 individuals in their natural habitat
(Sanderson 1999; Scrocchi and Halloy
1986; Ziesler 1992). Moreover, the 3 skulls
were subadult specimens, and 1 of them
was lost at the beginning of the 20th cen-
tury. Based on such a small sample, no at-
tempt at a detailed description of the mor-
phology and biometry of this species has
* Correspondent: mcng310@mncn.csic.es
been made. Also, age and sex variations are
unknown.
The pampas cat has traditionally been
considered a unique, polymorphic species
(Lynchailurus pajeros, after Allen 1919, or
Oncifelis colocolo, after Wozencraft 1993),
although it has recently been split into 3
species (Garcı́a-Perea 1994): Lynchailurus
pajeros, L. colocolo, and L. braccatus. The
amount and nature of morphological vari-
ation within this species group have been
described in some detail by Garcı́a-Perea
(1994).
No museum specimens of the pampas cat
from the altiplano were found by Garcı́a-
Perea (1994), but recent reports indicate
that altiplano farmers keep skins of both
kinds of cat in their homes (A. Iriarte, L.
Villalba, and N. Bernal, in litt.). Although
recent molecular studies (Johnson et al.
1998) suggest that Oreailurus and Lyn-
chailurus are not closely related genera,
specimens of both genera can be confused
in altiplanic areas because of the external

