ArticlePDF Available

Abstract and Figures

1. The current classification of the Felidae was reviewed by a panel of 22 experts divided into core, expert and review groups, which make up the Cat Classification Task Force CCTF of the IUCN Cat Specialist Group. 2. The principal aim of the CCTF was to produce a consensus on a revised classification of the Felidae for use by the IUCN. 3. Based on current published research, the CCTF has fully revised the classification of the Felidae at the level of genus, species and subspecies. 4. A novel traffic-light system was developed to indicate certainty of each taxon based on morphological, molecular, biogeographical and other evidence. A concordance of good evidence in the three principal categories was required to strongly support the acceptance of a taxon. 5. Where disagreements exist among members of the CCTF, these have been highlighted in the accounts for each species. Only further research will be able to answer the potential conflicts in existing data. 6. A total of 14 genera, 41 species and 77 subspecies is recognised by most members of the CCTF, which is a considerable change from the classification proposed by Wozencraft (2005), the last major revision of the Felidae. 7. Future areas of taxonomic research have been highlighted in order to answer current areas of uncertainty. 8. This classification of the Felidae will be reviewed every five years unless a major new piece of research requires a more rapid revision for the conservation benefit of felid species at risk of extinction.
No caption available
… 
No caption available
… 
No caption available
… 
No caption available
… 
No caption available
… 
Content may be subject to copyright.
ISSN 1027-2992
CAT
news
A revised taxonomy of the Felidae
Special
Issue
I
I
N° 11 | WINTER 2017
CATnews Special Issue 11 Winter 2017
02
CATnews is the newsletter of the Cat Specialist Group, a component
of the Species Survival Commission SSC of the International Union
for Conservation of Nature (IUCN). It is published twice a year, and is
available to members and the Friends of the Cat Group.
For joining the Friends of the Cat Group please contact
Christine Breitenmoser at ch.breitenmoser@kora.ch
Original contributions and short notes about wild cats are welcome Send
contributions and observations to ch.breitenmoser@kora.ch.
Guidelines for authors are available at www.catsg.org/catnews
This Special Issue of CATnews has been produced with support
from the Taiwan Council of Agriculture’s Forestry Bureau, Zoo Leipzig and
the Friends of the Cat Group.
Design: barbara surber, werk’sdesign gmbh
Layout: Christine Breitenmoser and Tabea Lanz
Print: Stämpfli Publikationen AG, Bern, Switzerland
ISSN 1027-2992 © IUCN/SSC Cat Specialist Group
Editors: Christine & Urs Breitenmoser
Co-chairs IUCN/SSC
Cat Specialist Group
KORA, Thunstrasse 31, 3074 Muri,
Switzerland
Tel ++41(31) 951 90 20
Fax ++41(31) 951 90 40
<u.breitenmoser@vetsuise.unibe.ch>
<ch.breitenmoser@kora.ch>
Cover Photo: p.42-43 from Linnaeus 1758
This report should be cited as:
Kitchener A. C., Breitenmoser-Würsten Ch., Eizirik E., Gentry A., Werdelin L., Wilting A., Yamaguchi N., Abramov A. V., Christiansen P., Driscoll
C., Duckworth J. W., Johnson W., Luo S.-J., Meijaard E., O’Donoghue P., Sanderson J., Seymour K., Bruford M., Groves C., Hoffmann M.,
Nowell K., Timmons Z. & Tobe S. 2017. A revised taxonomy of the Felidae. The final report of the Cat Classification Task Force of the IUCN/
SSC Cat Specialist Group. Cat News Special Issue 11, 80 pp.
The designation of the geographical entities in this publication, and the representation of the material, do not imply the expression of any
opinion whatsoever on the part of the IUCN concerning the legal status of any country, territory, or area, or its authorities, or concerning the
delimitation of its frontiers or boundaries.
A new taxonomy of the Felidae
03
Introduction
The main task of the IUCN SSC Cat Specialist Group is the continuous review of the conservation status of all cat species
and subspecies according to The IUCN Red List of Threatened Species process. A critical subject in this task is the
systematic classification of the cat family, the Felidae. The taxonomy of cats has undergone considerable changes in the
past, not only at the level of species and subspecies, but even at the level of genus. The classification presently used by the
Cat Specialist Group was published in Wild Cats – Status Survey and Conservation Action Plan edited by K. Nowell and
P. Jackson (published by IUCN 1996) and is based on the state of research in the early 1990s. Since then mainly studies
using more advanced morphological, biogeographical and, foremost, molecular techniques have provided new insights
into cat phylogeny and variation, suggesting several important changes with regard to species and subspecies, and the
evolutionary relationships between genera and species. These changes may impact on the Red List process and on the
listing of taxonomic units in international treaties and national legislation. Therefore the classification used by IUCN
institutions has significance beyond the Red List. The Cat Specialist Group initiated a review of the present taxonomic
system of the Felidae by an expert group, the Cat Classification Task Force CCTF. Their Terms of Reference were endorsed
by Dr Simon Stuart, IUCN/SSC Chair 2008-2016.
Goal
The CCTF presents, on behalf of the Cat Specialist Group and the IUCN Red List Unit, and based on the best science and
expert knowledge presently available, an updated and practical classification of the Felidae, including genera, species and
subspecies, and the most likely geographical ranges of all taxa.
Principles
The starting point of the CCTF is the classification used by the Cat Specialist Group based on Nowell & Jackson (1996)
and the classification (species, subspecies) used in the present version of The IUCN Red List of Threatened Species (www.
iucnredlist.org), generally based on Wozencraft (2005). The CCTF has considered and reviewed all recent taxonomic
reviews and scientific publications on the taxonomy of cats to propose an updated classification. The review was
based mainly on new molecular and morphological research, but also considered general evolutionary, phylogenetic,
palaeontological, biogeographical, behavioural and physiological evidence, especially in cases where molecular genetics
and morphology are in disagreement. Conventional rules of zoological taxonomic nomenclature have to be respected, but
traditions in the use of non-scientific names (from Jackson et al. 1996) – especially in cases where subspecies are merged –
are also considered in order to produce a classification of cats useful for the practical work of the Cat Specialist Group and
conservation in general. The CCTF has suggested a set of principles and criteria for decisions regarding the acceptance
of proposed species and subspecies that can also be applied in future reviews. In case of uncertainty or lack of consensus,
introduction
A revised taxonomy of the Felidae
Executive summary
1. The current classification of the Felidae was reviewed by a panel of 22 experts divided into core, expert and
review groups, which make up the Cat Classification Task Force CCTF of the IUCN Cat Specialist Group.
2. The principal aim of the CCTF was to produce a consensus on a revised classification of the Felidae for use
by the IUCN.
3. Based on current published research, the CCTF has fully revised the classification of the Felidae at the level
of genus, species and subspecies.
4. A novel traffic-light system was developed to indicate certainty of each taxon based on morphological,
molecular, biogeographical and other evidence. A concordance of good evidence in the three principal
categories was required to strongly support the acceptance of a taxon.
5. Where disagreements exist among members of the CCTF, these have been highlighted in the accounts for
each species. Only further research will be able to answer the potential conflicts in existing data.
6. A total of 14 genera, 41 species and 77 subspecies is recognised by most members of the CCTF, which is
a considerable change from the classification proposed by Wozencraft (2005), the last major revision of the
Felidae.
7. Future areas of taxonomic research have been highlighted in order to answer current areas of uncertainty.
8. This classification of the Felidae will be reviewed every five years unless a major new piece of research
requires a more rapid revision for the conservation benefit of felid species at risk of extinction.
CATnews Special Issue 11 Winter 2017
04
Table 1: Members of the Cat Classification Task Force who have contributed to this report.
Core Group
Andrew Kitchener (Chair CCTF) National Museums Scotland, UK
Christine Breitenmoser-Würsten Co-Chair IUCN/SSC Cat Specialist Group, Switzerland
Eduardo Eizirik PUCRS, Porto Alegre, Brazil
Anthea Gentry The Natural History Museum, London, UK
Lars Werdelin Naturhistoriska riksmuseet, Stockholm, Sweden
Andreras Wilting Leibniz Institute for Zoo and Wildlife Research, Berlin, Germany
Nobuyuki Yamaguchi Qatar University, Doha, Qatar
Expert Group
Alexei Abramov Zoological Institute, Russian Academy of Sciences, Saint-Petersburg,
Russia
Per Christiansen
Carlos Driscoll Wildlife Institute of India, Dehradun, India
Will Duckworth IUCN/SSC Cat Specialist Group, UK
Warren Johnson Smithsonian Institution, USA
Shu-Jin Luo Peking University, China
Erik Meijaard Borneo Futures, Jakarta, Indonesia
Paul O’Donoghue University of Chester, UK
Jim Sanderson Small Wild Cat Conservation Foundation, USA
Kevin Seymour Royal Ontario Museum, Canada
Review Group
Mike Bruford University of Cardiff, UK
Colin Groves Australian National University, Acton, Australia
Mike Hoffmann Zoological Society London, UK
Kristin Nowell IUCN/SSC Cat Specialist Group, USA
Zena Timmons National Museums Scotland, UK
Shanan Tobe Arcadia University, USA
we use a conservative approach. Additional cat specialists reviewed the proposals of the CCTF, especially with regard to
the distribution of the taxa (e.g. the borders between neighbouring subspecies). Finally, after a peer-review process, the
proposals of the CCTF are published here and formally adopted as the current cat classification used by the IUCN/SSC
Cat Specialist Group until the next revision. An important role of the CCTF was to identify key areas for future research in
order to resolve current taxonomic uncertainties. We suggest future reviews every five years in order to keep pace with
future research, but so as not to be too disruptive for legislators, field workers, captive breeding programmes, museums,
educators and other cat workers, for whom an unstable and changing taxonomy could cause an enormous amount of
work and may lead to inconsistent approaches.
The Cat Classification Task Force CCTF comprised three groups:
1. Core Group members set out general principles of the approach to be taken, reviewed current evidence, consulted
with experts and drew up the proposed new classification of the felids.
2. Expert Group members provided specific and critical expert advice on species and subspecies, morphology, genetics,
biogeographical areas, etc., to support the core group. Membership was not fixed and varied as specific expertise was
required by the core group.
3. Review Group members provided a robust peer review of the proposed cat classification. Expert and Core Group
members assumed also the role of reviewers at this stage.
The CCTF was chaired by Dr. Andrew Kitchener, Principal Curator of Vertebrates at National Museums Scotland,
Edinburgh, UK.
Urs Breitenmoser and Christine Breitenmoser-Würsten
introduction
A new taxonomy of the Felidae
05
Stages in the development of the revised felid taxonomy
1. Establish general principles, definitions, and approach by Core
Group
- request input of Expert Group
- invite Expert Group to submit any further evidence
2. Review of genera and likely included species
- request input of Expert Group
- invite Expert Group to submit any further evidence
3. Review of species focussing on those of key significance
- request input of Expert Group
- invite Expert Group to submit any further evidence
4. Review of subspecies focussing on those of key significance
- request input of Expert Group
- invite Expert Group to submit any further evidence
5. Core group assembles draft report
6. Draft report reviewed by Review and Expert Groups
7. Final report drafted and reviewed for publication
General principles for CCTF to consider including defini-
tion of terms
In order to overcome any misunderstandings of approach owing to dif-
ferent definitions of technical terms, the CCTF has defined these as
used in the Task Force. This includes also the interrelationships be-
tween different technical terms and the taxonomic approaches that
have been taken, e.g. naming of clines, relationship between species,
subspecies, Evolutionary Significant Units ESUs, etc. and how to deal
with the domestic cat.
Suggested required definitions and the relevant relationships between
them in a hierarchical taxonomic order:
1. Genera – the CCTF has adopted an arbitrary cut-off date of the be-
ginning of the Pliocene (5.2 Mya ± 0.5 Mya) to define genera, following
Hennig (1965). This is consistent with the major diversification of felid
lineages during the late Miocene (Johnson et al. 2006).
2. Species – a group of individuals that share diagnostic morphologi-
cal and molecular characteristics and distinct evolutionary lineages
and biogeographical histories that allows them to be distinguished
from other species. Species are generally expected to have had a most
recent common ancestor with other species of at least 800,000 years
ago based on the divergence times of Li et al. (2016). Species may
hybridise with other species to a limited extent, but basic morphology,
behaviour and ecology remain unaffected except in areas of introgres-
sion.
3. Subspecies – a group of individuals within a species that mostly
share morphological and molecular characteristics that distinguish
them from most other individuals within a species and that occupy
a distinct part of the geographical range of the species. These distin-
guishing characters are not expected to be 100% diagnostic and gene
flow is also expected between subspecies where ranges are contigu-
ous.
4. Units
4.1. Ecotypes – Populations within a species that exhibit morpho-
logical and/or physiological adaptation to a particular environment
or habitat that differs from the environment/habitat of neighbouring
populations, but which display little or no genetic differentiation.
These populations exhibit phenotypic plasticity in the face of varying
environmental conditions.
4.2. ESUs – Evolutionary significant units are populations within a
species that may not be morphologically distinct, but which exhibit a
distinct evolutionary history such that they may represent local long-
term adaptation to environmental conditions or habitats. They may
represent species or subspecies awaiting recognition.
4.3. MUs – Management units are populations within a species that
are considered distinctive enough to warrant separate conservation
management to that of the other populations. They may be equiva-
lent to subspecies or to populations that are genetically differentiated
but which are not taxonomically distinctive, but which may be locally
adapted and considered worthy of separate conservation management.
5. Clines – Clines occur where species change gradually over geo-
graphical space, but all populations are genetically contiguous. Even
though (particularly the end) populations along a cline may appear
morphologically distinct, they do not represent distinct taxa.
6. Domestic cat – Following the ruling of the International Commis-
sion for Zoological Nomenclature (Opinion 2027; International Com-
mission on Zoological Nomenclature 2003), the domestic descendant
of the North African wildcat should be treated as a separate taxon,
which here is regarded as a full species, Felis catus (Gentry et al. 2004).
A system for indicating taxonomic certainty
Most scientific names for species and subspecies have little or no
scientific basis. Many are based on one, a few or even no specimens,
with few or no comparisons with related taxa and hence, in particular,
many subspecies names are likely to be invalid. However, for many
species and subspecies recent research based on more than one line
of evidence is not available. Currently there is no system for indicating
taxonomic certainty of particular taxa. Such a system would indicate
to users of taxonomies the reliability and rigour behind classifications
as well as highlighting areas where urgent research is needed.
Therefore, we propose a simple traffic-light system to indicate the
likely reliability of species and subspecies given available evidence.
At least three lines of correlated evidence are required for taxonomic
certainty:
1. Morphological – taxa are diagnosably distinct on the basis of
several characters (e.g. skull, pelage) in comparison with all other
members of a species or genus (excluding hybrids) from throughout
their respective geographical ranges. Average differences and size
differences alone are not considered reliable indicators of taxonomic
distinctiveness. Pelage characteristics may be especially variable
within species and hence may be of poor diagnostic value. Care
must also be taken that apparent differences are not clinal, espe-
cially where gaps in formerly contiguous distributions have occurred
recently.
2. Genetic – taxa are genetically distinct based on a variety of ge-
netic information, including mtDNA, Y-chromosome markers, Single
Nucleotide Polymorphisms SNPs, etc., but care should be taken that
alternative explanations, such as genetic drift, founder effects and
population bottlenecks, could explain apparent genetic distinctive-
ness of no taxonomic significance.
3. Biogeographical – distinct taxa are more likely to be recognised
where there are distinct geographical barriers relevant to the taxon,
e.g. rivers, seas, mountains, deserts, or where geological events, such
as sea-level changes, or volcanic eruptions are broadly coincident
with coalescence times, or where recolonisations following climate
change are consistent with former refugia. Phylogeographical patterns
goals, principles and working processes
CATnews Special Issue 11 Winter 2017
06
- Three or more lines of correlated evidence as outlined
above.
- Two or more lines of correlated evidence as outlined above
plus reasonable inferences based on data from closely re-
lated species; taxon likely to be distinct. Further research
required.
- One or no lines of evidence; status of taxon currently un-
known, but considered unlikely to be valid. Further research
required.
- Despite recent research, no evidence for distinctiveness,
which may formerly have been suggested, or based on in-
complete or erroneous data, or alternatively con(sub)speci-
ficity demonstrated.
of similar species or those that occur in similar habitats in the same
geographical range may be useful to infer probable taxa.
Care must be taken to avoid misinterpretation of all lines of evidence
caused by recent anthropogenic impacts, which may have isolated
populations by extirpation in intervening geographical areas. Ancient
hybridisation between taxa may also give false indications of con(sub)
specifity, leading to erroneous conclusions about taxonomic status.
Clinal variation may be interpreted incorrectly as two or more appar-
ently distinctive populations owing to poor or incomplete sampling.