110

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 February 2002 GARCÍA-PEREA—MORPHOLOGY OF ANDEAN MOUNTAIN CATS
similarity in their coat patterns. Native
farmers even give them the same name in
many localities (L. Villalba and N. Bernal,
in litt.). Field researchers have found it dif-
ficult to distinguish the 2 species in sight-
ings, skins stuffed by native people, or
skulls found occasionally while conducting
fieldwork. For example, some of the Boli-
vian records of O. jacobita included in Yen-
sen et al. (1994) belong to L. pajeros (see
‘‘Conclusions’’). This situation is caused by
the paucity of museum specimens of O. ja-
cobita available for comparative purposes
and the lack of published information pro-
viding field researchers with effective tools
to identify specimens.
Awareness of these difficulties led me to
conduct a study focused on providing a key
for differentiating small wild cats living in
the area. The goals of my study were char-
acterization of external and skull morphol-
ogies of O. jacobita and elaboration of a
key for differentiating altiplanic individuals
of the Andean mountain cat and the pampas
cat. I have included, wherever possible,
some comparisons with the domestic cat
(Felis catus), because feral specimens of
the domestic form of the African wild cat
(Felis silvestris lybica group, after Nowell
and Jackson 1996) seem to be widespread
in the altiplano and other remote places of
South America. However, the high amount
of variation in skull and pelage character-
istics of these feral specimens makes it im-
possible to treat these differences in depth
now. A recent revision of the information
available on O. jacobita, based on litera-
ture, has been released (Yensen and Sey-
mour 2000). In this article, I am providing
firsthand data from the largest sample ever
studied of the rare Andean mountain cat.
MATERIALS AND METHODS
A sample of 44 specimens (3 skulls, 39 skins,
and 2 skulls plus skins) of O. jacobita, 50 spec-
imens (29 skulls and 38 skins) of L. pajeros, and
16 specimens (15 skulls and 6 skins) of F. catus
kept in 14 American and European institutions
was studied (Appendix I). Specimens were from
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111
FIG. 1.—A) Dorsal view of subadult skull of
Oreailurus jacobita (based on MVZ 116317)
showing an unossified interparietal bone (IB)
and a developing sagittal crest (SC). B) Ventral
view of a juvenile skull of Lynchailurus pajeros
(based on CBF 2960) showing upper milk den-
tition (dC, dP3, dP4) and unossified sphenooc-
cipital synchondrosis (SS).
Peru, Bolivia, Chile, and Argentina. Specimens
were identified to species a priori by skull and
skin characters, based on the information pro-
vided by the few descriptive papers available
(e.g., Cabrera 1961; Garcı́a-Perea 1994) and my
own data. Only specimens of pampas cats from
the altiplano and nearby areas were considered,
in order to remove geographic variation from the
sample (Garcı́a-Perea 1994).
For morphological characterization of O. ja-
cobita and for purposes of comparison among
species, a relative age was assigned to each in-
dividual. Age was estimated on skulls based on
the developmental stages of several morpholog-
ical structures (Barone 1976; Garcı́a-Perea 1991,
1996; Garcı́a-Perea et al. 1996; Gaunt 1959). Ju-
veniles (1st year) can be identified as those hav-
ing deciduous dentition either erupting, fully
erupted, or being replaced by permanent teeth
(Fig. 1B), and as those lacking sagittal crest or
showing only an incipient structure. Subadults
(2nd year) can be identified as those having per-
manent dentition, unossified sphenooccipital
synchondrosis, and interparietal bone, and a de-
veloping sagittal crest (Fig. 1A). Adults are
specimens more than 2 years old, showing to-
tally ossified sphenooccipital synchondrosis and
interparietal bone. The word adult is used here
with an ontogenetic meaning, indicating that
specimens have ended their growth and have
 112 JOURNAL OF MAMMALOGY
reached their definitive physical characteristics
(i.e., not related to reproductive condition).
Skulls were classified by age as follows: O. ja-
cobita, 2 subadults and 3 adults; L. pajeros, 5
juveniles, 5 subadults, and 19 adults; and F. ca-
tus, 15 adults.
No criteria have been established for estimat-
ing age from felid skins except relative size, to
use which we need to know already the size of
adult specimens, and this was not the case for
the Andean mountain cat. However, the large
sample (n 5 20) of O. jacobita skins studied
from Catamarca, Argentina, offered a unique op-
portunity to analyze age variation. Considering
the fact that these specimens came from a rela-
tively small area, I assumed that they represent-
ed a single population. All skins were preserved
with the same tanning method, so I assumed that
size differences I observed reflected real differ-
ences in size between individuals. Five external
measurements were recorded wherever avail-
able, either from the collection labels or from
measurement of specimens of O. jacobita: head
and body length, tail length, hind foot length,
ear length, and body weight.
Based on these premises, I used the variables
head and body length and tail length on the
skins, searching for differences in global size
that could be attributed to age. A comparison of
values obtained allowed me to establish 3 size
classes. Because males are likely larger than fe-
males, as in most felids (Garcı́a-Perea 1994,
1996, in litt.; Garcı́a-Perea et al. 1985), some
differences in size could be attributed to sexual
dimorphism. However, the 3 size groups identi-
fied showed noticeable differences in spotting
pattern and color, something unusual among
males and females of the same species of wild
felid but linked to growth in many species such
as the lion, the cougar, or the lynx (Ewer 1973;
R. Garcı́a-Perea, in litt.). I assumed therefore
that the 3 size groups represented 3 age classes,
covering age periods perhaps not strictly equiv-
alent to those described based on skull (values
of head and body length and tail length in mil-
limeters): juveniles (n 5 1, head and body length
5 500, tail length about 330); subadults (n 5 8,
head and body length 5 640–660, tail length 5
330–420); and adults (n 5 11, head and body
length 5 740–850, tail length 5 410–480).
Twenty skull and teeth variables commonly
used for carnivores were measured on each
specimen of O. jacobita and L. pajeros (Fig. 2),
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Vol. 83, No. 1
using a digital caliper accurate to 0.02 mm. De-
scriptive statistics (mean, standard error of
mean, range) were calculated for each variable.
Because I had only 2 subadult and 3 adult spec-
imens of O. jacobita, no statistical tests were
applied to test differences observed between the
2 age groups. However, significant differences
have been found between subadults and adults
of other felid species (Andersen and Wiig 1984;
Garcı́a-Perea et al. 1996); so differences among
species were tested only on adult specimens.
Thus, the sample used for that purpose consisted
of 3 O. jacobita, 19 L. pajeros, and 14 F. catus
(craniometric data of F. catus were recorded
from literature—Pocock 1951—because only
qualitative characters were available from the 15
skulls mentioned earlier). For dental measure-
ments, 5 skulls of O. jacobita were considered,
because carnassials erupt with their definitive di-
mensions. Differences among pairs of species
were tested by Mann–Whitney U-test and Stu-
dent’s t-test, using SPSS software (SPSS Inc.
1993).
Further, 119 morphological characters were
checked on each skull to record the character
states present in the sample, but only 10 of them
showed variation when specimens of Oreailurus
and Lynchailurus were compared (Figs. 3–5):
palate—position of anterior palatine foramina
relative to palatine-maxilla suture; rostrum—
shape of nasal bones; zygomatic arches—size of
dorsal postorbital process of jugal; basicran-
ium—shape of presphenoid body; bullae—po-
sition of vagina processus hyoideus relative to
stylomastoid foramen; bullae—shape of paroc-
cipital processes; teeth—1 ridge between 2
grooves in lingual side of upper canine; teeth—
size of parastyle related to protocone of P4;
teeth—metaconid on m1; and bullae—relative
size of ecto- and entotympanic chambers.
Seven characters related to body proportions,
external anatomy, and coat pattern were exam-
ined on each skin wherever possible (Figs. 6 and
7): legs—black, conspicuous rings around fore-
arms; ears—color of dorsal region; body—spi-
nal crest (band of long, erectile fur from shoul-
ders to base of tail); body—arrangement and
color of blotches or rosettes on sides; face—col-
or of rhinarium; tail—number, size, and color of
rings; and tail length—percentage of head and
body length represented.
A Spanish summary of the results is available
at the web site of Cat Action Treasury (CAT),
 February 2002 GARCÍA-PEREA—MORPHOLOGY OF ANDEAN MOUNTAIN CATS 113

FIG. 2.—Variables measured on each skull. P4, m1: upper and lower carnassials, respectively, in
occlusal views. GLS—greatest length of skull, CBL—condylobasal length, BAL—basilar axis length,
ZW—zygomatic width, PL—palatal length, FW—facial width (distance between tips of postorbital
processes), IOW—interorbital width, POW—postorbital width, CH—neurocranial height, RL—ros-
tral length, IBD—interbullae distance, MW—mastoid width, RWC—rostral width across the canines,
SCL—sagittal crest length, ML—mandible length, MRH—height of mandible ramus, P4L—length
of upper P4, P4W—maximum width of P4, Lm1—length of lower m1, Wm1—maximum width of m1.

www.felidae.org. Spanish translations of figure
legends and titles of tables are available from
the author.
RESULTS AND DISCUSSION
Age-related variation in skulls of O. ja-
cobita.—The youngest specimens of this
species studied were 2 subadults (MVZ
116317, UG D2337). Besides the small size
compared with adults (Table 1) and an un-
ossified sphenooccipital synchondrosis
(Fig. 1A), subadults have different skull
proportions: postorbital processes are less
developed, zygomatic arches are narrower,
and sagittal crest is shorter (i.e., lower val-
ues of facial width, zygomatic width, and
sagittal crest length). Also, postorbital con-
strictions of adults are narrower (lower val-
ues of postorbital width). These results are
consistent with those found for other felid
species, such as the Lynx (Andersen and
Wiig 1984; Garcı́a-Perea 1991) and other
carnivores (e.g., Genetta genetta—Craw-
ford-Cabral 1981).
These ontogenetic changes must be con-
sidered carefully, because several characters
described by Pocock (1941) and Cabrera
(1940, 1961) to differentiate Oreailurus
(Colocolo after Pocock) from Lynchailurus
and other South American felids showed
age-related variation, and the skull of O. ja-
cobita they based their results on was a sub-
adult (Philippi 1873). For example, Cabrera
(1961:203) described the sagittal crest of

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 114 JOURNAL OF MAMMALOGY
FIG. 3.—Qualitative characters and propor-
tions differing in skulls of A) and B) Oreailurus
jacobita and C) and D) Lynchailurus pajeros.
NB: nasal bones, SC: sagittal crest, PB: pre-
sphenoid body, EC: ectotympanic chamber of
bulla, EN: entotympanic chamber of bulla, APF:
anterior palatine foramina, C: upper canine, P3:
3rd upper premolar, P4: upper carnassial, M1:
1st upper molar.
Oreailurus as ‘‘only well developed in its
occipital tip,’’ and this is true for all sub-
adults and 1 adult of small skull size
(MUSM 6015), but not for 2 other adults
(CBF 445, CGECM 027), which have long
sagittal crests. It must be noted that values
of sagittal crest length increase with age in
other felids, sometimes remarkably so, and
that sagittal crest length is positively cor-
related with skull size (Garcı́a-Perea 1996).
Age-related variation in skins of O. ja-
cobita.—The only juvenile examined
(MACN 37.34) had a higher number of
blotches, smaller than those of adults and
of markedly darker color, especially those
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Vol. 83, No. 1
FIG. 4.—Qualitative characters and propor-
tions differing in skulls of A) Felis catus, B)
Oreailurus jacobita, and C) Lynchailurus pajer-
os. EC: ectotympanic chamber of bulla, EN: en-
totympanic chamber of bulla, PPJ: postorbital
process of jugal, PP: paroccipital process, P2:
upper 2nd premolar. Black arrows highlight cer-
tain anatomical differences mentioned in text.
from the ventral parts, which are chocolate
over a creamy background. Markings from
the sides are rosette-like, reddish-brown in-
side with darker borders. Tail rings are nar-
rower, closer to each other, and darker
brown than those of adults. Head is gray,
as in adults, but also darker. Black markings
on front legs are more conspicuous in the
kitten, but not forming rings.
In subadults, the general color becomes
lighter. Compared with adults, the ground
color is creamy instead of gray, and blotch-
es look reddish. As in the juvenile, blotches
are smaller and more numerous in subadults
than in adults and are distributed irregular-
ly. There is a blackish band along the back
(nonerectile), lacking the reddish tinge of
that in adults. Some subadults show con-
spicuous black markings on the front legs,
but these never form rings. Belly spots are
darker than those of adults. Tail rings are
more like those of adults than those of the
juvenile, but still narrower.
From the previous descriptions, data pre-
viously published (Garcı́a-Perea 1994), and
information provided subsequently (under
‘‘Coat characteristics of O. jacobita’’), it
follows that subadults of O. jacobita could
be mistaken for adult Lynchailurus from the
 February 2002 GARCÍA-PEREA—MORPHOLOGY OF ANDEAN MOUNTAIN CATS 115

FIG. 7.—Main external differences observed

FIG. 5.—Qualitative differences between up-
per carnassials (P4) of A) Lynchailurus pajeros
and B) Oreailurus jacobita. Pr: protocone, Pa:
parastyle, Ec: ectostyle.
between A) Oreailurus jacobita and B) Lyn-
chailurus pajeros from the altiplano. Black ar-
rows indicate characters differentiating the 2
species.

ual dimorphism of this species. However,

altiplano (especially in sightings), unless
the details of tail rings, front leg stripes, and
spinal crest are identified.
Skull and body measurements of O. ja-
cobita.—Values for skull and teeth variables
measured on specimens of O. jacobita are
summarized in Table 1. Because the only
skull of known sex studied was a male (Ta-
ble 1), it is risky to say anything about sex-
the relatively small size of specimen
MUSM 6015, an old adult with totally os-
sified sutures and rough braincase surface
(Table 1), suggests that MUSM 6015 could
be a female. This pattern of females being
smaller than males occurs in other felid spe-
cies (e.g., Lynx pardinus—Garcı́a-Perea et
al. 1985).
Data suggest that, although some large

FIG. 6.—External appearance of adult Oreailurus jacobita. Based on pictures of a living individual
and on museum specimens (see Appendix I).