Other lines of supporting evidence may also be useful:
4. Behavioural – e.g. predisposition to taming
5. Ecological – e.g. use of distinct habitats with appropriate
adaptations
6. Reproductive – e.g. seasonality or not of reproductive cycles.
Therefore, we propose a simple traffic-light system:
Species Subspecies Morphology Molecular Biogeography Certainty Comments
Neofelis nebulosa nebulosa ++ ++ ++
macrosceloides -(-) -Possible skull differences from
nebulosa, but could be clinal
brachyura -(-) ?+ Genetically and morphologically
similar to nebulosa
Neofelis diardi diardi ++ ++ ++
borneensis +++ ++
Skulls distinguishable between
subspecies, but pelage variation
poorly known
Below are two examples of the use of the traffic-light system. These summary tables are presented at the end of each species section to provide
a quick-to-read summary to help in rapid assessment of taxonomic certainty of taxa within and between species. Key: ++ good evidence within
category, e.g. skull/pelage; mtDNA/nDNA; + some evidence or reasonable, inference within category; - was investigated, but no evidence to
support distinction; o has never been investigated. If the symbol is in brackets, the validity of evidence is considered uncertain.
Genus Neofelis
Genus Leptailurus
Species Subspecies Morphology Molecular Biogeography Certainty Comments
Leptailurus serval serval ++ ++ ++
brachyurus + o o
constantina + o ++ Possibly distinct, but could be
synonym of serval
lipostictus + o ++ Possibly distinct, but could be
synonym of serval
phillipsi + + +
togoensis + o o
taxonomic certainty
A new taxonomy of the Felidae
07
A short history of felid systematics
Today at least 38 species of cats are recognised throughout the world
(excluding only Australasia and the polar regions), although recent
morphological and molecular research suggests that there may be
a few more. Traditionally cats have been classified into two main
groups; the big cats, mostly of the genus Panthera, and the smaller
cats, with the cheetah left as an odd afterthought, representing a very
early divergence from the felid line (Pocock 1917). This basic classifi-
cation stood the test of time throughout most of the twentieth century
until new techniques and analyses became available. The key charac-
teristic that was used to separate the big cats (Pantherinae) from the
smaller cats (Felinae) is the presence in big cats of an elastic ligament
in the hyoid apparatus below the tongue, which apparently allowed
big cats to roar, but not purr. Conversely, the bony hyoid of smaller
cats allowed them to purr but not roar. The other key characteristic,
which allowed for the separation of the cheetah in its own subfamily,
the Acinonychinae, was the absence of cutaneous sheaths to protect
the retracted claws. However, recent studies of hyoid structure and
vocal abilities of cats have found that this simple correlation does not
hold. While it is true that some big cats roar (e.g. lion, leopard), not all
are able or confirmed able to do so, despite having an elastic hyoid. It
was found that the fundamental difference between the mostly roar-
ing non-purring cats and the rest was the structure of the larynx (Hast
1989). Long, fleshy, elasticated vocal folds within the larynx of big cats
resonate to produce a roar, whereas the smaller cats, including the
cheetah, have simpler vocal folds that only allow purring.
Although the number of species of cats is fairly well known (with a
handful of exceptions), the number of genera that have been recog-
nised is very variable. From a proliferation of genera or subgenera dur-
ing the 19th century, there was a lumping together into a handful during
the middle of the 20th century, followed by a final flourish and re-rec-
ognition of many of the 19th century names towards the end of the
20th century. Therefore, at one extreme only two or three genera were
used to classify all felids, whereas today there are varying opinions,
with as many as 18 being recognised on the basis of several studies of
morphology and genetics. This uncertainty is probably due to the high
degree of similarity in basic body plan between all felid species, with
the exception of the cheetah, and a lack of congruence between dif-
ferent sets of characters. Perhaps the recent radiation of the cat family
as we know it today, coupled with the constraints of prey capture and
the processing of a highly carnivorous diet, have resulted in a limited
range of variation within the felids.
Two major developments towards the end of the 20th century have
helped change our view of felid systematics. Firstly, the development
of molecular techniques, including the polymerase chain reaction, has
allowed the sequencing of mitochondrial genes and nuclear microsat-
ellites, in particular, so that differences in the sequences of base pairs
of DNA can be elucidated. Secondly, the advent of personal computers
coupled with powerful software, and the use of advanced statistical
methods, have allowed vast datasets of morphological and molecular
characters to be analysed in order to produce dendrograms of relation-
ships between species. By adding in a time element by correlating
known evolutionary events with divergences between species, it has
also been possible to calibrate phylogenies using the so-called mo-
lecular clock, for comparison with the fossil record. Encouragingly, in
broad outline there is a high degree of agreement between the various
morphological and molecular phylogenies, but inevitably some species
have been difficult to pin down. Even a study of lipids from the anal
sacs of 16 cat species showed a high degree of agreement with more
typical phylogenies (Bininda-Emonds et al. 2001).
To some extent the advent of molecular techniques, in particular, has
helped overcome some of the taxonomic conundrums of the past. For
example, on the basis of its highly specialised morphology for cursorial
hunting, the cheetah was often placed in its own subfamily and re-
garded as representing a very early offshoot of the felid line. Another
example is the caracal, which was often regarded as a close relative
of the bobcat, Eurasian, Canadian and Iberian lynxes of the northern
hemisphere, presumably on the basis of its tufted ears and short tail.
However, molecular phylogenies based on a variety of techniques
demonstrate that the cheetah is found within the main felid radiation
(the pantherine lineage) and is closely related to the puma (Johnson et
al. 2006, Li et al. 2016). The caracal was also found to have diverged
from this pantherine lineage as part of its own lineage with the ser-
val and African golden cat and is unrelated to the lynxes, which do
form a monophyletic clade, thus confirming Werdelin’s (1981) earlier
study based on morphology. Molecular techniques can also be used on
some recent fossils of extinct taxa; a recent study (Barnett et al. 2005)
showed that the sabre-toothed cats, Smilodon and Homotherium, are
sister taxa to the crown group of modern felids and not in Panthera as
previously shown.
The main Old World lineage of small cats comprises those species that
belong to the genus Felis proper, including the wildcats, sand cat, jun-
gle cat, and the domestic cat. The most recent molecular data suggest
that the Felinae and Pantherinae diverged about 11.5 million years
ago and that the eight felid lineages diverged sequentially from 4.23
to 10.67 million years ago (Li et al. 2016). Felis proper diverged from
the Leopard Cat lineage about 7.25 million years ago. The most basal
lineage of the Felinae is the Caracal lineage, followed by the Ocelot
lineage, the Lynx and Bay Cat lineages, the Puma lineage, and finally
the Leopard Cat and Domestic Cat lineages. Some genera and species
have been difficult to place in the felid phylogeny. For example, Oto-
colobus has been associated as a basal member of the Domestic Cat
lineage, but in the latest phylogeny it is basal in the Leopard Cat line-
age. The African and Asiatic golden cats were formerly considered to
be close, but they occupy different lineages, i.e. the Asiatic golden cat
and the bay cat form a distinct genus, Catopuma, in south east Asia,
but the apparently morphologically similar African golden cat is clos-
est to the caracal (both in the genus Caracal), which has no close rela-
tionship to the lynxes. The marbled cat (Pardofelis) is also found in the
Bay Cat lineage, but is more distantly related to Catopuma. The once
aberrant cheetah (genus Acinonyx) is in the same lineage as the puma
(Puma) and jaguarundi (Herpailurus), although some include the jagua-
rundi in Puma. Fossil cheetahs (genus Miracinonyx) in North America
that date back to three million years ago are very similar to pumas,
supporting a close relationship between the lineages, although some
palaeontologists regard the similarity between Old World and Ameri-
can cheetahs as being due to convergent evolution (Adams 1979).
As defined here, Miracinonyx would be subsumed within Puma. The
Puma lineage probably diverged more than eight million years ago on
the basis of the molecular clock (Johnson et al. 2006, Li et al. 2016).
As would be expected the lynxes form a coherent group (genus Lynx),
supported by molecular and morphological analyses, which seems to
have evolved to exploit the radiation of lagomorphs in the northern
hemisphere. The molecular clock suggests a common ancestor dating
back some 3.48 million years ago (Li et al. 2016).
history of felid systematics
CATnews Special Issue 11 Winter 2017
08
A recent molecular phylogeny (Li et al. 2016) showed that the diver-
gence and radiation of small South American cats occurred almost
10 million years ago, which was long before the Panama land bridge
formed about 3-5 million years ago to join South and North America.
Therefore, this radiation occurred in North America and indeed cats
are not known from South America until a maximum of 1.8 million
years ago. This isolation and radiation in the Americas is supported
by differences in chromosome numbers between the two groups; the
small South American cats of this group have two fewer chromosomes
than the 38 of other lineages. These New World cats belong to the
genus Leopardus, although more genera were recognised until re-
cently, including Oncifelis (Geoffroy’s cat and guigna), Oreailurus (An-
dean mountain cat) and Lynchailurus (pampas cat). However, natural
hybridisation is frequent among some species of Leopardus, including
Geoffroy’s cat, pampas cat and tigrinas, emphasizing their close rela-
tionships (Li et al. 2016).
The big cats also form a monophyletic group comprising the clouded
leopards (Neofelis) and the remaining big cats (Panthera). Molecular
data suggest a common ancestor some 5.67 million years ago, which
at first sight seems discordant with a fossil record that goes back only
2-3 million years ago (although a recent putative snow leopard an-
cestor, Panthera blytheae from Tibet, has been dated to 4.4 million
years ago and may be almost six million years old (Deng et al. 2011,
Tseng et al. 2013). However, the clouded leopards represent the earli-
est divergence from this lineage and if the common ancestor of this
group was also a rain forest inhabitant, it is unlikely that fossils will
have survived, owing to poor preservation conditions. Therefore the
fossil record of this group is likely to be deficient. Within the genus
Panthera, recent molecular analyses have concluded that the snow
leopard and tiger are sister species and diverged earlier from the an-
cestors of the jaguar, leopard and lion, of which the latter are also
sister species (Davis et al. 2010, Li et al. 2016). Clearly more research
needs to be done in all fields, not just the molecular side. The recent
radiation of the Felidae and their conservative morphology will prob-
ably continue to give systematists trouble for some time to come and
result in continuing instability in the classification of felid species at
the generic level and above.
Bininda-Emonds et al. (1999) published the first attempt to combine
carnivoran phylogenies from different molecular and morphological
studies. In the case of the felids 40 part or whole phylogenies were
combined. The combined phylogeny for felids still places the cheetah
as a distinct lineage from the other cats, but puts the marbled cat back
into the Pantherinae, the African golden cat into a group with Asiatic
golden cat and bay cat, finds the Pallas’s cat and serval as basal to
the Felis group, but otherwise places species into the same groups as
described above. All in all it demonstrates once more that although
there are well-defined groups within the Felidae, their interrelation-
ships are still uncertain. This was recently updated by Nyakatura &
Bininda-Emonds (2012) with similar results.
As mentioned earlier there may still be some cat species waiting to be
recognised and this review will highlight where there is clear or some
evidence for this. The other contentious area concerning felid system-
atics is the number of felid subspecies that should be recognised.
There is a plethora of subspecies names associated with the cat spe-
cies, but it must be remembered that in almost all cases these are not
based on scientific research, but have been used as handy labels for one
or a handful of (often atypical) specimens from particular geographical
locations. Putting aside questions of how we define and recognise sub-
species, which are common to all animals, there is a clear need for taxo-
nomic revisions (based on a variety of studies) of all species to determine
whether geographical variation within species is present or not, and if
so, whether it is clinal or discrete and of taxonomic significance. Many
of the problems of lack of concordance between traditional classifica-
tions and molecular studies have arisen because of the misplaced as-
sumption that these classifications have some basis in science. Where
revisions have been made, there is a great deal of common ground.
In recent years there has been a growth in the application of the phy-
logenetic species concept (PSC) in taxonomic revisions, particularly
in ornithology. The PSC defines species on the basis of populations,
which have at least one unique diagnosable character. In ornithology
the application of the PSC has seen many former subspecies raised
to species level. Groves (2001) applied this concept to primates as
the only feasible alternative to other species concepts and this has
also resulted in a proliferation of primate species. There is concern
that uncritical application of this species concept will lead to a huge
rise in the number of recognised cat species, with implications for
the conservation of many more endangered species (e.g. Zachos et
al. 2013, but see also Meijaard & Rawson 2015). However, we have
taken a conservative approach that relies on at least three independ-
ent lines of existing evidence to confirm the recognition of species
and subspecies. While our review will not be the last word written
on felid taxonomy, we hope the approach we have taken will provide
a solid baseline against which future changes can be made and will
offer taxonomic stability that will provide confidence for current and
future conservation management of many endangered species.
A brief review of modern felid genera
The starting point for our discussion will be Johnson et al. (2006), in
which 11 genera are recognised. We have followed Hennig (1965) in
recognising as distinct genera those lineages that diverged in the late
Miocene, c. 5 Mya. On this basis, the following genera are recognised.
1. Felis – there is a general consensus on this genus, although the
number of included species is uncertain.
2. Otocolobus – this monotypic genus is retained for the unusual Pal-
las’s cat, which is weakly aligned with Prionailurus or, more unlikely,
with Felis.
3. Prionailurus – this genus is strongly supported, although there
has previously been discussion about the inclusion of the rusty-spotted
cat, which diverged early, but it does appear morphologcally to be in
this genus.
4. Puma – Johnson et al. (2006) include the puma and jaguarundi in
this genus. Johnson et al. (2006) found the jaguarundi to be the sis-
ter species to the puma and hence included it in the genus Puma,
although Agnarsson et al. (2010) found that the jaguarundi was not
a sister species to Puma and retained Herpailurus. There are dis-
tinct differences in morphology and behaviour between the two, and
Segura et al. (2013) found that cranial development between Puma
and Acinonyx was more similar to each other than between Puma
and Herpailurus. Chimento et al. (2014) included the jaguarundi and
Puma pumoides in the subgenus Herpailurus within the genus Puma
in their morphological review of Puma pumoides. The problem in
employing Hennig’s (1965) criterion for generic recognition is that
this depends on the dating of the divergence between the puma and
jaguarundi lineages, which in turn depends on which fossils are used
to calibrate the molecular tree. For example, although Johnson et al.
(2006) date this divergence more recently at a mean of 4.17 Mya,
history of felid systematics
A new taxonomy of the Felidae
09
Barnett et al. (2005) found that the divergence date for the jagua-
rundi lineage varied from a mean of 5.03 Mya to a mean 7.42 Mya,
depending on whether Pseudaelurus or Proailurus is used to cali-
brate the molecular tree. However, these dates fall within the late
Miocene, which would lead to retention of Herpailiurus as a distinct
genus (but see Li et al. 2016). There is no clear resolution of this mat-
ter, in which case the CCTF has been asked to retain a conservative
position, so that provisionally Herpailurus has been retained as a
distinct genus until further evidence is available.
5. Herpailurus – see discussion under Puma above.
6. Acinonyx – there is general consensus on this genus, which is
clearly defined.
7. Lynx – there is a general consensus on this genus, which is clearly
defined.
8. Leopardus – in the recent past this genus has been further subdi-
vided into four genera (Leopardus, Oncifelis, Lynchailurus and Oreai-
lurus), but the recent radiation of these species, natural hybridisation
and the close similarity in skull morphology between these species
supports Johnson et al.’s (2006) conclusion of a single genus.
9. Caracal – formerly the African golden cat (Profelis) and serval (Lep-
tailurus) were in separate genera. Using Hennig’s (1965) criterion the
serval would be retained in a separate genus, but Profelis is subsumed
into Caracal.
10. Leptailurus - see Caracal
11. Pardofelis – the marbled cat is morphologically very distinct from
the Asiatic golden cat and bay cat (in a way that the margay is not
distinct from the ocelot or other Leopardus spp.). Further based on Li
et al. (2016), the marbled cat split from the Asiatic golden cat and
the bay cat about 5.5 Mya. Therefore, applying Hennig’s (1965) crite-
rion together with the morphological differences, it is recommended
separating the marbled cat from the latter two, thereby reinstating
Catopuma for the bay cat and Asiatic golden cat.
12. Catopuma - see Pardofelis
13. Neofelis – there is a general consensus on this genus.
14. Panthera – there is a general consensus on this genus with the
possible exception of the inclusion of the snow leopard, which has
been separated into its own genus, Uncia. However, recent molecular
studies show that the snow leopard is the sister species to the tiger
(Davis et al. 2010, Li et al. 2016), so that either these two should be
separated from the other Panthera spp. or all should be retained in
Panthera, which would also be supported by Hennig’s (1965) criterion
and which we support.
Therefore, the CCTF recognises three genera, Herpailurus, Leptailu-
rus and Catopuma, in addition to the 11 recognised by Johnson et al.
(2006), although this could be reduced to a total of 13 if further re-
search supports the inclusion of Herpailurus in Puma.
References
Adams D. B. 1979. The cheetah: Native American. Science 205, 1155-1158.
Barnett R., Barnes I., Phillips M. J., Martin L. D., Harington C. R., Leonard J. A.
& Cooper A. 2005. Evolution of the extinct sabretooths and the American
cheetahlike cat. Current Biology 15, 589-590. Supplemental data.
Bininda-Emonds O. R. P., Gittleman J. L. & Purvis A. 1999. Building large trees
by combining phylogenetic information: a complete phylogeny of the ex-
tant Carnivora (Mammalia). Biological Reviews 74, 143-175.
Bininda-Emonds O. R. P., Decker-Flum D. M. & Gittleman J. L. 2001. The utility
of chemical signals: An example from the Felidae. Biological Journal of the
Linnaean Society 72, 1-15.
Chimento N. R., Derguy M. R. & Hemmer H. 2014. Puma (Herpailurus)
pumoides (Castellanos, 1958) nov. comb. Comentarios sistemáticos
y registro fósil. Estudios Geológicos, Serie Correlación Geológica, 30,
92-134.
Collier G. E. & O’Brien S. J. 1985. A molecular phylogeny of the Felidae: Im-
munological distance. Evolution 39, 473-487.
Cracraft J., Feinstein J., Vaughn J., and Helm-Bychowski K. 1998. Sorting out
tigers (Panthera tigris): mitochondrial sequences, nuclear inserts, system-
atics, and conservation genetics. Animal Conservation 1, 139-150.
Culver M., Johnson W. E., Pecon-Slattery J. & O’Brien S. J. 2000. Genomic
ancestry of the American puma (Puma concolor). Journal of Heredity 91,
186-197.
Davis B. W., Li G. & Murphy W. J. 2010. Supermatrix and species tree methods
resolve phylogenetic relationships within the big cats, Panthera (Carnivora:
Felidae). Molecular Phylogenetics and Evolution 56, 64-76.
Deng T., Wang X., Fortelius M., Li Q., Wang Y., Tseng Z. J., Takeuchi G. T.,
Saylor J. E., Säilä L. K. & Xie G. 2011. Out of Tibet: Pliocene woolly rhino
suggests high-plateau origin of Ice Age megaherbivores. Science 333,
1285-1288.
Garcia-Perea R. 1994. The pampas cat group (genus Lynchailurus Severtzov,
1858) (Carnivora: Felidae), a systematic and biogeographic review. Ameri-
can Museum Novitates No. 3096, 1-36.
Gentry A., Clutton-Brock J. & Groves C. P. 2004. The naming of wild animal
species and their domestic derivatives. Journal of Archaeological Science
31, 645-651.
Groves, C.P. 2001. Primate taxonomy. Smithsonian Institution Press Washing-
ton, D.C.
Hast M. H. 1989. The larynx of roaring and non-roaring cats. Journal of Anat-
omy 163, 117-121.
Hennig W. 1965. Phylogenetic systematics. Annual Review of Entomology 10,
97-116.
International Commission on Zoological Nomenclature. 2003. Usage of 17
specific names based on wild species which are pre-dated by or contem-
porary with those based on domestic animals (Lepidoptera, Osteichthyes,
Mammalia): Conserved. Bulletin of Zoological Nomenclature 60, 81-84.
Jackson P., Farrell Jackson A., de Crem J. & Devitre D. 1996. Les félins. Laus-
anne and Paris: Delachaux et Niestlé.
Janczewski D. N., Yuhki N., Gilbert D. A., Jefferson G. T. & O’Brien S. J. 1992.
Molecular phylogenetic inference from saber-toothed cat fossils of Rancho
La Brea. Proceedings of the National Academy of Sciences of the United
States of America 89, 9769-9773.
Johnson W. E., Culver M., Iriarte J. A., Eizirik E., Seymour K. L. & O’Brien S. J.
1998. Tracking the evolution of the elusive Andean mountain cat (Oreailu-
rus jacobita) from mitochondrial DNA. The Journal of Heredity 89, 227-232.
Johnson W. E., Pecon-Slattery J., Eizirik E., Kim J-H., Menotti-Raymond M., Bo-
nacic C., Cambre R., Crawshaw P., Nunes A., Seuánez H. N., Moreira M. A.
M., Seymour K. L., Simon F., Swanson W. & O’Brien S. J. 1999. Disparate
phylogeographic patterns of molecular genetic variation in four closely re-
lated South American small cat species. Molecular Ecology 8, 79-94.
Johnson W. E., Eizirik E., Pecon-Slattery J., Murphy W. J., Antunes A., Teel-
ing E. & O’Brien S. J. 2006. The late Miocene radiation of modern Felidae:
A genetic assessment. Science 311, 73-77.
Kitchener A. C. 1999. Tiger distribution, phenotyoic variation and conservation
issues. In Riding the tiger. Tiger conservation in human-dominated land-
scapes. Seidensticker J., Christie S. & Jackson P. (Eds). Cambridge Univer-
sity Press, Cambridge, pp. 19-39.
Kitchener A. C. & Dugmore A. J. 2000. Biogeographical change in the tiger.
Animal Conservation 3, 113-124.
review of modern felid genera
CATnews Special Issue 11 Winter 2017
10
Larson S. E. 1997. Taxonomic re-evaluation of the jaguar. Zoo Biology 16, 107-120.
Leyhausen P. 1979. Cat behavior. Garland STPM Press, New York.
Li G., Davis B. W., Eizirik E. & Murphy W. J. 2016. Phylogenomic evidence
for ancient hybridization in the genomes of living cats (Felidae). Genome
Research 26, 1-11.
Martin L. D. 1980. Functional morphology and the evolution of cats. Transac-
tions of the Nebraska Academy of Sciences 8, 141-154.
Masuda R., Lopez J. V., Pecon-Slattery J., Yuhki N. & O’Brien S. J. 1996. Mo-
lecular phylogeny of mitochondrial cytochrome b and 12S rRNA sequences
in the Felidae: Ocelot and domestic cat lineages. Molecular Phylogenetics
and Evolution 6, 351-365.
Meiaard E. & Rawson B. 2015. The phylogenetic species concept and its role
in Southeast Asian mammal conservation. In Taxonomic tapestries: The
threads of evolutionary, behavioural and conservation research. Behie A.
M. & Oxenham M. F. (Eds). The Australian National University Press, Can-
berra, pp. 345-360.
Nowak R. 1996. Walker’s mammals of the world. 6th edition. Johns Hopkins
University Press, Baltimore.
Nowell K. & Jackson P. 1996. Wild cats. Status survey and conservation action
plan. IUCN, Gland.
Nyakatura K. & Bininda-Emonds O. R. 2012. Updating the evolutionary history
of Carnivora (Mammalia): A new species-level supertree complete with
divergence time etsimates. BMC Biology 10:12.
Pocock R. I. 1917. The classification of existing Felidae. Annals and Magazine
of Natural History (8th series) 20, 329-350.
Salles L. O. 1992. Felid phylogenetics: Extant taxa and skull morphology (Feli-
dae, Aeluroidea). American Museum Novitates No. 3047, 67 pp.
Segura V., Prevosti F. & Cassini G. 2013. Cranial ontogeny in the Puma line-
age, Puma concolor, Herpailurus yagouaroundi, and Acinonyx jubatus (Car-
nivora: Felidae): A three-dimensional geometric morphometric approach.
Zoological Journal of the Linnean Society 169, 235-250.
Thenius E. 1967. Zur Phylogenie der Feliden (Carnivora, Mamm.). Zeitschrift für
zoologische Systematik und Evolutionforschung 5, 129-143.
Trigo T. C., Schneider A., de Oliveira T. G., Lehugeur L. M., Silveira L., Freitas,
T. R. O. & Eizirik E. 2013. Molecular data reveal complex hybridization and
a cryptic species of Neotropical wild cat. Current Biology 23, 2528-2533.
Tseng Z. J., Wang X., Slater G. J., Takeuchi G. T., Li Q., Liu J. & Xie G. 2014.
Himalayan fossils of the oldest known pantherine establish ancient origin
of big cats. Proceedings of the Royal Society B: Biological Sciences 281
(1774), 20132686.
Werdelin L. 1981. The evolution of lynxes. Annales Zoologici Fennici 18, 37-71.
Zachos F. E, Apollonio M., Bärmann E. V., Festa-Bianchet M., Göhlich U., Habel
J. C., Haring E., Kruckenhauser L., Lovari S., McDevitt A. D., Pertoldi C.
2013. Species inflation and taxonomic artefacts – A critical comment on
recent trends in mammalian classification. Mammalian Biology-Zeitschrift
für Säugetierkunde 78, 1-6.
Species Accounts
The sequence of species below follows the phylogenetic tree of Li
et al. (2016) from the most basal member of each lineage, starting
with the domestic cat lineage. We provide maps based on the most
recent Red List assessments (www.iucnredlist.org) with approximate
locations of subspecies. For monotypic species we do not show a dis-
Museum abbreviations
AMNH American Museum of Natural History, New York, NY, USA
ANSP Academy of Natural Sciences, Philadeplphia, PA, USA
BMNH Natural History Museum, London, UK
MACN Museo Argentino de Ciencias Naturales “Bernardino Rivadavia”, Buenos Aires,
MCZ Museum of Comparative Zoology, Cambridge, Massachusetts, USA
ZMB Museum für Naturkunde, Berlin, Germany
MLP Museo de La Plata, Buenos Aires, Argentina
MNCN Museo Nacional de Ciencias Naturales, Madrid, Spain
MNHN Museum national d’Histoire Naturelle, Paris, France
MNHNM Museo Nacional Historia Natural, Montevideo, Uruguay
MSNM Museo Civico di Storia Naturale, Milan, Italy
MZS Musée zoologique de la ville de Strasbourg, Strasbourg, France
NMW Naturhistorisches Museum Wien, Vienna, Austria
NRM Naturhistoriska riksmuseet, Stockholm, Sweden
NSM - National Museum of Nature and Science, Tokyo, Japan
RMCA Royal Museum for Central Africa, Tervuren, Belgium
RMNH Naturalis Biodiversity Center, Leiden, The Netherlands
ROM Royal Ontario Museum, Toronto, Canada
SMF Senckenberg Forschungsinstitut und Naturmuseum, Frankurt am Main, Germany
TM Ditsong: National Museum of Natural History, Pretoria, Republic of South Africa
USNM National Museum of Natural History, Smithsonian Institution, Washington DC, USA
ZFMK Zoologisches Forschungsmuseum Alexander König,Bonn, Germany
ZMMU Zoological Museum of Moscow State University, Moscow, Russia;
ZIN Zoological Institute of the Russian Academy of Sciences, Saint-Petersburg, Russia (each specimen from ZIN has two collection num-
bers – one for skin, and another for skull (skeleton)).
review of modern felid genera
tribution map as they are available on the IUCN Red List website. We
illustrate each species with at least one photograph.
Non-scientific names abbreviations
E – English; F – French; G – German; Sp – Spanish
A new taxonomy of the Felidae
11
Family Felidae Fischer, 1817; 372
Subfamily Felinae Fischer, 1817; 372
Domestic Cat lineage
Genus Felis Linnaeus, 1758; 41
The genus Felis usually includes between four and six species. Here
we provisionally recognise six species.
Felis chaus
E: Jungle cat, swamp cat; F: Chat de marais, chat de jungle, chaus; G:
Rohrkatze, Sumpfluchs; Sp: Gato de la jungla, gato de los pantanos.
Up to 10 subspecies have been recognised (Wozencraft, 2005):
Felis chaus chaus Schreber, 1777a; 414 and 1777b; pl. 110B.
Locality from where species was first described: ”... wohnt in den sump-
figen mit Schilf bewachsenen oder bewaldeten Gegenden der Steppen
um das kaspische Meer, und die in selbiges fallenden Flüsse. Auf der
Nordseite des Terekflusses und der Festung Kislar siehet man ihn selten,
und gegen die Wolga hin gar nicht; desto häufiger aber bei der Münd-
ung des Kur, und in den persischen Landschaften Gilan und Masand-
eran” [= lives in marshes overgrown with reeds or forested areas of the
steppes around the Caspian Sea, and the same in the surrounding rivers.
On the north side of the Terek River around the fortress Kislar it is rarely
seen, and not at all towards the Volga; but more often at the mouth of
the Kur River, and in the Persian provinces of Ghilan and Mazanderan],
i.e. Terek River, Northern Caucasus, Dagestan, Russia.
Type: None designated, based on Gueldenstaedt’s (1776) Chaus.
Distribution: Turkestan, Caucasus, Iran and Baluchistan, Pakistan.
Felis chaus nilotica de Winton, 1898; 292.
Type locality: near Cairo [Egypt].
Holotype: BMNH 1898.6.5.1 adult male skin and skull.
Distribution: Egypt, the delta district extending westwards to Mersa
Matruh, 155 miles W of Alexandria and S along Nile to Fayum, Quena
Province and probably Mina Province.
Felis chaus furax de Winton, 1898; 293.
Type locality: Jericho.
Holotype: BMNH 1864.8.17.4 male skull .
Distribution: S Syria and Iraq.
Felis chaus prateri Pocock, 1939; 298.
Type locality: Jacobabad, on the Upper Sind Frontier [Pakistan].
Holotype: BMNH 1832.2.1.67 male skin and skull .
Distribution: Sind from upper frontier to Larkana and Karachi in the
west to Thar Parkar in SE Pakistan.
Felis chaus kelaarti Pocock, 1939; 300.
Type locality: Cheddikulam, N.P., Ceylon [= Sri Lanka].
Holotype: BMNH 1932.2.1.58 young male skull and skin.
Distribution: Sri Lanka and S India, south of the Kistna River up to
about 1500 metres.
Felis chaus oxiana Heptner, 1969; 1259.
Type locality: “Tigrovaya Balka” Nature Reserve in the lower Vakhsh
flow (tributary of the Amu Darya river) [Tadjikistan].
Holotype: ZMMU S-77271 adult male skull and skin.
Distribution: Turkestan (= C Asia).
© J. Tiwari
jungle cat
Felis chaus affinis Gray, 1830; pl. 3.
Type locality: Gangootri, Tehri Garwhal, N India.
Syntypes: BMNH 1837.6.10.40 and BMNH 1837.6.10.41 skins and
skulls.
Distribution: The Himalayas from Kashmir to Sikkim and probably the
Naga Hills, Assam, at altitudes ranging from about 300-2300 metres
or more.
Felis chaus kutas Pearson, 1832; 75.
Type locality: Midnapore, Bengal, c.70 miles W of Calcutta.
Holotype: Museum of the Asiatic Society Bengal mounted skin; lost?
Distribution: N Peninsular India from Bengal to Cutch and ranging from
460 metres in Darbhanga north of Ganges to 46 metres in Cutch.
Felis chaus fulvidina Thomas, 1928; 834.
Type locality: Originally given as “Kampong Tomb, Annam”, but this
was an error. Should be “Komphong Thom, Cambodia” (Duckworth et
al. 2005).
Type: BMNH 1928.7.1.36 skin.
Distribution: Cambodia and probably the range of the species in Viet-
nam, Lao PDR, Thailand and Myanmar (Burma), up to 1500 meters in
the Chin Hills.
Felis chaus maimanah Zukowsky, 1914; 139.
Type locality: Maimana, Afghanistan.
Holotype: Skin in Indian Museum, Kolkata, now lost (see Heptner &
Sludskii 1972; 328).
Distribution: Afghanistan.
N.B. Heptner (1969) and Heptner & Sludskii (1972) identify this taxon
as F. lybica caudata=ornata.
CATnews Special Issue 11 Winter 2017
12
In addition the following subspecies is sometimes recognised from
southern India, but would be included in kelaarti above:
Felis chaus valbalala Deraniyagala, 1955: 201.
Type locality: Karnool, [S India].
Holotype: BMNH 1932.2.1.58, adult male skin and skull.
Distribution: S India, S of the Kistna River.
Discussion
Until recently there had been no morphological or molecular study of
geographical variation in jungle cats. Mukherjee & Groves (2007) ex-
amined the skull morphometrics of jungle cats from throughout their
geographical range except SE Asia. They found that the skulls of west-
ern cats were much larger than those of eastern cats, such that all In-
dian populations were similar, but distinguishable from western ones.
Mukherjee et al. (2010) examined variation in mitochondrial genes
NADH5 and cytochrome b in Indian populations and found some de-
gree of substructuring between northern and southern populations,
but this was very recent and probably not sufficient to support subspe-
cies distinctions. On the basis of these two studies it might be possible
to infer two subspecies, with a possible additional subspecies in SE
Asia, which has so far not been examined in detail:
Felis chaus chaus Schreber, 1777 (incl. oxiana, nilotica, furax).
Distribution: Egypt and the Middle East to Turkestan, Uzbekistan, Ka-
zakhstan and Afghanistan.
Felis chaus affinis Gray, 1830 (incl. prateri, kutas).