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 116 JOURNAL OF MAMMALOGY Vol. 83, No. 1

TABLE 1.—Values (in millimeters) of 20 skull and teeth variables measured on 2 subadult and 3
adult specimens of O. jacobita. Sex is indicated where known. Additional data from 1 skull from
the literature are included. Specimen numbers refer to museums listed in Appendix I.

MVZ UG MUSM CBF CGECM From

Variable 116317 D2337 6015 445 027 literaturea
Age class Subadult Subadult Adult Adult Adult Subadult
Sex Male
Greatest length of skull 99.9 104.4 100.4 114.8 108.4 105.0
Condylobasal length 95.8 98.2 94.4 107.3 102.6 99.0
Rostral length 34.8 37.0 35.0 37.2 37.2
Rostral width across
the canines 24.8 25.0 24.8 27.3
Mastoid width 45.9 47.7 44.8 47.3
Palatal length 38.1 40.2 38.5 44.7 43.5
Interorbital width 20.6 23.2 23.8 26.7 24.1
Facial width 41.8 46.3 48.1 51.2 47.1
Postorbital width 29.0 29.9 29.1 28.4 25.9
Zygomatic width 69.1 72.9 71.2 79.7 78.5 71.0
Basilar axis length 33.8 33.5 31.7 35.3
Neurocranial height 34.9 35.5 36.0 38.4 35.6
Interbullae distance 15.0 12.8 13.5 13.0 13.5
Sagittal crest length 6.7 11.8 10.8 43.5 43.7
Length of P4 13.9 13.2 12.8 14.0 12.5 14.0
Maximum width of P4 7.1 6.9 6.9 7.2 7.1
Mandible length 66.9 70.2 65.1 75.5 72.6
Height of mandible ramus 29.6 30.6 28.7 34.2 34.4
Length of ml 11.0 10.3 9.8 11.4 11.3
Maximum width of ml 4.6 4.5 4.5 4.7 5.0
a
Skull illustrated by Philippi (1873) and measured by Cabrera (1961).

specimens of the pampas cat have skull di-
mensions similar to those of O. jacobita,
the latter species is significantly larger
(greatest length of skull 12% larger, con-
dylobasal length 14% larger) than altiplanic
specimens of L. pajeros and F. catus (great-
est length of skull 19% larger, condylobasal
length 21% larger; Table 2). Carnassial
teeth (P4 and m1) are also significantly
more massive in O. jacobita than in L. pa-
jeros and in F. catus (P4 length, m1 length;
Table 2). Note that average values and rang-
es of skull measurements of adult O. jacob-
ita given in Table 2 are larger that those
offered by Yensen and Seymour (2000:2),
because their sample included subadult
specimens.
Observation of skulls also shows other
differences: dorsal profile of skull is more
flat and elongated in O. jacobita than in L.
pajeros; dorsal profile of rostrum is lower
and more flat in O. jacobita than in L. pa-
jeros; adult specimens of O. jacobita usu-
ally show a long sagittal crest, which is
never so well developed in adult specimens
of L. pajeros from the altiplano; and audi-
tory bullae are relatively smaller and more
separate in O. jacobita (Figs. 3 and 4) than
in L. pajeros (interbullae distance; Table 2).
An elongated muzzle places postorbital pro-
cesses of O. jacobita closer to the middle
of the skull (Figs. 3A and 4B), as Yensen
and Seymour (2000) mention, but not as
much as Pocock (1941) indicates. As I not-
ed above, Pocock and Cabrera based their
descriptions on the subadult specimen illus-
trated by Philippi (1873).
Values of external measurements ob-
tained on tanned skins from Catamarca
were useful for establishing age classes, but
they could not be used for descriptive pur-
poses because they were likely overesti-