Distribution: East Afghanistan, Indian subcontinent and Sri Lanka.
Felis chaus fulvidina Thomas, 1928.
Distribution: SE Asia, possibly including China.
Groves (pers. comm.) states that there are external characters as well
as craniodental characters (Mukherjee & Groves 2007), which strongly
distinguish those from the west (more or less, Iran westward) from
those from the east, which he would be inclined to separate specifi-
cally. Their habitat requirements seem to be different as well – the
western ones are riverine specialists, whereas the eastern ones are
much more evenly spread.
A comprehensive phylogeographical study is required to understand
better geographical variation in Felis chaus.
Subspecies Morphology Molecular Biogeography Certainty Comments
Felis chaus chaus ++ ++ ++
Felis chaus oxiana + o + Synonym of chaus?
Felis chaus maimanah + o Synonym of chaus?
Felis chaus nilotica + o +
Felis chaus furax + o + Synonym of chaus?
Felis chaus affinis + + +
Felis chaus prateri + - o Synonym of affinis?
Felis chaus kutas + - o Synonym of affinis?
Felis chaus kelaarti + o o Synonym of affinis?
Felis chaus valbalala + - + Synonym of affinis?
Felis chaus fulvidina + o +
F. c. chaus
F. c. affinis F. c. fulvidana
Distribution of tentative subspecies of the
jungle cat. Borders between subspecies are
speculative.
jungle cat
A new taxonomy of the Felidae
13
References
Deraniyagala P. E. P. 1955. A new subspecies jungle cat from South India. Spo-
lia Zeylanica 27, 291.
Duckworth J. W., Poole C. M., Tizard R. J., Walston J. L. & Timms R. J. 2005.
The jungle cat Felis chaus in Indochina: A threatened population of a wide-
spread and adaptable species. Biodiversity & Conservation 14, 1263-1280.
Fischer von Waldheim, G. 1817. Adversaria Zoologica. Fasciculus primus.
Quaedam ad Mammalium systema et genera illustranda. Mémoires de la
Société impériale des naturalistes de Moscou 5, 357-446 [=428].
Gray J. E. 1830. Illustrations of Indian Zoology; chiefly selected form the col-
lection of Major-General Hardwicke, vol 1: pl. 3 London: Treuttel, Wurtz,
Treuttel, Jun. and Richter.
Gueldenstaedt A. I. 1776. Chaus, animal feli affine descriptum. Novi commen-
tarii Academiae Scientiarum Imperialis Petropolitanae 20, 483-500.
Heptner V. G. 1969. On systematics and nomenclature of Palearctic cats. Zoo-
logicheskii Zhurnal 48, 1258-1260.
Heptner V. G. & Sludskii A. A. 1972. Mammals of the Soviet Union. Vol. II,
part 2. Carnivora (hyaenas and cats). Moscow: Vysshaya Shkola. [English
translation published in 1992, Smithsonian Institution Libraries, Washing-
ton D.C.].
Mukherjee S. & Groves C. 2007. Geographic variation in jungle cat (Felis chaus
Schreber, 1777) (Mammalia, Carnivora, Felidae) body size: Is competition
responsible? Biological Journal of the Linnean Society 92, 63-172.
black-footed cat
Mukherjee S., Krishnan A., Tamma K., Home C. R. N., Joseph S., Das A. &
Ramakrishnan U. 2010. Ecology driving genetic variation: A comparative
phylogeography of jungle cat (Felis chaus) and leopard cat (Prionailurus
bengalensis) in India. PLoS ONE 5(10): e13724.
Pearson J. 1832. Proceedings of the Societies. Journal of the Asiatic Society
of Bengal 1, 75.
Pocock R. I. 1939. Mammals of British India, vol. 1. Taylor and Francis, London.
Pocock R. I. 1951. Catalogue of the genus Felis. British Museum (Natural His-
tory), London.
Schreber J. C. D. 1777a. Die Säugthiere in Abbildungen nach der Natur mit
Beschreibungen, vol. 3(24), 409-424. Wolfgang Walther, Erlangen.
Schreber J. C. D. 1777b. Die Säugthiere in Abbildungen nach der Natur mit
Beschreibungen, vol. 3(25), 425-440. Wolfgang Walther, Erlangen.
Thomas O. 1928. The Delacour exploration of French Indo-China – Mammals.
III. Mammals collected during winter 1927-28. Proceedings of the Zoologi-
cal Society of London 98, 831-841.
Wozencraft W. C. 2005. Order Carnivora. In Mammal species of the world. A
taxonomic and geographic reference (3rd ed.). Wilson D. E. & Reeder D. M.
(Eds). Johns Hopkins University Press, Baltimore, pp. 532-628.
de Winton W. E. 1898. Felis chaus and its allies, with descriptions of new
subspecies. Annals and Magazine of Natural History (7th series) 2, 291-294.
Zukowsky L. 1914. Drei neue Kleinkatzenrassen aus Westasien. Archiv für
Naturgeschichte Berlin 80, 124-141.
© A. Sliwa
Felis nigripes
E: Black-footed cat; F: Chat à pieds noirs; G: Schwartzfußkatze; Sp:
Gato patinegro, gato de pies negros.
Felis nigripes is typically divided into two subspecies (Wozencraft
2005):
Felis nigripes nigripes Burchell, 1824; 592.
Type locality: the town of Litákun, [= Dithakong, near Kuruman, N Cape
Province, South Africa].
Holotype: Incomplete skin seen by Burchell.
Distribution: Kalahari of Botswana, Namibia and northern Cape.
Province, South Africa (Sliwa 2013).
Distinguishing characters: Lighter in colour, tawny or off-white; bands
running from nape often broken into spots or short stripes; spots
brownish black or tawny (Sliwa 2013).
Felis nigripes thomasi Shortridge, 1931; 119.
Type locality: Thorn Kloof (Carlisle Bridge), C.P. (= Cape Province), [East-
ern Cape, South Africa].
Holotype: Albany Museum no. 6333 adult male skin and skeleton.
Distribution: Karoo of central and southern South Africa (Sliwa 2013).
Distinguishing characters: Cinnamon-buff; bands from nape strongly
developed and run unbroken to base of tail; three distinctive throat
rings; spots are satiny black (Sliwa 2013).
Discussion
Pocock (1951) pointed out that variation in the pelage coloration
of skins from the nominate race suggests the differences between
these two subspecies are less than described. The supposed bio-
geographical barrier of the Orange River is also not consistent with
the distribution of the two supposed morphological types. Sliwa
(2013) considered the two putative subspecies to represent the
ends of a cline, with animals of the appearance of both subspe-
cies occurring near Kimberley. Therefore, it seems likely that these
subspecies are not valid and that this is a monotypic species, with
some clinal variation:
Felis nigripes Burchell, 1824.
Distribution: Southern Africa.
CATnews Special Issue 11 Winter 2017
14
sand cat
Subspecies Morphology Molecular Biogeography Certainty Comments
Felis nigripes nigripes ++ ++ ++
Felis nigripes thomasi + o o Included in nigripes
Felis margarita
E: Sand cat; F: Chat des sables; G: Sandkatze, Saharakatze; Sp: Gato
de las arenas, gato del Sahara.
Typically four subspecies are recognised (Wozencraft 2005):
Felis margarita margarita Loche, 1858; 49, pl.1.
Type locality: environs de Négonça (Sahara) [Algeria].
Holotype: No longer survives.
Distribution: North Africa.
Felis margarita harrisoni Hemmer, Grubb and Groves, 1976; 301.
Type locality: northern edge of Umm as Samin, Oman, 21°55’ N /
55° 30’ E.
Holotype: BMNH 1977.430 adult male skull and skin.
Distribution: Arabian Peninsula, Sinai, Israel.
Felis margarita thinobia (Ognev, 1927; 356).
Type locality: Repetek, Turkmenistan.
Holotype: ZMMU S-14226 adult male skull and skin.
Distribution: The deserts of Karakum and Kizilkum, Central Asia, and
Iran (Lay et al. 1970).
Felis margarita scheffeli Hemmer, 1974; 32.
Type locality: Nushki-Wüste, Westpakistan [Nuski Desert, W Paki-
stan].
Holotype: SMF 38326 skull, skeleton and skin of an adult female im-
ported alive in 1970-72.
Distribution: Pakistan.
Discussion
To date there have been no phylogeographical studies. There appear
to be differences in pelage coloration and markings and skull size be-
tween North African sand cats and those from Pakistan (A. Kitchener,
pers. obs.). The pelages of Arabian sand cats resemble that of some
North African sand cats, and others resemble those of Pakistani and
Turkmenian sand cats, which resemble each other, although the former
tend to be greyer and the latter yellower. Preliminary genetic data (H.
Senn, pers. comm.) support the distinctiveness of North African sand
cats, albeit weakly. Therefore, it is possible that there are only two
subspecies:
F. margarita margarita
F. margarita thinobia
Distribution of tentative subspecies of sand cat. Borders between
subspecies are speculative.
Felis margarita margarita Loche, 1858.
Distribution: North Africa.
Distinguishing characters: Smaller size with yellowish pelage and of-
ten marked with spots and stripes.
Felis margarita thinobia (Ognev, 1927).
Distribution: Southwest Asia and the Arabian Peninsula.
Distinguishing characters: Large size, greyer pelage with fewer mark-
ings.
© A. Sliwa
References
Burchell W. J. 1824. Travels in the interior of southern Africa, vol. 2. Longman,
Hurst, Rees, Orme, Brown and Green, London.
Pocock R. I. 1951. Catalogue of the genus Felis. British Museum (Natural His-
tory), London.
Shortridge G. C. 1931. Felis (Microfelis) nigripes thomasi ssp. nov. Records of
the Albany Museum 4, 119-121.
Sliwa A. 2013. Felis nigripes Black-footed cat. In Mammals of Africa, vol. 5.
Kingdon J. & Hoffmann M. (Eds). Bloomsbury, London pp. 203-206.
Wozencraft W. C. 2005. Order Carnivora. In Mammal species of the world. A
taxonomic and geographic reference (3rd ed.). Wilson D. E. & Reeder D. M.
(Eds). Johns Hopkins University Press, Baltimore, pp. 532-628.
A new taxonomy of the Felidae
15
Chinese mountain cat
Subspecies Morphology Molecular Biogeography Certainty Comments
Felis margarita margarita ++ ++ ++
Felis margarita harrisoni + o + Synonym of thinobia?
Felis margarita scheffeli + o +
Felis margarita thinobia ++ o ++ Probably distinct and includes
scheffeli
References
Hemmer H. 1974. Felis margarita scheffeli, eine neue Sandkatzen-Unterart aus
der Nushki-Wüste, Pakistan. Senckenbergia biologica 55, 29-34.
Hemmer H., Grubb P. & Groves C. P. 1975. Notes on the sand cat, Felis marga-
rita Loche, 1858. Zeitschrift für Säugetierkunde 41, 286-303.
Lay D. M., Anderson J. A. W. & Hassinger J. D. 1970. New records of small
mammals from West Pakistan and Iran. Mammalia 34, 98-106.
Loche V. 1858. Description d’une nouvelle espèce de chat. Revue et magasin
de zoologie pure et appliquée, série 2, 10, 49-50.
Ognev S. I. 1927. A new genus and species of cat from the Trans-Caspian re-
gion. Ezhegodnik Zoologicheskogo Muzeya Imperatorskoi Akademii Nauk
27, 356-362.
Schauenberg P. 1974. Données nouvelles sur le chat des sables Felis margarita
Loche, 1858. Revue Suisse de Zoologie 81, 949-969.
Wozencraft W. C. 2005. Order Carnivora. In Mammal species of the world. A
taxonomic and geographic reference (3rd ed.). Wilson D. E. & Reeder D. M.
(Eds). Johns Hopkins University Press, Baltimore, pp. 532-628.
Felis bieti
E: Chinese steppe cat, Chinese mountain cat, Chinese desert cat; F:
Chat de Biet; G: Graukatze, Gobikatze; Sp: Gato de Biet, gato desierto
de China.
Various subspecies have been described for Felis bieti, including (fol-
lowing Pocock 1951):
Felis bieti bieti Milne-Edwards, 1892; 671.
Type locality: Tengri-Nor à Batang, restricted to the environs de Ton-
golo et de Ta-tsien-lou [Central Sichuan, China] by Pousargues (1898;
357).
Diagnosis: Ears match coloration of back with red ear tufts, yellowish
grey pelage, faint transverse stripes, whitish ventrally.
Holotype: MNHN-ZM-MO-1891-391 mounted skin and skull.
Distribution: Provinces of Qinghai, Sichuan and possibly Gansu, China
(He et al. 2004, Webb et al. 2016).
Felis bieti chutuchta Birula, 1917; 1.
Type locality: Nor locality, Goizso area, Gobi Desert, [China].
Holotype: Adult female skull (ZIN 9377) and skin (ZIN 9880).
Distribution: Gobi Desert, China.
Distinguishing characters: Redder and more distinctly striped than bieti.
Felis bieti vellerosa Pocock, 1943; 172.
Type locality: near Yu-Lin-fu, 4000 ft, on the borders of Ordos and
NE Shensi [near Yulin, 1200 metres, on the borders of Ordos and NE
Shaanxi, China].
Holotype: BMNH 1909.1.1.11 skin.
Distribution: Known only from type locality.
Distinguishing characters: Bicoloured ears (grey proximally, black dis-
tally) with long black ear tuft, black genal stripes, legs striped.
Discussion
There has been no recent taxonomic study of this species since Groves
(1980), although Driscoll et al. (2007) showed that Felis bieti was basal
to Felis silvestris (sensu lato) according to mtDNA, but a sister taxon to
Felis lybica ornata from microsatellites, which suggests that Felis bieti
may have an ancient hybrid origin, possibly during the last glaciation
when the distribution of F. l. ornata was apparently restricted to a very
small area in Central Asia (Kitchener & Rees 2009). F. bieti is morpho-
logically distinct and is supposedly sympatric with F. l. ornata, which
would also preclude its recognition as a subspecies of F. silvestris/
lybica. However, C. Driscoll (pers. obs.) maintains this species as a
subspecies within F. silvestris (sensu lato).
The skull of putative subspecies chutuchta is similar to that of lybica
(Groves 1980, A. Abramov, pers. obs.) and doubt has also been cast on
vellerosa, although A. Kitchener (pers. obs.) believes that this speci-
men is a trade skin outside its geographical distribution or possibly a
Felis chaus, while Groves (1980) believes it to be F. catus.
Therefore, given its restricted distribution and distinct morphology, Fe-
lis bieti is recognised here as a monotypic species.
© A. Guillemont
CATnews Special Issue 11 Winter 2017
16
European wildcat
Felis silvestris
E: European wildcat, Caucasian wildcat; F: Chat forestier, chat sau-
vage d’Europe, chat sylvestre; G: Europäische Wildkatze, Waldkatze;
Sp: Gato montés, gato silvestre.
Felis silvestris, as defined here, includes only the forest cats of Eu-
rope. Many subspecies have been described, but there are no recent
morphological and molecular studies of geographical variation in
Europe and beyond. Wozencraft (2005) recognised the following sub-
species:
Felis silvestris silvestris Schreber, 1777; 397 – conserved by Opin-
ion 465 of the International Commission of Zoological Nomenclature
(International Commission on Zoological Nomenclature, 1957).
Locality from where the species was described: Unknown but fixed as
“vielleicht Nordfrankreich” [perhaps N France] by Haltenorth, 1953 and
“Germany” by Pocock (1951).
Type: None designated.
Distribution: Mainland Europe from Spain to eastern Europe, including
Bulgaria, Rumania, southern Poland, western Russia.
Felis silvestris caucasica Satunin, 1905; 154.
Type locality: Borjomi, Georgia, Caucasus.
Type: Museum Tiflis [Georgian National Museum] female.
Distribution: Caucasus, Turkey.
Felis silvestris grampia Miller, 1907; 396.
Type locality: Invermoriston District, Inverness, Scotland.
Type: BMNH 1904.1.25.3 male skin and skull.
Distribution: N and C Scotland, formerly all of Britain.
Discussion
There is a cline in pelage flank stripes in Europe from distinctly striped
animals in the west to faintly striped animals in the east (A. Kitchener,
pers. comm.). This may reflect divergence in Pleistocene refugia in
southern Europe followed by recolonisation and introgression follow-
ing the end of the last glacial.
Based on current geographical isolation, it seems likely that there are
two subspecies of F. silvestris:
Felis silvestris silvestris Schreber, 1777.
Distribution: Europe, including Scotland, Sicily and Crete.
Felis silvestris caucasica Satunin, 1905.
Distribution: Caucasus, Turkey.
Subspecies Morphology Molecular Biogeography Certainty Comments
Felis bieti bieti ++ ++ ++
Felis bieti chutchta ++ o o This is probably a form of Felis lybica
Felis bieti vellerosa ++ o o This may be Felis chaus or F. catus
© KORA
Felis bieti Milne-Edwards, 1892.
Distribution: Provinces of Qinghai, Sichuan and possibly Gansu, China.
References
Birula A. 1917. De Felibus asiaticis duabus novis. Annuaire du Musée Zo-
ologique de l’Academie Impériale de St. Pétersbourg 21: Itinéraires: Nou-
velles et Faites Divers: I-II.
Driscoll C. A., Menotti-Raymond M., Roca A. L., Hupe K., Johnson W. E., Gef-
fen E., Harley E. H., Delibes M., Pontier D., Kitchener A. C., Yamaguchi
N., O’Brien S. J. & Macdonald D. W. 2007. The Near Eastern origin of cat
domestication. Science 317, 519-523.
Groves C. P. 1980. The Chinese Mountain cat. Carnivore 3, 35-41.
He L., Garcia-Perea R., Li M. & Wei F. 2004. Distribution and conservation
status of the endemic Chinese mountain cat, Felis bieti. Oryx 38, 55-61.
Kitchener A. C. & Rees E. E. 2009. Modelling the dynamic biogeography of the
wildcat: Implications for taxonomy and conservation. Journal of Zoology
London 279, 144-155.
Milne-Edwards A. 1892. Observations sur les mammifères du Thibet. Revue
Génerale des Sciences Pures et Appliquées 3, 670-672.
Pocock R. I. 1943. A new desert cat (Felis) from North China. Proceedings of
the Zoological Society of London B 113, 172-175.
Pocock R. I. 1951. Catalogue of the genus Felis. British Museum (Natural His-
tory), London.
Webb R., Francis S., Telfer P. & Guillemont A. 2016. Chinese mountain cat and
Pallas’s cat co-existing on the Tibetn plateau in Sichuan. Cat News 63,
31-33.
A new taxonomy of the Felidae
17
Subspecies Morphology Molecular Biogeography Certainty Comments
Felis silvestris silvestris ++ ++ ++
Felis silvestris grampia + + + Doubtfully distinct
Felis silvestris caucasica ++ o ++ Probably distinct
steppe and bush cats of Africa and Asia
Distribution of tentative subspecies of European wildcat. Borders
between subspecies are speculative.
However, C. Driscoll (pers. comm.) retains lybica, cafra, ornata and bieti
within Felis silvestris as subspecies following Discroll et al. (2007), while
C. Groves (pers. comm.) regards all these as valid phylogenetic species.
References
Driscoll C. A., Menotti-Raymond M., Roca A. L., Hupe K., Johnson W. E., Gef-
fen E., Harley E. H., Delibes M., Pontier D., Kitchener A. C., Yamaguchi
N., O’Brien S. J. & Macdonald D. W. 2007. The Near Eastern origin of cat
domestication. Science 317, 519-523.
International Commission on Zoological Nomenclature. 1957. Opinion 465.
Variation under the Plenary Powers of the specific name silvestris Schre-
ber, [1777], as published in the combination Felis (catus) silvestris, for the
European wild cat (Class Mammalia). Opinions and declarations rendered
by the International Commission on Zoological Nomenclature 16, 43-52.
Miller G. S. 1907. Some new European Insectivora and Carnivora. Annals and
Magazine of Natural History (7th series) 20, 389-398.
Pocock R. I. 1951. Catalogue of the genus Felis. British Museum (Natural His-
tory), London.
Satunin K. A. 1905. Die Säugetiere des Talyschgebietes und der Mughan-
steppe. Mlekopitayushchie Talysha i Mugani. Mitteilungen des Kaukasis-
chen Museums - Izvestiya Kavkazskago Muzeya 2, 87-402.
Schreber J. C. D. 1777. Die Säugthiere in Abbildungen nach der Natur mit
Beschreibungen, vol. 3(23). Wolfgang Walther, Erlangen.
Wozencraft W. C. 2005. Order Carnivora. In Mammal species of the world. A
taxonomic and geographic reference (3rd ed.). Wilson D. E. & Reeder D. M.
(Eds). Johns Hopkins University Press, Baltimore, pp. 532-628.
Felis lybica
E: African wildcat, Indian desert cat; F: Chat ganté, chat sauvage
d’Afrique, chat orné, chat sauvage d’Asie ; G: Nubische Falbkatze, Asi-
atische Wildkatze; Sp: Gato silvestre, gato montés.
Felis lybica, as defined here, includes the steppe and bush cats of Af-
rica and Asia. Very many subspecies have been described throughout
the extensive geographical distribution of this species (Pocock 1951;
distributions of subspecies below are taken from here). The subspe-
cies listed by Wozencraft (2005) are as follows:
Felis lybica lybica Forster, 1780; 313.
Type locality: in der Gegend der alten Stadt Kapsa [= in the region of
the ancient town of Gafsa, Tunisia].
Holotype: Based on “chat du desert from Capsa, Lybie” of Buffon
(1776; 233) based on unpublished correspondence from Bruce.
Distribution: Semi-deserts of North Africa from Morocco, Algeria and
Tunisia to Egypt and up the Nile to Sudan and eastwards to Suakin and
Massowah and the E coast of Sinai.
Distinguishing characters: Light, buff or sandy coloration, pale with
reddish spots ventrally, ochreous ears, whitish face.
Felis lybica ocreata Gmelin, 1791; 27 and 79.
Type locality: Ras el Feel, Abyssinia [= Ethiopia].
Holotype: Based on the booted lynx of Bruce (1790; 146).
Distribution: Ethiopia.
Distinguishing characters: Similar to lybica, but grey with more black
speckling and reddish or yellow wash.
Felis lybica haussa Thomas and Hinton, 1921; 2.
Type locality: Zinder [about 300 miles south of Aïr, Niger].
Holotype: BMNH 1921.2.11.16 male skin and skull.
Distribution: Zinder, Niger and Franiso, near Kano, Nigeria.
Distinguishing characters: Similar to lybica, but smaller skull.
Felis lybica foxi Pocock, 1944a; 71.
Type locality: Kabwir, 7000 ft. on the slopes of the Panyam Plateau,
[Bauchi Province, N Nigeria].
Holotype: BMNH 1912.11.7.5 male skin.
Distribution: Panyam Plateau, N Nigeria.
Distinguishing characters: Darker than haussa, similar to sarda, with
reddish face, but less thick fur, spinal area and crown less black, and
speckling on flanks buffy.
Felis lybica rubida Schwann, 1904; 422.
Type locality: Monbuttu [Belgian Congo = Democratic Republic of
Congo].
Holotype: BMNH 1887.12.1.6 young male skin and skull.
Distribution: Democratic Republic of Congo.
Distinguishing characters: Pale brown or cinnamon coloration, with
almost no black speckling except dorsal line, spotted.
F. s. silvestris
F. s. caucasica
CATnews Special Issue 11 Winter 2017
18
steppe and bush cats of Africa and Asia
Felis lybica ugandae Schwann, 1904; 424.
Type locality: Mulema, Uganda.
Holotype: BMNH 1903.11.7.8 young adult male skin and skull.
Distribution: Mongalla in South Sudan, Garamba in Democratic Re-
public of Congo, Uganda, Kenya and possibly Tanzania.
Distinguishing characters: More black speckling than rubida, duller
coloration, striping not always present.
Felis lybica tristrami Pocock, 1944b; 125.
Type locality: Ghor Seisaban, Moab [Palestine = Israel].
Holotype: BMNH 1893.1.29.3 female skin and skull.
Distribution: Israel, Jordan, Syria, Lebanon, W and S Arabia.
Distinguishing characters: Similar to sarda, but paler, less luxurious
pelage, less black on back of metatarsus.
Felis lybica iraki Cheesman, 1921; 331.
Type locality: Koweit, Arabia [= Kuwait].
Holotype: BMNH 1920.1.19.2 male skin and skull.
Distribution: Kuwait and Iraq.
Distinguishing characters: Similar to tristrami, but pelage tawnier
above, spinal band undifferentiated, face and feet whiter.
Felis lybica gordoni Harrison, 1968; 283.
Type locality: Wadi Suwera 6 miles west of Sohar, Batinah coast of
Oman.
Holotype: BMNH 1968.608 female skin and skull.
Distribution: Oman and UAE.
Distinguishing characters: Compared with other Arabian lybica, very
pale grey, lacking olivaceous tint of tristrami, brown spinal stripe from
shoulders.
Felis lybica nesterovi Birula, 1917; 1.
Type locality: Nachr-Chazasch, Mesopotamia [= Iraq].
Holotype: Adult female ZIN 9374 (skull), ZIN 27643 (skin).
Distribution: Iraq and S Iran.
Distinguishing characters: Similar to ornata, but longer fur.
Felis lybica reyi Lavauden, 1929; 1023.
Type locality: Forêt d’Aunes des bords de la lagune de Biguglia (Sud
de Bastia) [Corsica].
Holotype: MNHN-ZM-MO-1932-3806 female skin and skull.
Distribution: Corsica.
Distinguishing characters: Compared with sarda, darker pelage, short-
er tail, and backs of ears dark brown without a trace of red.
Felis silvestris cretensis Haltenorth, 1953; 29.
Type locality: Kanea auf Kreta [= Chania, Crete, Greece].
Holotype: BMNH 1905.12.2.14 skin.
Distribution: Crete.
Distinguishing characters: Similar to lybica, but with tail similar to
silvestris.
Comments: C. Groves (pers. obs.) considers that this is probably Felis
catus. Felis silvestris occurs also on Crete (Matschei 2015, A. Kitch-
ener, pers. obs.). It could be a hybrid between F. silvestris, F. lybica
and/or F. catus.
Felis lybica jordansi Schwarz, 1930; 223.
Type locality: Santa Margarita, Mallorca, Balearen [Majorca, Spain].
Holotype: ZFMK 83.186 male skull and skin.
Distribution: Balearic Islands.
Distinguishing characters: More strongly striped with brighter legs
than lybica. C. Groves (pers. obs.) has measured the cranial volume of
the holotype and it is Felis catus.
Felis lybica cafra Desmarest, 1822; 540.
Type locality: Kaffraria [South Africa].
Syntypes: MNHN-ZM-MO-2002-321 and MNHN-ZM-MO-2002-322
mounted skins (skulls inside).
Distribution: Originally S of the Orange River from Cape of Good Hope
and Little Namaqualand in W to Eastern Cape and KawZulu Natal and
N to Transvaal. Now southern Africa.
Distinguishing characters: Similar to ugandae, but occurs in two colour
phases (iron grey with black and whitish speckling, and tawny grey
with less speckling), both of which have thicker coat, and development
of black pigment on fore legs.
Felis lybica mellandi Schwann, 1904; 423.
Type locality: Mpika, NE Rhodesia [= Muchinga Provine, Zambia].
Holotype: BMNH 1904.3.11.2 skin.
Distribution: Malawi, Zambia and S Democratic Republic of Congo.
Distinguishing characters: Similar to ugandae, but coloration above more
uniform, brighter coloration on ears, and faint or absent striping on flanks.
Felis lybica griselda Thomas, 1926; 180.
Type locality: Fifty miles south of Dombe Grande, Benguella, Angola.
Holotype: BMNH 1925.5.16.1 skin.
Distribution: From S Angola and Namibia eastwards into Botswana.
Distinguishing characters: Similar to cafra, but paler, brighter ochreous
ears, paler pelage, and coat pattern less distinct.
Felis lybica ornata Gray, 1830; pl.2.
Type locality: Nusserabad, Rajputana [India].
Holotype: BMNH 1848.8.14.3 skin.
Distribution: W and C India S of the Ganges.
Distinguishing characters: Greyish sandy cat covered in irregular black
or brown spots.
Felis lybica caudata (Gray, 1874; 31).
Type locality: “Cocan”, Bokhara; near the river Dyanan. Cocan, or Kho-
kan, is situated on the Sir Daria; and I suppose that the Dyanan is a
branch of the river Sir, which falls into the sea of Aral (Gray 1874:
31). Birula (1912; 226) clarified the type locality as “Jana Darya River,
which is the south branch of Syr Darya River in its lower reach”, nei-
ther Bukhara nor Kokand.
Holotype: BMNH 1873.7.22.12 skin and skull.
Distribution: Turkestan (C Asia) as far E as Tian Shan and S into Iran
and Afghanistan.
Distinguishing characters: Similar to ornata, but larger and more luxu-
riant winter pelage, larger teeth.
Felis lybica chutuchta Birula, 1917; 1.
Type locality: Nor locality, Goizso area, Gobi Desert [China].
Holotype: Adult female ZIN 9377 (skull), ZIN 9880 (skin).
Distribution: Gobi Desert, China.
Distingushing characters: Reddish body and ears with distinct trans-
verse stripes.
A new taxonomy of the Felidae
19
steppe and bush cats of Africa and Asia
Discussion
Driscoll et al. (2007) identified three distinct clades within this species,
which we identify tentatively as subspecies. However, it should be
noted that samples were not available from some key areas through-
out the geographical range, e.g. much of North, West and East Africa.
In contrast, C. Groves (pers. comm.) recognises the three clades as
representing distinct species, while C. Driscoll (pers. comm.) includes
these, silvestris and bieti within Felis silvestris. We tentatively identify
the following subspecies within F. lybica:
Felis lybica lybica Forster, 1780.
Distribution: E, W and N Africa, Arabian Peninsula, Middle East, Cor-
sica, Sardinia and Crete; probably intergrades with ornata in Iraq.
F. l. lybica in Saudi Arabia (Photo C. Barichirvy & T. Wacher).
Felis lybica cafra Desmarest, 1822.
Distribution: Southern Africa; exact boundary with lybica uncertain,
but may lie in Mozambique or Tanzania.
F. l. cafra in Botswana (Photo P. Meier).
Felis lybica ornata Gray, 1830.
Distribution: SW and C Asia, Afghanistan, Pakistan, India, Mongolia
and China.
Distinguishing characters: Light-coloured pelage with black spots.
F. l. ornata in India (Photo D. Khandal).
References
Birula A. 1912. Materialy po sistematike i geograficheskomu rasprostraneniyu
mlekopitayushchikh. III. Carnivora, sobrannye N. A. Zarudnym v Persii v
1896, 1898, 1900-1901 i 1903-1904. Ezhegodnik Zoologicheskogo Muzeya
Imperatorskoi Akademii Nauk 27, 219-280.
Birula A. 1917. De Felibus asiaticis duabus novis. Annuaire du Musée Zo-
ologique de l’Academie Impériale de St. Pétersbourg 21: Itinéraires: Nou-
velles et Faites Divers: I-II.
Bruce J. 1790. Select specimens of Natural History collected in travels to dis-
cover the source of the Nile in Egypt, Arabia, Abyssinia and Nubia. Vol V.
London: Robinson and Robinson.
Buffon G.-L. L., Comte de 1776. Addition à l’article du lynx & à celui du caracal,
volume IX , pages 231 & 262. In Histoire naturelle générale et particu-
lière. Servant de suite à l’histoire des animaux quadrupeds. Supplement,
Tome troisième. Buffon, G.-L. L., Comte de & Daubenton L.-J.-M. (Eds). De
l’Imprimière royale, Paris, pp. 229-233.
Cheesman R. E. 1921. Report on the mammals of Mesopotamia collected by
members of the Mesopotamian Expeditionary Force, 1915 to 1919. Journal
of the Bombay Natural History Society 27, 323-346.
Desmarest A. G. 1822. Mammalogie ou description des espèces de mammifères.
Encyclopèdie Méthodique. Mammiferes Supplement. Agasse, Paris.
Driscoll C. A., Menotti-Raymond M., Roca A. L., Hupe K., Johnson W. E., Gef-
fen E., Harley E. H., Delibes M., Pontier D., Kitchener A. C., Yamaguchi
N., O’Brien S. J. & Macdonald D. W. 2007. The Near Eastern origin of cat
domestication. Science 317, 519-523.
Forster G. R. 1780. Herrn von Buffon’s Naturgeschichte der vierfüssigen Thiere.
Mit Vermehrungen, aus dem Französischen übersetzt, vol. 6. J. Pauli, Ber-
lin.
Gmelin J. F. 1791. Anmerkungen zu James Bruce Reise nach Abyssinien. In An-
hang zu James Bruce Reisen in das Innere von Africa, nach Abyssinien an
die Quellen des Nils – welcher Berichtigungen und Zusäzze aus der Natur-
geschichte von J. F. Gmelin und aus der alten, besonders orientalischen
Litteratur von verschiedenen Gelehrten enthält, vol. 2, pp. 1-38. Rinteln:
In der Expedition der theologischen Annalen, Lepizig: In Commission bey
Joh. Ambrosius Barth.
Gray J. E. 1830. Illustrations of Indian Zoology; chiefly selected from the col-
lection of Major-General Hardwicke, vol 1: pl. 3. Treuttel, Wurtz, Treuttel,
Jun. and Richter, London.