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 February 2002 GARCÍA-PEREA—MORPHOLOGY OF ANDEAN MOUNTAIN CATS
mated. In fact, specimen MVZ 116317
(Pearson 1957), a subadult male, shows
lower values of body measurements (on the
flesh) than those measured on subadult Ca-
tamarca specimens (Table 3). These differ-
ences are likely a consequence of change in
size due to the tanning method. Average
values given by Yensen and Seymour
(2000), based on the five specimens record-
ed in the literature (also given in Table 3),
are just suggestive, because age of speci-
mens is unknown, except for 1 subadult,
and subadult felids are significantly smaller
than adults (e.g., Garcı́a-Perea 1996;
Garcı́a-Perea et al. 1996).
Regarding the 10 qualitative traits show-
ing variation in skulls of O. jacobita and L.
pajeros, character states observed on the
sample are shown in Table 4. These char-
acters confirm that, as Kuhn (1973) men-
tioned, the relative sizes of ecto- and ento-
tympanic chambers of auditory bulla are di-
agnostic for O. jacobita in the altiplano, not
the presence of external groove, or sulcus,
separating both chambers (Pocock 1941),
which is not always obvious, as the sulcus
does not appear better developed in all old
individuals, as Seymour (1999) claims. In
O. jacobita, the ectotympanic chamber is
equal to or larger than the entotympanic
(Table 4), representing more than 50% of
bullar volume. However, this character is
not unique to O. jacobita, because L. colo-
colo from central Chile, and some species
of Felis (e.g., F. manul, F. margarita) also
have an ectotympanic chamber equal to or
larger than the entotympanic chamber
(Garcı́a-Perea 1994; Pocock 1916).
As Cabrera (1961) suspected, the 5 skulls
of O. jacobita lack P2 (similar to L. paje-
ros). Its dental formula is then I 3/3, C 1/
1, P 2/2, M 1/1, total 28.
Feral cats show a high amount of varia-
tion in qualitative characters of skull. For
this reason, I am providing in Table 5 some
typical, although not exclusive, skull char-
acters that, if restricted to the altiplano, can
help to identify F. catus (Fig. 4A).
Coat characteristics of O. jacobita.—A
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117
general view of an adult O. jacobita show-
ing the most conspicuous features charac-
terizing the coat pattern of this species is
shown in Fig. 6—based on photographs of
living individuals and on museum study
skins. Former illustrations of O. jacobita
shown by Cabrera and Yepes (1940:plate
29), by Seidensticker and Lumpkin (1991:
48), and by Redford and Eisenberg (1992:
plate 9h) fail to reflect accurately external
characters such as shape and distribution of
body markings, tail proportions, and tail
rings of the species.
The ground color of an adult O. jacobita
is ashy-gray, with yellowish-brown, irreg-
ular blotches arranged in vertical series on
the flanks (transverse lines, when observed
from the top). The head and face are gray,
with cheeks and areas around the lips white.
As in many other felids, 2 dark brown lines
run across the cheeks, converging laterally.
In some specimens, 2 dark gray lines start
above the eyes, running up to the space be-
tween the ears. From there, 2 wide yellow-
ish-brown bars run laterally down to the
base of the neck, sometimes with 1 or 2
bars of the same color between them. Ears
are gray, with darker borders. There is a
rusty-black band (nonerectile fur) along the
back. Some black spots occur in the ulnar
region of forelegs, but never forming com-
plete rings like those in L. pajeros. How-
ever, hind limbs may have 1 or 2 narrow,
dark rings, dorsally black and ventrally red-
dish. Belly is white or creamy with light
brown spots laterally and black spots me-
dially. One or 2 dark brown, transverse
stripes occur in the ventral part of the neck,
sometimes incomplete.
Tail is long (around 66–75% of head and
body length in fresh specimens), bushy, and
cylindrical, showing 6–9 rings varying
from black to dark brown. The 3 or 4 distal
rings are remarkably wider (up to 60 mm
wide) than those of the proximal half.
Two adult females had well-developed
mammae with no fur around them, sug-
gesting that they were rearing kittens when
captured. Both females had 2 pairs of mam-
mae located in abdominal position, and
 118 JOURNAL OF MAMMALOGY Vol. 83, No. 1

TABLE 2.—Descriptive statistics for 20 cranial variables of adult O. jacobita and L. pajeros and
of 8 variables of adult F. catus (after Pocock 1951); n 5 sample size. Level of significance is indicated
for Mann–Whitney U-test (comparing O. jacobita with each of the other species) and Student’s t-
test (comparing L. pajeros with F. catus). Symbols: *, P # 0.05; **, P # 0.005; and ***, P # 0.001;
ns, not significant (P . 0.05).

Skull size (mm)

n X̄ SE Range L. pajeros F. catus
Greatest length of skull
O. jacobita 3 107.9 4.165 100.4—114.8 * **
L. pajeros 19 96.6 0.795 89.2—103.2 **
F. catus 14 90.9 1.734 81.0—99.0
Condylobasal length
O. jacobita 3 101.4 3.769 94.4—107.3 ** **
L. pajeros 19 89.0 0.690 82.3—94.4 **
F. catus 14 83.8 1.726 73.0—92.0
Rostral length
O. jacobita 3 36.5 0.733 35.0—37.2 ns
L. pajeros 19 32.9 0.447 29.1—37.0
Rostral width across the canines
O. jacobita 2 26.1 1.250 24.8—27.3 ns
L. pajeros 19 23.4 0.439 20.3—28.9
Mastoid width
O. jacobita 2 46.1 1.250 44.8—47.3 *
L. pajeros 19 41.8 0.418 38.9—45.6
Palatal length
O. jacobita 3 42.2 1.899 38.5—44.7 **
L. pajeros 18 34.7 0.388 31.3—37.7
Interorbital width
O. jacobita 3 24.9 0.921 23.8—26.7 ** **
L. pajeros 19 18.4 0.258 16.6—21.0 *
F. catus 14 17.2 0.422 15.0—19.0
Facial width
O. jacobita 3 48.8 1.234 47.1—51.2 *
L. pajeros 19 44.4 0.576 38.5—47.6
Postorbital width
O. jacobita 3 27.8 0.971 25.9—29.1 ns ns
L. pajeros 19 27.7 0.320 25.2—30.2 ***
F. catus 14 31.1 0.601 27.0—34.0
Zygomatic width
O. jacobita 3 76.5 2.656 71.2—79.7 * *
L. pajeros 17 68.7 0.859 60.1—73.3 ns
F. catus 14 65.1 1.664 56.0—73.0
Basilar axis length
O. jacobita 2 33.5 1.800 31.7—35.3 ns
L. pajeros 19 31.5 0.349 29.1—34.3
Neurocranical height
O. jacobita 3 36.7 0.874 35.6—38.4 ns
L. pajeros 19 36.5 0.349 34.4—40.3

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 February 2002 GARCÍA-PEREA—MORPHOLOGY OF ANDEAN MOUNTAIN CATS 119

TABLE 2.—Continued.