Gray J. E. 1874. On the steppe-cat of Bokhara (Chaus caudatus). Proceedings
of the Zoological Society of London 42, 31-33.
Haltenorth T. 1953. Die Wildkatzen der alten Welt. Eine Übersicht über die
Gattung Felis. Geest & Portig, Leipzig.
Harrison D. L. 1968. The mammals of Arabia: Carnivora, Hyracoidea, Artiodac-
tyla, vol. 2. Ernest Benn Ltd, London.
Lavauden L. 1929. Sur le chat sauvage de la Corse. Comptes Rendus Hebdo-
madaires des Séances de l’Académie des Sciences, Paris 189, 1023-1024.
Pocock R. I. 1944a. The races of the North African wild cat. Proceedings of the
Zoological Society of London 114, 65-73.
Pocock R. I. 1944b. The South African races of the wild cat (Felis lybica). Pro-
ceedings of the Zoological Society of London 114, 297-301.
Pocock R. I 1944c. The wild cat (Felis lybica) of Palestine. Annals and Maga-
zine of Natural History (11th series) 11, 125-130.
Pocock R. I. 1951. Catalogue of the genus Felis. British Museum (Natural His-
tory), London.
Schwann H. 1904. On Felis ocreata, better known as Felis caligata, and its sub-
species. Annals and Magazine of Natural History (7th series) 13, 421-426.
Schwarz E. 1930. Die Wildkatze der Balearen. Zoologischer Anzeiger 91, 223-
224.
CATnews Special Issue 11 Winter 2017
20
steppe and bush cats of Africa and Asia
Distribution of tentative subspe-
cies of steppe and bush cats of
Africa and Asia. Borders between
subspecies are speculative.
Thomas O 1926. Some new African Mammalia. Annals and Magazine of Natu-
ral History (9th series) 17, 180-184.
Thomas O. & Hinton M. A. C. 1921. Captian Angus Buchanan’s Aïr Expedition.
II. On the mammals (other than ruminants) obtained during the expedition
to Aïr (Asben). Novitates Zoologicae 28, 1-13.
Wozencraft W. C. 2005. Order Carnivora. In Mammal species of the world. A
taxonomic and geographic reference (3rd ed.). Wilson D. E. & Reeder D. M.
(Eds). Johns Hopkins University Press, Baltimore, pp. 532-628.
Subspecies Morphology Molecular Biogeography Certainty Comments
Felis lybica lybica ++ ++ ++
Felis lybica ocreata + o + Doubtfully distinct
Felis lybica haussa + o o
Felis lybica foxi + o o
Felis lybica rubida + o o
Felis lybica ugandae + o o
Felis lybica tristrami + o + Doubtfully distinct
Felis lybica iraki + o o
Felis lybica gordoni + o + Doubtfully distinct
Felis lybica nesterovi + o o
Felis lybica reyi + o + Introduced by humans
Felis lybica jordansi + o + Introduced by humans = Felis catus
Felis lybica cretensis + o + Introduced by humans?
Felis lybica cafra ++ ++ ++
Felis lybica mellandi + o o
Felis lybica griselda + o o
Felis lybica ornata ++ ++ ++
Felis lybica caudata + o o
Felis lybica chutuchta + o o
F. l. lybica
F. l. ornata
F. l. cafra
A new taxonomy of the Felidae
21
domestic cat, Pallas‘s cat
Leopard Cat lineage
Genus Otocolobus Brandt, 1842; 38.
Otocolobus manul
E: Pallas’s cat, manul; F: Manul, chat de Pallas; G: Manul; Sp: Gato
manul, gato de Pallas.
Wozencraft (2005) recognised three subspecies of Otocolobus manul:
Otocolobus manul manul (Pallas, 1776; 692).
Locality from where the species was described: Frequens in rupetri-
bus, apricis totius Tatariae Mongoliaeque desertae = Kulusutai,
Borzya District, Chita Province, USSR [= Russia] (Heptner & Sludskii
1972).
Type: None designated.
Distribution: China (Gansu), Mongolia, Kazakhstan, southern Siberia
(Altai, Tuva, Transbaikalia).
Otocolobus manul nigripectus (Hodgson, 1842; 276).
Type locality: from Tibet
Types: Three syntypes, all skins: BMNH 1845.1.8.209; BMNH
1858.6.24.68; BMNH 1858.6.24.112
Distribution: Tibet and Kashmir.
Distinguishing characters: Winter coat silvery-grey with more black
in it, wool paler, head spotted thickly with black, back and tail stripes
more distinct (Pocock 1951).
Otocolobus manul ferrugineus Ognev, 1928; 1013.
Type locality: from mountain ridge of Missanev, Kopet-Dag Mountains,
Turkmenistan.
Holotype: Male skin (ZIN 28013) and skull (ZIN 15065).
Distribution: C Asia (Turkmenistan, Uzbekistan, Tadjikistan), N Iran,
Afghanistan and Baluchistan.
Distinguishing characters: Reddish dorsally, black markings incon-
spicuous or red.
© P. Meier
Felis catus
E: Domestic cat, feral cat; F: Chat domestique; G: Katze; Sp: Gato do-
mestico.
Felis catus Linnaeus, 1758; 42
Locality from where species was first described: Sweden (Pocock 1951).
Type: None designated.
Distribution: Worldwide, except Antarctica.
Discussion
Domesticated mostly from a lineage of Felis lybica lybica from Meso-
potamia (Driscoll et al. 2007). Following Opinion 2027 of the Interna-
tional Commission on Zoological Nomenclature (2003), the domestic
cat is treated as a distinct taxon, Felis catus (Gentry et al. 2004).
References
Driscoll C. A., Menotti-Raymond M., Roca A. L., Hupe K., Johnson W. E., Gef-
fen E., Harley E. H., Delibes M., Pontier D., Kitchener A. C., Yamaguchi
N., O’Brien S. J. & Macdonald D. W. 2007. The Near Eastern origin of cat
domestication. Science 317, 519-523.
Gentry A., Clutton-Brock J. & Groves C. P. 2004. The naming of wild animal
species and their domestic derivatives. Journal of Archaeological Science
31, 645-651.
International Commission on Zoological Nomenclature (2003). Opinion 2027.
Usage of 17 specific names based on wild species which are pre-dated
by or contemporary with those based on domestic animals (Lepidoptera,
Osteichthyes, Mammalia): Conserved. Bulletin of Zoological Nomenclature
60, 81-84.
Linnaeus, C. 1758. Systema Naturae per regna tria naturae, secundum classis,
ordines, genera, species cum characteribus, differentiis, synonymis, locis.
10th edition, vol. 1. Holmiae: Laurentii Salvii.
Pocock R. I. 1951. Catalogue of the genus Felis. London: British Museum
(Natural History).
© KORA
CATnews Special Issue 11 Winter 2017
22
Pallas‘s cat
Distribution of tentative subspecies of Pallas’s cat. Borders between subspecies are speculative.
Subspecies Morphology Molecular Biogeography Certainty Comments
Otocolobus manul manul ++ ++ ++
Otocolobus manul ferrugineus + Doubtfully distinct
Otocolobus manul nigripectus + + Possibly distinct
Discussion
There have been no recent molecular or morphological studies. The
subspecies ferrugineus is variably rufescent (Pocock 1939, A. Kitch-
ener, pers. obs.) and is said to intergrade with typical manul. It seems
likely that this variability in erythrism is explained mostly by simple
Mendelian inheritance, with completely orange animals (homozygous)
and those with a mixture of orange and grey (heterozygous) for the O
gene. Perhaps there is selection for more erythristic forms in the SW
of the species’ range? It is possible that this species is monotypic and
shows clinal variation in pelage coloration.
Pocock (1951) found complete overlap in coloration between skins of all
putative subspecies and it may turn out that variation is largely clinal.
We suggest the tentative recognition of only two subspecies:
Otocolobus manul manul (Pallas, 1776), including ferrugineus.
Distribution: China (Gansu), Mongolia, C Asia and Kazakhstan, S Sibe-
ria, Iran, Afghanistan and Pakistan.
Otocolobus manul nigripectus (Hodgson, 1842).
Distribution: Tibet, Kashmir, Nepal and Bhutan.
Further research is required to understand geographical variation in
Otocolobus manul.
References
Brandt J. F. 1842. Observations sur le manoul (Felis manul Pallas). Bulletin Sci-
entifique. Académie Impériale des Sciences de Saint Petersbourg 9, 37-39.
Heptner V. G. & Sludskii A. A. 1972. Mammals of the Soviet Union. Vol. II,
part 2. Carnivora (hyaenas and cats). Moscow: Vyshshaya Shkola. [English
translation published in 1992, Smithsonian Institution Libraries, Washing-
ton, D.C].
Hodgson B. H. 1842. Notice of the mammals of Tibet, with descriptions and
plates of some new species. Journal of the Asiatic Society of Bengal 11,
275-289; coloured drawing p. 288-289.
Ognev S. 1928. On a new form of the steppe cat from the Transcaspian region.
Comptes Rendus de l’Académie des Sciences de l’URSS, 308-310.
Pallas P. S. 1776. Reise durch verschiedene Provinzen des russischen Reichs,
vol. 3. St. Petersbourg.
Pocock R. I. 1939. Mammals of British India, vol. 1. Taylor and Francis, London.
Wozencraft W. C. 2005. Order Carnivora. In Mammal species of the world. A
taxonomic and geographic reference (3rd ed.). Wilson D. E. & Reeder D. M.
(Eds). Johns Hopkins University Press, Baltimore, pp. 532-628.
O. m. manul
O. m. nigripectus
A new taxonomy of the Felidae
23
Genus Prionailurus Severtzov, 1858; 387.
This genus contains five species.
Prionailurus rubiginosus.
E: Rusty-spotted cat; F: Chat rougâtre, chat rubigineux; G: Rostkatze;
Sp: Gato rubiginosa, gato rojizo.
Wozencraft (2005) recognised two subspecies:
Prionailurus rubiginosus rubiginosus (I. Geoffroy Saint-Hilaire,
1831; 140).
Type locality: “les bois de lataniers qui couvrent une hauteur voisine de
Pondichéry, connue sous le nom de Coteau”.
Holotype: MNHN-ZM-AC-A1791 juvenile incomplete skull.
Distribution: India, Nepal and ?Pakistan.
Distinguishing characters: Grey, darker and drabber on back, limbs
paler than flanks; spots and stripes on back black, brownish black to
brown on flanks, sometimes faint.
Prionailurus rubiginosus phillipsi Pocock, 1939; 278.
Type locality: Mousakanda, Gammaduwa, C.P., 3,000 ft [Sri Lanka].
Holotype: BMNH 1935.4.8.2 skin and skull.
Distribution: Sri Lanka.
Distinguishing characters: Darker, richer and less grey than P. r. rubigi-
nosus with flank spots brownish to rusty brown. Pocock (1939) found
no differences in skull measurements between rubiginosus and phil-
lipsi.
However, a third subspecies is sometimes recognised:
Prionailurus rubiginosus koladivius Deraniyagala, 1956; 113.
Type locality: Kathiraveli (E. P.) [Sri Lanka].
Holotype: Colombo National Museum 384.
Distribution: Lowland dry zone of E Sri Lanka.
Distinguishing characters: Darker head than P. r. phillipsi, blue-grey;
dorsal spots and stripes blackish, flank spots dark brown.
Discussion
Variation within subspecies is unclear. There appear to be two colour
morphs both in India and Sri Lanka, i.e. typical pale brown or rusty
spots, and dark brown to blackish spots. In Sri Lanka the dark-spotted
form is said to inhabit the lowland dry zone, but it is unknown whether
dark-spotted animals occupy drier habitats in India (Deraniyagala,
1956). There has been no phylogeographical study of Prionailurus ru-
biginosus.
We note several other taxonomic differentiations between Sri Lanka’s
wet and dry zone. Groves (1998) argued that within Sri Lanka there
are two species of Loris, one (Loris tardigradus) in the island’s wet
Subspecies Morphology Molecular Biogeography Certainty Comments
Prionailurus rubiginosus rubiginosus ++ ++ ++
Prionailurus rubiginosus phillipsi + o ++
Prionailurus rubiginosus koladivius + o +
rusty-spotted cat
Distribution of tentative subspecies of the rusty-spotted cat. Bor-
ders between subspecies in Sri Lanka are speculative.
P. r. koladivius
P. r. phillipsi
zone and the other (Loris lydekkerianus) in the dry country and me-
dium altitudes of Sri Lanka and also in southern India. A similar, but
not identical, pattern is also found in Trachypithecus vetulus, which
has two subspecies in the wet zone, one in the dry zone and at me-
dium altitudes, and one on the high mountains (Groves 2001). Groves
& Meijaard (2005) found that chevrotains (genus Moschiola) from Sri
Lanka’s wet zone were distinct in pelage, in body proportions, and in
skull proportions, and differed more from both the Indian and dry zone
Sri Lankan taxa than the two latter differed from each other. However,
there is no such differentiation among carnivorans.
In view of the uncertainties over whether dark-spotted and rusty-spot-
ted forms represent colour morphs, ecotypes or subspecies, we retain
three subspecies:
P. r. rubiginosus
© V. Athreya
CATnews Special Issue 11 Winter 2017
24
Prionailurus rubiginosus rubiginosus (I. Geoffroy Saint-Hilaire,
1831).
Dustribution: India and Nepal
Prionailurus rubiginosus phillipsi Pocock, 1939.
Distribution: Wet forest zone of Sri Lanka.
Prionailurus rubiginosus koladivius Deraniyagala, 1956.
Distribution: Lowland dry zone of E Sri Lanka.
References
Deraniyagala P. E. P. 1956. A new subspecies of rusty spotted cat from Ceylon.
Spolia Zeylanica 23, 113-114.
Geoffroy-Saint-Hilaire I. 1831. Mammifères. In Bélanger C. 1831. Voyage aux
Indes-Orientales par le nord de l’Europe, les provinces du Caucases, la
Géorgie, l’Arménie et la Perse, suivi des détails topographiques, statis-
Prionailurus planiceps
E: Flat-headed cat; F: Chat à tête plate; G: Flachkopfkatze; Sp: Gato
cabeciancho.
Prionailurus planiceps is normally treated as a monotypic species:
Prionailurus planiceps (Vigors and Horsfield, 1827; 450, plate XII).
Type locality: Sumatra.
Holotype: BMNH 1855.12.24.247 skin and skull.
Distribution: Borneo, Sumatra, Malay Peninsula, Thailand.
Discussion
Kitchener (1993) suggested that there may be subspecific differ-
entiation between Sumatra/Malay Peninsula and Borneo, but so
far no molecular or morphological data are available, which could
support this view. Luo et al. (2014) reported molecular genetic vari-
ation with two individuals from the Malay Peninsula and Borneo
respectively, which, based on mtDNA sequences, did not share
haplotypes.
Until such studies are completed, this species is treated here as
monotypic.
Prionailurus planiceps (Vigors and Horsfield, 1827; 450, plate XII).
Distribution: Borneo, Sumatra, Malay Peninsula, Thailand, Burma
Species Morphology Molecular Biogeography Certainty Comments
Prionailurus planiceps ++ ++ ++ Distinct monotypic species
tiques et autre sur le Pégou, les Iles de Jave, de Maurice et de Bourbon,
sur le Cap-de-bonne-Espérance et Sainte-Hélène, pendant les années 1825,
1826, 1827, 1828 et 1829 publié sous les auspices de ll. ee. mm. les Minis-
tres de la Marine et de l’Intérieur, 3. Zoologie. Arthus Bertrand, Paris.
Groves C. P. 1998. Systematics of tarsiers and lorises. Primates 39, 13-27.
Groves C. P. 2001. Primate Taxonomy. Smithsonian Institution Press, Wash-
ington D.C.
Groves C. P. & Meijaard E. 2005. Interspecific variation in Moschiola, the In-
dian chevrotain. Raffles Bulletin of Zoology, Supplement 12, 413-421.
Pocock R. I. 1939. The Fauna of British India, including Ceylon and Burma.
Mammalia – Vol. I. Primates and Carnivora (in part), Families Felidae and
Viverridae. Taylor and Francis, London.
Wozencraft W. C. 2005. Order Carnivora. In Mammal species of the world. A
taxonomic and geographic reference (3rd ed.). Wilson D. E. & Reeder D. M.
(Eds). Johns Hopkins University Press, Baltimore, pp. 532-628.
flat-headed cat
References
Kitchener A. C. 1993. A new look at subspecies in the Felidae. Lifeline,
May 1993, pp. 6-13.
Luo S.-J., Zhang Y., Johnson W. E., Miao L., Martelli P., Antunes A., Smith
J. L. D. & O’Brien S. J. 2014. Sympatric Asian felid phylogeography
reveals a major Indochinese-Sundaic divergence. Molecular Ecology
23, 2072-2092.
Vigors N. A. & Horsfield T. 1827. Descriptions of two species of the genus
Felis, in the collections of the Zoological Society. Zoological Journal 3,
449-451, plate XII.
© J. Sanderson
A new taxonomy of the Felidae
25
fishing cat
Distribution of tentative subspecies of the fishing cat. Borders be-
tween subspecies are speculative.
Prionailurus viverrinus
E: Fishing cat; F; Chat pêcheur, chat viverrin; G: Fischkatze; Sp: Gato
pescador
Wozencaft (2005) recognised two subspecies of the fishing cat:
Prionailurus viverrinus viverrinus (Bennett, 1833; 68).
Type locality: from the continent of India; probably the Malabar coast
(Pocock 1939).
Holotype: BMNH 1855.12.24.252 skin and part skull.
Distribution: Pakistan, Sri Lanka, India E to Indochina.
Prionailurus viverrinus rhizophoreus Sody, 1936; 45.
Type locality: Pamanoekan, North coast of West Java.
Holotype: RMNH.MAM 33859 skull and skin.
Distribution: Java.
Distinguishing character: Shorter basal length of skull than P. v. viver-
rinus form Siam [= Thailand] (Sody 1936).
Discussion
Despite being commonly cited in the literature, there are no records
that the fishing cat has ever occurred on Sumatra and there are only a
few uncertain records from peninsular Malaysia (Sody 1949, Van Bree
& Momin Khan 1992, Duckworth et al. 2009). Pocock (1939) was una-
ble to discern any geographical differentiation based on pelage colora-
tion and markings between fishing cats from throughout their range.
Luo et al. (2014) described the phylogeographical pattern of fishing
cats from northern Indochina based on multiple mitochondrial and nu-
clear markers, but specimens from other regions were lacking. This
species needs urgent research into its geographical variation, because
it is mostly intensely threatened throughout its geographical range.
Until a more comprehensive analysis becomes available, we recognise
the following subspecies:
Prionailurus viverrinus viverrinus (Bennett, 1833).
Distribution: India, Sri Lanka, Pakistan, Bangladesh, Indochina, Nepal,
and possibly Bhutan.
Prionailurus viverrinus rhizophoreus Sody, 1936.
Distribution: Java.
Distinguishing characters: Smaller basal length of skull than P. v. viver-
rinus from Siam [=Thailand] (Sody 1936).
References
Bennett E. T. 1833. Felis viverrina. Proceedings of the Zoological Society of
London 1, 68-69.
Duckworth J. W., Shepherd C. R., Semiadi G., Schauenberg P., Sanderson J.,
Roberton S. I., O’Brien T. G., Maddox T., Linkie M., Holden J. & Brickle N.
W. 2009. Does the fishing cat inhabit Sumatra? Cat News 51, 4-9.
Luo S.-J., Zhang Y., Johnson W. E., Miao L., Martelli P., Antunes A., Smith J.
L. D. & O’Brien S. J. 2014. Sympatric Asian felid phylogeography reveals a
major Indochinese-Sundaic divergence. Molecular Ecology 23, 2072-2092.
Pocock R. I. 1939. Fauna of British India, Mammals vol. 1. Taylor and Francis,
London.
Sody H. J. V. 1936. Seventeen generic, specific and subspecific names for
Dutch East Indian mammals. Natuurkundig Tijdschrift voor Nederlandsch
Indië 96, 42-55.
Sody H. J. V. 1949. Notes on some Primates, Carnivora and the babirusa from
the Indo-Malayan and Indo-Australian regions. Treubia 20, 121-190.
Van Bree P. J. H. & Momin Khan M. K. B. 1992. On a fishing cat, Felis (Prion-
ailurus) viverrina Bennett, 1833, from continental Malaysia. Zeitschrift für
Säugetierkunde 57, 179-180.
Wozencraft W. C. 2005. Order Carnivora. In Mammal species of the world. A
taxonomic and geographic reference (3rd ed.). Wilson D. E. & Reeder D. M.
(Eds). Johns Hopkins University Press, Baltimore, pp. 532-628.
Subspecies Morphology Molecular Biogeography Certainty Comments
Prionailurus viverrinus viverrinus ++ ++ ++
Prionailurus viverrinus rhizophoreus + o ++
P. v. viverrinus
P. v. rhizophoreus
© A. Sliwa
CATnews Special Issue 11 Winter 2017
26
mainland leopard cat
P. b. euptilurus in the Russian Far East (Photo L. Kerley).
Prionailurus bengalensis
E: Leopard cat; F: Chat-léopard du Bengale; G: Bengalkatze; Sp: Gato
bengali, gato de Bengala
This widespread species is usually recognised as having several subspe-
cies. Wozencraft (2005) recognised the following eleven subspecies:
Prionailurus bengalensis bengalensis (Kerr, 1792; 151).
Type locality: The coast of Bengal; restricted to S Bengal, India.
Holotype: From Pennant (1781; 164) a male that “swam on board a
ship at anchor off the coast of Bengal and produced young afterwards
with female cats in England”. The specimen’s remains were seen at
Hammersmith. However Pocock (1939) noted: “Although the story
hardly bears the impress of truth and Pennants’s description agrees
better with rich-coloured examples of the Sumatran race than with
any Indian skins I have seen, I adhere to the traditional acceptance of
Bengal as the locality and restrict it to the coast of that province to the
west of the Ganges.” A skin in the Natural History Museum (BMNH
1879.11.21.562) is labelled as a holotype, but originates from the In-
dian Museum, which seems unlikely since the specimen was last seen
in Hammersmith, London.
Distribution: Peninsular India, Burma, Thailand, Indochina.
Distinguishing characters: Short, thin coat, tail slender in winter
months. Ground colour ranges from ochreous buff to buffish white
on flanks, but darker on head and back. Spots large and well spaced,
sometimes solid and may run in chains.
Prionailurus bengalensis alleni Sody, 1949; 181.
Type locality: Nodoa [= Dan Xian], Hainan Dao, China.
Lectotype: AMNH M-59961 skin and skull adult male (Goodwin,
1956; 1).
Distribution: Hainan.
Distinguishing characters: Slightly smaller than those from the main-
land (P. b. chinensis) and nasals of males seems to be a trifle shorter
than in P. b. chinensis (Sody 1949).
Prionailurus bengalensis chinensis (Gray, 1837; 577).
Type locality: China.
Holotype: BMNH GERM 120a skin and skull.
Distribution: China.
Distinguishing character: Yellowish grey.
Prionailurus bengalensis horsfieldii (Gray, 1842; 260).
Type locality: India, Bhotan [= Bhutan].
Holotype: BMNH 1879.11.21.285 adult male skin and skull.
Distribution: Kashmir, Kumaon, Nepal and Bhutan; limits to N and NE
unknown.
Distinguishing characters: Larger skull than bengalensis, more luxuri-
ant coat and bushy tail in winter. Ground colour paler and not so richly
ochreous.
Prionailurus bengalensis euptilurus (Elliot, 1871; 761).
Type locality: Amur River, 60 km below mouth of Zeya River, Amur
Province, Russia.
Holotype: BMNH 1873.11.20.1 skin. Based on Felis undata of Radde,
1862; 106.
Distribution: Amur and Ussuri regions, Russia, NE China, Korean Pen-
insula.
Distinguishing characters: Ground colour light brownish yellow mixed
with grey. Spots reddish brown and rather oblong on flanks; darker
and browner on hind quarters and back. Tail thick and bushy with in-
complete rings.
Prionailurus bengalensis trevelyani Pocock, 1939; 273.
Type locality: Near Gilgit, Kashmir, 5000 feet.
Type: BMNH 1932.4.9.2 young adult male skin and skull.
Distribution: Northern Kashmir and the Upper Punjab in the drainage
area of the Indus and Jhelum, approximately 74° longitude, also Balu-
chistan.
Distinguishing characters: Fuller longer coat (36-40 mm long) than
horsfieldii and paler grey, sometimes nearly silvery ground colour, but
buff tinge low on flanks and on limbs.
Prionailurus bengalensis javanensis (Desmarest, 1816; 115).
Type locality: Java.
Holotype: MNHN-ZM-MO-2001-326 mounted skin.
Distribution: Java and Bali.
Distinguishing characters: Ground colour dark brownish grey; darker
on midline of back than on flanks. Four dark longitudinal stripes on
nape of neck are narrow and of equal width or inner ones wider than
outer ones. Markings blackish brown and are elongate dark spots on
midline of back and rounder on flanks.
Prionailurus bengalensis borneoensis Brongersma, 1935; 26.
Type locality: Rantau, SE Borneo.
Holotype: RMNH.MAM 467 male skin.
Distribution: Borneo.
Distinguishing characters: Ground colour ferruginous to tawny, but
darker than Sumatran animals. Nape stripes; inner pair narrower than
outer ones. Spots very dark, almost black; fewer elongate spots on
midline of back.
Prionailurus bengalensis sumatranus (Horsfield, 1821; pl. and text).
Type locality: Bencoolen, Sumatra [= Benkulu, Sumatra].
Holotype: BMNH GMCM 125a male skin.
Distribution: Sumatra.
Distinguishing characters: Ground colour ferruginous. Nape stripes;
outer pair broad, inner pair narrow. Spots dark brown to blackish; spots
on flanks elongate, but may be very small, roundish and numerous.
A new taxonomy of the Felidae
27
mainland leopard cat
Distribution of tentative subspecies of the mainland leopard cat.
Borders between subspecies are speculative.
Mainland leopard cat in captivity (Photo A. Sliwa).
Prionailurus bengalensis heaneyi Groves, 1997; 377.
Type locality: Puerto Princesa, Palawan.
Holotype: FMNH 62896 subadult male skin and skeleton.
Distribution: Palawan.
Distinguishing characters: Grey-fawn with small dark brown flank
spots.
Prionailurus bengalensis rabori Groves, 1997; 336.
Type locality: Canlaon, Negros Oriental.
Holotype: FMNH 74326 adult female skin and skull.
Distribution: Negros, Cebu and Panay, Philippines.
Distinguishing characters: Dark ochery to buffy fawn, but not as bright
as borneoensis.
Discussion
In addition Wozencraft (2005) recognised the Iriomote cat as a distinct
species:
Prionailurus iriomotensis (Imaizumi, 1967; 75).
Type locality: Haimida, Iriomote.
Holotype: NSM M 10890 adult male skin and almost complete skeleton.
Distribution: Iriomotejima, Ryukyu Islands, Japan.
However, recent molecular studies have clearly demonstrated that
P. iriomotensis is a leopard cat, and its skull morphology confirms this
(A. Kitchener, pers. obs.; contra Leyhausen & Pfleiderer 1999), who
considered its skull morphology unique and hence the taxon to be rec-
ognised as a distinct species. The pelage coloration is similar to that
of leopard cats from northern China and this form has almost certainly
arisen from a human introduction. Molecular studies confirm that P. iri-
omotensis is a leopard cat (Masuda et al. 1994, Masuda & Yoshida
1995, Suzuki et al. 1995), although estimated divergence times vary
from 100,000 to 200,000 years ago.
A recent molecular studiy, based on 1,792 bp of concatenated mtDNA
haplotypes (spanning cytochrome b, ATPase8 and 16S ribosomal DNA),
X-linked gene (PLP), 2,154 bp of concatenated Y-chromosome haplo-
types of intronic regions of three genes (DBY, SMCY3 and UTY11) and
one Y-linked microsatellite SMCY7-STR, has demonstrated that there
is a deep genetic divergence between Sundaland (the late Pleistocene
land mass connecting the islands of Sumatra, Borneo and Java) and
mainland leopard cats, with possible overlap in the Malay Peninsula
(Luo et al. 2014, see also Tamada et al. 2008 for first demonstration
of this deep split). Even more recently, Li et al. (2016) have shown a
“species-level” difference between mainland and island leopard cats.
However, it is unclear where the boundary between these two species
occurs, although there appear to be clear morphological differences
between the two; Sunda leopard cats have small solid spots while
mainland leopard cats have larger blotches filled with a lighter colo-
ration (A. Kitchener, pers. obs.).
Based on mitochondrial genomes, Patel et al. (2017) confirmed this
deep split, which they dated to more than 900,000 years ago, although
they tentatively regarded P. bengalensis as a single species, because
mainland and Sunda Island clades were sympatric in peninsular Ma-
laya, but this could also be regarded as evidence for two species.
Therefore, based on these recent molecular studies, coupled with clear
morphological differences, possible sympatry between two forms and
biogeographical separation, we recognise two species:
Prionailurus bengalensis (Kerr, 1792).
E: Mainland leopard cat; F: Chat-léopard du Bengale; G: Bengalkatze;
Sp: Gato bengali, gato de Bengala
Distribution: Mainland Asia from Pakistan to South East Asia, China
and the Russian Far East, Tsushima Island and Iriomote Island, Japan.
There is no comprehensive molecular review of mainland leopard cats
and hence it is unclear how many subspecies to recognise. Luo et al.
(2014) found no genetic differences between Chinese (chinensis) and
Indochinese (bengalensis) leopard cats. Tamada et al. (2008) found
two clades among mainland leopard cats; a northern clade comprising
continental Far Eastern, Korean (euptilurus), Iriomote I. (iriomotenis),
Tsushima I. and Taiwanese leopard cats and a southern clade of Indo-
chinese cats. This dichotomy is reinforced by the inclusion of Indian
leopard cats, which fall into the same clade as the Indochinese ani-
mals (Mukherjee et al. 2011). Based on mitochondrial genomes, Patel
et al. (2017) confirmed this split between northern and southern forms
although the boundary between the two is still uncertain. Patel et al.
(2017) also showed that many previously accepted subspecies could
not be recognised.
Thus we tentatively recognise two leopard cat subspecies in mainland
Asia:
P. b. bengalensis
P. b. euptilurus
CATnews Special Issue 11 Winter 2017
28
Sunda leopard cat
Prionailurus javanensis (Desmarest, 1816).
E: Sunda leopard cat; F: Chat-léopard de la Sonde; G: Sundakatze; Sp:
Gato de las Islas de la Sonda.
Distribution: Java, Bali, Borneo, Sumatra, Palawan, Negros, Cebu and
Panay, Philippines, possibly Malay Peninsula; probably introduced to
Philippines with possible exception of Palawan.
There is some morphological variation amongst the different island
populations of Sunda leopard cat. Javan animals appear to be distinct
from Sumatran and Bornean animals, which are more similar to each
other based on pelage coloration. It is possible that leopard cats from
the Philippines are the result of human introductions, although it is
more likely that those from Palawan are indigenous, having colonised
from Borneo during glaciations when sea levels were lower. A recent
phylogeographical study by Patel et al. (2017) has clarified the rela-
tionship between these putative island subspecies, confirming that
only two can be recognised:
Prionailurus javanensis javanensis (Desmarest, 1816).
Distribution: Java and Bali.
Distinguishing characters: Ground colour of pelage is brownish grey
(Brongersma 1935).
Prionailurus javanensis sumatranus (Horsfield, 1821).
Distribution: Sumatra, Borneo, and Palawan, Negros, Cebu and Panay,
the Philippines.
Distinguishing characters: Pelage ground coloration variable, ranging
from ferruginous to tawny, buffy-fawn and grey-fawn (Brongersma
1935; Groves 1997).
References
Brongersma L. D. 1935. Notes on some Recent and fossil cats, chiefly from the
Malay archipelago. Zoologische Mededeelingen 18, 1-89.
Desmarest A. G. 1816. Chat. Nouveau dictionnaire d’histoire naturelle 6,
73-123.
Elliot D. G. 1871. Remarks on various species of Felidae with a description of
a new species from north-western Siberia. Proceedings of the Zoological
Society of London 39, 758-761.
Goodwin G. G. 1956. The status of Prionailurus bengalensis alleni Sody. Amer-
ican Museum Novitates 1767, 1-3.
Gray J. E. 1837. Description of some new or little known Mammalia, princi-
pally in the British Museum collection. Magazine of Natural History, and
Journal of Zoology, Botany, Mineralogy, Geology and Meteorology N. S.
1, 577-587.
Gray J. E. 1842. Descriptions of some new genera and fifty unrecorded species
of Mammalia. Annals and Magazine of Natural History 10, 255-267.
Groves C. P. 1997. Leopard-cats, Prionailurus bengalensis (Carnivora: Felidae)
from Indonesia and the Philippines, with the description of two new sub-
species. Zeitschrift für Säugetierkunde 62, 330-338.
Gyldenstolpe N. 1919. A list of the mammals at present known to inhabit
Siam. Journal of the Natural History Society of Siam 3, 127-175.
Sunda leopard cat from Borneo (Photo Jo Ross and A. Hearn).
Subspecies Morphology Molecular Biogeography Certainty Comments
Prionailurus bengalensis bengalensis ++ ++ ++
Prionailurus bengalensis alleni + o ++
Prionailurus bengalensis chinensis + + o Same genetic clade as
bengalensis
Prionailurus bengalensis horsfieldii + + +
Prionailurus bengalensis euptilurus ++ ++ + Boundary with bengalensis
uncertain
Prionailurus bengalensis iriomotensis + + ++ More research required to de-
termine distinctiveness on island;
probably introduced by humans
Prionailurus bengalensis bengalensis (Kerr, 1792).