Skull size (mm)

n X̄ SE Range L. pajeros F. catus
Inter-bullar distance
O. jacobita 3 13.3 0.167 13.0—13.5 **
L. pajeros 19 11.0 0.228 9.1—12.8
Sagittal crest length
O. jacobita 3 32.7 10.933 10.8—43.7 ns
L. pajeros 19 18.5 1.176 10.0—29.6
Length of P4
O. jacobita 5 13.2 0.296 12.5—14.0 *** ***
L. pajeros 19 11.6 0.104 11.0—12.6 ***
F. catus 14 10.1 0.152 9.0—11.0
Maximum width of P4
O. jacobita 5 7.0 0.060 6.9—7.2 ***
L. pajeros 19 5.5 0.086 4.6—6.2
Mandible length
O. jacobita 3 71.1 3.099 65.1—75.5 ** *
L. pajeros 18 61.8 0.695 54.2—65.5 ns
F. catus 14 60.5 1.455 51.0—66.0
Height of ramus of mandible
O. jacobita 3 32.4 1.868 28.7—34.4 ns
L. pajeros 19 27.8 0.505 21.6—30.9
Length of ml
O. jacobita 5 10.8 0.308 9.8—11.4 *** ***
L. pajeros 11 9.1 0.132 8.3—9.6 ***
F. catus 14 7.5 0.111 7.0—8.0
Maximum width of ml
O. jacobita 5 4.7 0.093 4.5—5.0 ***
L. pajeros 11 4.1 0.060 3.8—4.4

both were captured in December. Taking
into account that weaning usually occurs in
small felids about 2 months after birth
(Ewer 1973; Kitchener 1991), birth of both
litters could have occurred around October,
during austral spring.
Pelage of both O. jacobita and L. pajeros
differ conspicuously in 7 external charac-
ters, illustrated in Fig. 7 and described in
Table 4. Characteristics described are con-
sistent with those observed by L. Villalba
and N. Bernal (in litt.)
Tail of L. pajeros is shorter than that of
O. jacobita. It is necessary to be careful
when checking tail proportions, because
measurements taken on fresh specimens
differ from those taken on collection skins.
Regarding O. jacobita, data recorded from
literature (fresh specimens, n 5 5), indicate
that its tail length is about 66–75% of head
and body length, whereas data recorded
from collection specimens (n 5 11) give a
figure of 52–65%. Similarly, data on fresh
specimens of L. pajeros recorded from
specimen labels (n 5 9) indicate that tail
length is about 40–50% of head and body
length, whereas these variables measured
on collection specimens yield values that
are 34–38% of the length (n 5 9). My data
on fresh specimens of O. jacobita agree
with those given by Yensen and Seymour
(2000)—60–75% of length—but data on L.
pajeros given by these authors (ca. 30%)
seem to refer to collection specimens.

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 120 JOURNAL OF MAMMALOGY Vol. 83, No. 1

TABLE 3.—Values of 5 external measurements for O. jacobita, either found in the literature or measured on tanned collection skins for this
A great amount of variation is observed

Pine et al. 1979

in coat pattern of feral cats, too large to be

Matschie 1912
Cornalia 1865
Pearson 1957a

Cabrera 1961
Reference
treated here. However, I include in Table 5

This study
This study
the most striking characters of feral cats to
be considered when comparing with the 2
other wild genera, Oreailurus and Lynchail-
urus, occurring in the altiplanic area.
Body weight

CONCLUSIONS
4000
(g)

Felids are especially difficult to assess

morphologically. Usually, individual varia-
tion is moderate, and often odd character
states are found at very low frequencies. For
Length of
hind foot

this reason, finding diagnostic characters for

(mm)
133

110
115

closely related species is very difficult, and

it is safer to look for sets of traits character-
istic of a species, with a high probability of
finding them all together. Based on this pre-
Ear length

mise, a dichotomous key alone would not be

(mm)
63

53

enough for my purposes; so I am providing

sets of characters (Tables 4 and 5) that are
likely to occur together in a species, and if
one fails, we can confirm our identification
Tail length

330–420
410–485

by looking for a match with the others. From

(mm)

my experience, 1 or 2 characters are ex-

413
430
410
480
480

pected to be missing together. For F. catus,

only typical characters are provided, but we
must remember that they are not exclusive
body length

to F. catus and may not be present in some

Head and

640–660
740–850
(mm)

specimens. The following keys are only use-

ful within the limits of the altiplanic region,
577
600
850
640
640

because L. pajeros shows a certain amount

of geographic variation.
Key for skulls.—It is necessary to deter-
1
1
1
1
1
8
11
n

mine first whether the skull is from an adult

(with sphenooccipital synchondrosis ossi-
fied). If it is not, the key may not be useful.
After using the following keys, confirm
Subadult

Subadult
Age

identification using characters in Tables 4

Adult

Adult

and 5, especially for individuals with con-

study. n 5 sample size.

dylobasal length close to 94 mm (as certain

domestic cats could also have similar con-
Males and females
Males and females

dylobasal length).
MVZ 116317.

DICHOTOMOUS KEY FOR SKULLS

Sex

1a. Condylobasal length . 94 mm, lower

m1 length . 9.8 mm, dorsal profile of
Male

Male

skull flat and low, anterior chamber of

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 February 2002 GARCÍA-PEREA—MORPHOLOGY OF ANDEAN MOUNTAIN CATS 121

TABLE 4.—Character states observed on skulls of O. jacobita and L. pajeros, and external traits
characterizing skins of both species.

Characters Oreailurus jacobita Lynchailurs pajeros

Skull
Location of anterior palatine
foramina
Shape of nasal bones
Length of postorbital process of
jugal
Shape of body of presphenoid
Position of vagina processus hy-
oideus
Shape of paroccipital processes
Presence of lingual ridge in up-
per canine
Relative size of parastyle and
protocone of P4
Presence of metaconid on m1
Relative size of ectotympanic and Ectotympanic equal to or larger
entotympanic
Skin
Black rings on forelegs
Color of ears dorsally
Spinal crest of erectile fur
Pattern and color of body mark-
ings
Color of rhinarium
Tail rings
Tail length as percentage of head
and body length
At palatine-maxilla suture Posterior to palatine-maxilla suture
Slowly narrowing posteriorly Strongly narrowing posterior half
Short Long
Widened medially No medial widening
Posterior to stylomastoid foramen Medial to stylomastoid foramen
Long, separate, protruding Short, not protruding, cupped
ventrally around bulla
Absent Present
About the same size Larger than protocone or no proto-
cone
Present (80%) Absent
Ectotympanic smaller
Absent, only large spots Present, complete rings
Uniformly gray Black and cream and reddish
Absent Present
Vertical series of yellowish-brown Oblique series or rusty rosettes
blotches
Black Pink to coffee or dark mahogany
6–9 dark brown-to-black rings, Around 8 reddish rings, up to 20
some of them reaching up to 60 mm wide each
mm wide
Tail length about 66–75% of head Tail length about 40–50% of head
and body length and body length

bulla equal to or larger than posterior
. . . . . . . . . . . . . . . . . . . . . . . . . O. jacobita
1b. Condylobasal length # 94 mm, lower
m1 length , 9.6, dorsal profile high and
convex, anterior chamber of bulla small-
er than posterior . . . . . . . . . . . . . . . . . . 2
2a. Presence of lingual ridges on upper ca-
nines, lower m1 length . 8.0 mm, an-
terior chamber of bulla medially ex-
panded and inflated . . . . . . . . . . L. pajeros
2b. Absence of ridges on upper canines,
lower m1 length , 8.0 mm, anterior
chamber of bulla medially expanded but
not inflated . . . . . . . . . . . . . . . . . F. catus
Dichotomous key for skins.—Because of
the high amount of variation exhibited by
feral cats, this key may not be as effective
as the previous one for skulls; so I recom-
mend checking characters in Tables 4 and
5 carefully.
DICHOTOMOUS KEY FOR SKINS
1a. Medium-sized body; tail length around
66–75% of head and body length; no
spinal crest; no complete rings in fore-
legs; body sides with blotches arranged
in vertical series . . . . . . . . . . . O. jacobita
1b. Small-sized body; tail length ,50% of
head and body length; spinal crest pre-
sent; complete rings in forelegs; body
sides not marked or with rosettes ar-
ranged in oblique series . . . . . . . . . . . . 2
2a. Ears black and cream and reddish; body
sides with rusty rosettes arranged in
oblique series . . . . . . . . . . . . . . L. pajeros
2b. Ears uniformly colored; body sides not

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 122 JOURNAL OF MAMMALOGY
TABLE 5.—Skull and skin characters typical of
F. catus, absent in O. jacobita and L. pajeros.
Skull characters
P2 present
Unossified interparietal bone in adults
Posterior border of palate W-shaped
Ectotympanic expanded but no inflated
Skin characters
Plain color, no body markings
Medium-sized, slender tail, 2–5 black distal rings,
black tip
Ears of reddish color, or another color matching
body
marked or with another pattern different
from that described in 2a . . . . . . F. catus
Based on skull and skin characteristics
described earlier and from a detailed com-
parison with O. jacobita and L. pajeros ma-
terials, specimens CBF 2224, CBF 2960,
and CBF 2229, previously identified as O.
jacobita (Yensen et al. 1994) were identi-
fied as L. pajeros in this study.
RESUMEN
Recientes expediciones en busca del des-
conocido gato andino, Oreailurus jacobita,
se han encontrado con dificultades para
identificar esta especie a partir de avista-
mientos y pieles. Esto se debe a la escasez
de ejemplares en las colecciones (sólo se
conocı́an 3 cráneos hasta ahora) y a la au-
sencia de criterios para diferenciar esta es-
pecie de otros felinos simpátricos (por
ejemplo, Lynchailurus pajeros). Con el ob-
jeto de resolver estos problemas, se ha reali-
zado un estudio en el que se han identifi-
cado 3 nuevos cráneos de O. jacobita, ha-
biéndose examinado en total 5 cráneos y 41
pieles. Los cráneos de O. jacobita son un
12–14% más grandes que los de L. pajeros,
mostrando una gran cámara anterior en la
bula timpánica. El gato andino presenta se-
ries verticales de manchas pardo-amarillen-
tas en los flancos, ası́ como una larga cola
con abundante pelo y anchos anillos oscu-
ros. Se ofrecen dos claves para diferenciar
estas especies de felinos basadas en estos y
otros caracteres.
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Vol. 83, No. 1
ACKNOWLEDGMENTS
K. Nowell (Director, Cat Action Treasury)
and P. Jackson (former Chairman, Cat Specialist
Group, IUCN) gave me the opportunity to con-
duct this study. Funding was provided by Cat
Action Treasury, supported by the Leonard X.
Bosack and Bette M. Kruger Foundation. J. Gis-
bert (Museo Nacional de Ciencias Naturales,
MNCN, Madrid) drew Figs. 1–5 and 7. M. An-
tón drew Fig. 6, using pictures generously pro-
vided by J. Sanderson. I am indebted to the peo-
ple in charge of the collections listed in Appen-
dix I, who kindly provided access to them: E.
Vivar, N. Bernal, L. Villalba, M. Lucherini, E.
Luengos, J. L. Patton, H. J. Kuhn, A. Iriarte, J.
Yañez, J. C. Torres-Mura, R. Kraft, M. Piantan-
ida, O. Vaccaro, P. Jenkins, D. M. Hills, L. Gor-
don, M. Carleton, M. Rutzmosser, S. Anderson,
G. Musser, and B. Patterson. This study has also
benefited from grant DGICYT PB95-0114
(Spain, 1999), a Large Scale Facilities grant un-
der the Training and Mobility of Researchers
Programme (European Union, 1999), and from
a grant under the Agreement CSIC-Deutsche
Forschungsgemeinschaft (Spain–Germany,
1995). These funds were made available thanks
to J. Morales (MNCN, Madrid), P. Jenkins (The
Natural History Museum, London, United King-
dom), and G. Peters (Museum Alexander Ko-
enig, Bonn, Germany). Valuable comments of an
anonymous reviewer improved this article.
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APPENDIX I
Collections consulted for this study.—Ameri-
can Museum of Natural History, New York
(AMNH); The Natural History Museum, Lon-
don, United Kingdom (BM); Colección Bolivi-
ana de Fauna, Mastozoologı́a, La Paz, Bolivia
(CBF); Collection of the Grupo de Ecologı́a
Comportamental de Mamı́feros, Universidad
Nacional del Sur, Bahı́a Blanca, Argentina
(CGECM); Field Museum of Natural History,
Chicago, Illinois (FMNH); Museo Argentino de
Ciencias Naturales, Buenos Aires, Argentine
(MACN); Museum of Comparative Zoology,
 124 JOURNAL OF MAMMALOGY
Harvard University, Cambridge, Massachusetts
(MCZ); Museo Nacional de Historia Natural,
Santiago de Chile, Chile (MNHN); Museo de
Historia Natural Javier Prado, Lima, Peru
(MUSM); Museum of Vertebrate Zoology, Uni-
versity of California, Berkeley, California
(MVZ); Servicio Agrı́cola y Ganadero, Ministry
of Agriculture, Chile (SAG); University of Got-
tingen, Gottingen, Germany (UG); National Mu-
seum of Natural History, Washington D.C.
(USNM); Zoologisches Staatssammlung Mün-
chen, Munich, Germany (ZSM).
Catalog numbers.—Oreailurus jacobita.—
BM: 23.11.18.1; 23.11.18.2; 40.851. CBF:
00445; 02018; 02227; 02230. CGECM: 027.
MACN: 15.586; 29.200; 37.31; 37.32; 37.33;
37.34; 37.36; 37.37; 37.38; 37.106; 37.107;
37.108; 37.109; 37.111; 37.112; 37.114; 37.116;
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Vol. 83, No. 1
37.142; 37.143; 42.113; 9 uncataloged. MNHN:
1 uncataloged. MUSM: 6015; 1 uncataloged.
MVZ: 116317. SAG: 1 uncataloged. UG:
D2337. ZSM: 1984/12.
Lynchailurus pajeros.—BM: 26.5.3.6;
27.11.1.66; 27.11.1.67; 34.9.2.31; 34.11.4.5;
42.57. CBF: 02224; 02229; 02274; 02960;
06142; 06144; 06145; 06146. CGECM: 001;
022. FMNH: 21677; 25350; 49735; 52488;
68318. MACN: 14086; 17816; 25.33; 26186;
26187; 29765; 30103; 34322; 34326; 34565;
36230; 41163; 50446. MUSM: 01 J.O.; 415;
417; 418; 420; 421; 2150; 2151; 8467; 8468.
MVZ: 114777; 114942; 114943; 139613. ZSM:
1916/38; 1974/1.
Felis catus.—AMNH: 14079; 41557; 133972;
248700. CBF: 4 uncataloged. CGECM: 028; 1
uncataloged. MCZ: 51093. MVZ: 116007;
141633. USNM: 173336; 200285; 201072.