Distribution: S Asia from Pakistan to China and including probably the
Malay Peninsula.
Synonyms: horsfieldii, alleni, chinensis, trevelyani.
Prionailurus bengalensis euptilurus (Elliot, 1871).
Distribution: Manchuria, Russian Far East, Taiwan, Iriomote Island,
Tsushima Island.
Synonym: iriomotensis.
A new taxonomy of the Felidae
29
Sunda leopard cat
Horsfield T. 1821. Zoological researches in Java and the neighbouring islands.
Kingbury, Parbury and Allen, London.
Imaizumi Y. 1967. A new genus and species of cat from Iriomote, Ryukyu Is-
lands. Journal of the Mammalogical Society of Japan 3, 75-106.
Kerr R. 1792. The Animal Kingdom or zoological system of the celebrated Sir
Charles Linnaeus. Class I. Mammalia: Containing a complete systematic de-
scription, arrangement, and nomenclature, of all the known species and va-
rieties of the mammalia, or animals which give suck to their young, being a
translation of that part of the Systema Naturae as lately published with great
improvements by Professor Gmelin of Goettingen together with numerous ad-
ditions from more recent zoological writers and illustrated with copper plates.
Printed for A. Strahan, and T. Cadell, London, and W. Creech, Edinburgh.
Leyhausen P. & Pfleiderer M. 1999. Systematic status of the Iriomote cat (Pri-
onailurus iriomotensis Imaizumi, 1967) and the subspecies of the leopard
cat (Prionailurus bengalensis Kerr, 1792). Journal of Zoological Systemat-
ics and Evolutionary Research 37, 121-131.
Li G., Davis B. W., Eizirik E. & Murphy W. J. 2015. Phylogenomic evidence
for ancient hybridization in the genomes of living cats (Felidae). Genome
Research 26, 1-11.
Linnaeus C. 1758. Systema Naturae per regna tria naturae, secundum classis,
ordines, genera, species cum characteribus, differentiis, synonymis, locis.
10th edition, vol. 1. Holmiae: Laurentii Salvii.
Luo S.-J., Zhang Y., Johnson W. E., Miao L., Martelli P., Antunes A., Smith J.
L. D. & O’Brien S. J. 2014. Sympatric Asian felid phylogeography reveals a
major Indochinese-Sundaic divergence. Molecular Ecology 23, 2072-2092.
Masuda R. & Yoshida M. C. 1995. Two Japanese wildcats, the Tsushima cat
and Iriomote cat, show the same mitochondrial DNA lineage as the leop-
ard cat Prionailurus bengalensis. Zoological Science 12, 655-659.
Masuda R., Yoshida M. C., Shinyashiki F. & Bando G. 1994. Molecular phyloge-
netic status of the Iriomote cat Felis iriomotensis, inferred from mitochon-
drial DNA sequence analysis. Zoological Science 11, 597-604.
Mukherjee S., Krishnan A., Tamma K., Home C. R. N., Joseph S., Das A. &
Ramakrishnan U. 2010 Ecology driving genetic variation: A comparative
phylogeography of jungle cat (Felis chaus) and leopard cat (Prionailurus
bengalensis) in India. PLoS ONE 5(10): e13724.
Patel R. P., Wutke S., Lenz D., Mukherjee S., Ramakrishnan U., Veron G., Fickel
J., Wilting A. & Förster D. W. 2017. Genetic structure and phylogeography
of the Leopard cat (Prionailurus bengalensis) inferred from mitochondrial
genomes. Journal of Heredity esx017. doi: 10.1093/jhered/esx017
Pennant T. 1781. History of quadrupeds, vol. 1. B. White, London.
Pocock R. I. 1939. Fauna of British India, including Ceylon and Burma. Mam-
malia – Vol. I. Primates and Carnivora (in part), Families Felidae and Viver-
ridae. Taylor & Francis, London.
Radde G. 1862. Reisen im Süden von Ost-Sibirien in den Jahren 1855-1859
incl., im Auftrage der Kaiserlichen geographischen Gesellschaft aus-
geführt von Gustav Radde. Band 1. Die Säugethierfauna. St Petersburg:
Kaiserlichen Akademie der Wissenschaften.
Severtzov N. A. 1858. Notice sur la classification multisériale des Carnivores,
spécialement des Félidés, et les études de zoologie générale qui s’y rat-
tachent. Revue et magasin de zoologie pure et appliqué, Ser. 2, 10, 385-393.
Sody H. J. V. 1949. Notes on some Primates, Carnivora and the babirusa from
the Indo-Malayan and Indo-Australian regions. Treubia 20, 121-190.
Suzuki H., Hosoda T., Sakurai S., Tsuchiya K., Tsuchiya T., Munechika I. & Ko-
rablev V. P. 1994. Phylogenetic relationship between the Iriomote cat and
the leopard cat, Felis bengalensis, based on the ribosomal DNA. Japanese
Journal of Genetics 49, 397-405.
Tamada T., Siriaroonrat B., Subramaniam V., Hamachi M., Lin L.-K., Oshida T.,
Rerkamnuaychoke W. & Masuda R. 2008. Molecular diversity and phylo-
geography of the Asian leopard cat, Felis bengalensis, inferred from mi-
tochondrial and Y-chromosomal DNA sequences. Zoological Science 25,
154-163.
Wozencraft W. C. 2005. Order Carnivora. In Mammal species of the world. A
taxonomic and geographic reference (3rd ed.). Wilson D. E. & Reeder D. M.
(Eds). Johns Hopkins University Press, Baltimore, pp. 532-628.
Subspecies Morphology Molecular Biogeography Certainty Comments
Prionailurus javanensis javanensis ++ ++ ++
Prionailurus javanensis sumatranus ++ ++ ++
Prionailurus javanensis borneoensis + - ++ Consubspecific with sumatranus
Prionailurus javanensis heaneyi + - +
Probably introduced by humans;
consubspecific with sumatranus
Prionailurus javanensis rabori + - + Consubspecific with sumatranus
Distribution of tentative subspecies of the Sunda leopard cat. Bor-
ders between subspecies are speculative.
P. j. sumatranus
P. j. javanensis
CATnews Special Issue 11 Winter 2017
30
cheetah
Puma lineage
The Puma lineage contains three monotypic genera.
Genus Acinonyx Brookes, 1828; 16, 33.
The genus Acinonyx contains a single species, A. jubatus, which is read-
ily diagnosable from its distinctive morphology, including claws lacking
cutaneous sheaths, elongated lower limbs and skull lacking diastema.
Acinonyx jubatus
E: Cheetah; F: Guépard; G: Gepard; Sp: Guepardo, chita.
Smithers (1975) recognised the following five subspecies:
Acinonyx jubatus jubatus (Schreber, 1775; pl. 105; 1777; 392).
Type locality: Das Vaterland dieses Thieres ist das südliche Afrika;
man bekömmt die Felle vom Vorgebirge der guten Hofnung [The range
of this mammal is southern Africa; the skin came from the Cape of
Good Hope]. See Hollister (1911).
Holotype: Skin seen by Schreber.
Distribution: Southern Africa.
Acinonyx jubatus venaticus (Griffith, 1821; 93).
Type locality: India.
Holotype: From a sketch of a live animal by Mr Devis.
Distribution: SW Asia.
Acinonyx jubatus hecki Hilzheimer, 1913; 288.
Type locality: Senegal.
Holotype: Live animal in Berliner Zoologisches Garten.
Distribution: W Africa.
Acinonyx jubatus soemmeringii (Fitzinger, 1855; 245).
Type locality: von den Steppen der Kababisch im Süden der Bajuda-
Wüste [= from the steppes of the Kababish in the south of the Bayuda
Desert, Sudan].
Holotype: Male living in the Menagerie at Schönbrunn, Vienna.
Distribution: Sudan, Ethiopia, Horn of Africa.
Acinonyx jubatus raineyi Heller, 1913; 9.
Type locality: Ulu, Kapiti Plains, British East Africa [= Kenya].
Holotype: USNM 182321 adult male skin and skull.
Distribution: E Africa.
Wozencraft (2005) also listed velox Heller, 1913; 7 apparently in error,
but both these names are junior synonyms of ngorongorensis Hilzhei-
mer, 1913; 290.
Discussion
The most comprehensive phylogeographical study to date was by Char-
ruau et al. (2011). This analysis of mtDNA (NADH5, cytb and control
region) and 18 polymorphic nuclear microsatellites revealed a complex
star-shaped pattern in the mtDNA haplotype network, with suggestions
of geographical partitioning. For example, Asian, Arabian and north Af-
rican cheetahs tended to group together as did those from north-east
Africa, including Somalia, Sudan, Ethiopia and Djibouti. However, this
was not exclusive. East African cheetahs diverged into two different
lineages from those of southern Africa. C. Groves (pers. obs.) found that
Saharan cheetahs are probably distinguishable morphologically from
their small spots, but that Asian cheetahs did not seem to differ sig-
nificantly from African cheetahs. Charruau et al. (2011) concluded that
the following subspecies can be recognised genetically, although no
diagnostic morphological distinctions can be made currently:
Acinonyx jubatus jubatus (Schreber, 1775).
Distribution: Southern and eastern Africa.
Acinonyx jubatus soemmeringii (Fitzinger, 1855).
Distribution: NE Africa.
Acinonyx jubatus venaticus (Griffith, 1821).
Distribution: SW Asia and India.
Acinonyx jubatus hecki Hilzheimer, 1913.
Distribution: W and N Africa.
However, the divergence times between these lineages are very re-
cent (Charruau et al. 2011), e.g. 32,000-67,000 ya between jubatus and
venaticus, and 16,000-72,000 ya between jubatus and soemmeringii,
and the inclusion of ancient DNA samples from north Africa and south-
west Asia blurred the distinction between north African and Asian
cheetahs, suggesting isolation by distance. It is possible that there are
only two subspecies of cheetah; northern (venaticus) and southern/
eastern (jubatus), or perhaps none if further more comprehensive sam-
pling of museum specimens is carried out.
Distribution of tentative subspecies of cheetah. Borders between
subspecies are speculative.
A. j. venaticus
A. j. hecki
A. j. soemmeringii
A. j. jubatus
© P. Meier
A new taxonomy of the Felidae
31
References
Brookes J. 1828. Catalogue of the Anatomical and Zoological Museum of
Joshua Brookes. Gold and Walton, London.
Charruau P., Fernandes C., Orozco-Ter Wengel P., Peters J., Hunter L., Ziaie
H., Jourabchian A., Jowkar H., Schaller G., Ostrowski S., Vercammen P.,
Grange T., Schlötterer C., Kotze A., Geigl E.-M., Walzer C. & Burger P. A.
2011. Phylogeography, genetic structure and population divergence time
of cheetahs in Africa and Asia: Evidence for long-term geographic isolates.
Molecular Ecology 20, 706-724.
Fitzinger L. 1855. Bericht an die kaiserl. Akademie der Wissenchaften über
die von dem Herrn Consultatsverweser Dr. Theodor v. Heuglin für die
kaiserliche Menagerie zu Schönbrunn mitgebrachten lebenden Thiere.
Sitzungsberichte der Mathematisch-Naturwissenschaftlichen Classe der
kaiserlichen Akademie der Wissenschaften 17, 242-253.
Griffith E. 1821. General and particular descriptions of the vertebrated ani-
mals, arranged conformably to the modern discoveries and improvements
in zoology. Order Carnivora. Baldwin, Cradock and Joy, London.
Heller E. 1913. New races of carnivores and baboons from Equatorial Africa
and Abyssinia. Smithsonian Miscellaneous Collections 61, 1-12.
Hilzheimer M. 1913. Über neue Gepparden nebst Bemerkungen über die
Nomenklatur dieser Tiere. Sitzungsberichte der Gesellschaft Naturfor-
schender Freunde zu Berlin 1913, 283-292.
Hollister N. 1911. The nomenclature of the cheetahs. Proceedings of the Bio-
logical Society of Washington 24, 225-230.
Schreber J. C. D. 1775. Die Säugethiere in Abbildungen nach der Natur mit
Beschreibungen, vol.2 (15). Wolfgang Walther, Erlangen.
Schreber J. C. D. 1777. Die Säugethiere in Abbildungen nach der Natur mit
Beschreibungen, vol.3 (22). Wolfgang Walther, Erlangen.
Smithers R. H. N. 1975. 8.1 Family Felidae. In The mammals of Africa. An iden-
tification manual. Meester J. & Sezter H. W. (Eds). Smithsonian Institution
Press, Washington D.C.
Wozencraft W. C. 2005. Order Carnivora. In Mammal species of the world. A
taxonomic and geographic reference (3rd ed.). Wilson D. E. & Reeder D. M.
(Eds). Johns Hopkins University Press, Baltimore, pp. 532-628.
jaguarundi
Subspecies Morphology Molecular Biogeography Certainty Comments
Acinonyx jubatus jubatus ++ ++ ++
Acinonyx jubatus venaticus + ++ ++ Possibly includes hecki and soemmeringii
Acinonyx jubatus hecki + + + Possible synonym of venaticus
Acinonyx jubatus soemmeringii + + + Possible synonym of venaticus
Acinonyx jubatus raineyi + - - Synonym of jubatus
Genus Herpailurus Severtzov, 1858; 385.
As used here, this is a monotypic genus, but it may be included within
Puma.
Herpailurus yagouaroundi
E: Jaguarundi; F: Jaguarondi; G: Jaguarundi, Wieselkatze, Eyra; Sp:
Yaguarundi, onza, gato moro, gato eyra.
Wozencraft (2005) recognised eight subspecies of jaguarundi:
Herpailurus yagouaroundi yagouaroundi (É. Geoffroy Saint-Hi-
laire, 1803; 124).
Type locality: Paraguay, restricted to Cayenne, French Guiana by Her-
shkovitz (1951; 565).
Holotype: Based on l’yagouaroundi of Azara, who had two females
(1801; 171).
Distribution: E Venezuela, the Guianas and NE Brazil.
Herpailurus yagouaroundi ameghinoi (Holmberg, 1898; 485).
Type locality: San Luis [Argentina].
Holotype: Skin in Turin Museum.
Distribution: Argentina.
Herpailurus yagouaroundi cacomitli (Berlandiere, in Baird,1859; 12).
Type locality: Matamoros, Tamaulipas [Mexico].
Holotype: USNM 1426 adult female skull.
Distribution: E Mexico as far north as S Texas.
Herpailurus yagouaroundi eyra (G. Fischer, 1814; 228).
Type locality: Paraguay.
Holotype: Based on a live animal in the possession of Azara, which he
called l’eyra (D’Azara, 1801; 177).
Distribution: S Brazil, Paraguay and N Argentina in Misiones and the
Mediterranean and Chaco zones.
A very red jaguarundi from Eastern Amazonia, Brazil (Photo Pro-
jeto Gatos do Mato - Brasil).
CATnews Special Issue 11 Winter 2017
32
jaguarundi
Herpailurus yagouaroundi fossata (Mearns, 1901; 150).
Type locality: Merida, Yucatán [Mexico].
Holotype: USNM 7036 adult skull.
Distribution: Honduras, Belize, Guatemala and Yucatan, Mexico.
Herpailurus yagouaroundi melantho (Thomas, 1914; 350).
Type locality: Pozuzo, deparamento de Huánuco 800m [Peru].
Holotype: BMNH 1908.6.17.10 male skin and skull.
Distribution: Peru in the valleys of the Andes.
Herpailurus yagouaroundi panamensis (J. A. Allen, 1904: 71).
Type locality: Boqueron, Chiriqui, Panama.
Holotype: AMNH M-18946 subadult female skin and skull.
Distribution: E of Colombia and possibly Ecuador extending N to Pan-
ama and Costa Rica.
Herpailurus yagouaroundi tolteca (Thomas, 1898; 41).
Type locality: Tatemales, Sinaloa [Mexico].
Holotype: BMNH 1898.3.2.17 male skin and skull.
Distribution: W Mexico as far N as S Arizona.
Discussion
This species is polymorphic with at least three common pelage col-
ours. Da Silva et al. (2016) carried out ecological modelling of the two
principal coat colours; dark/grey pelage is associated mostly with wet,
dense forests, whereas the ancestral red coat colour is associated
principally with dry, open habitats. A recent phylogeographical study
by Ruiz-García & Pinedo-Castro (2013) based on three mitrochondrial
genes (ATP8, 16S rRNA and NADH5) found no evidence for subspe-
cies. On the basis of this study we regard Herpailurus yagouaroundi
as a monotypic species:
Herpailurus yagouaroundi (É. Geoffroy Saint-Hilaire, 1803).
Distribution: C and S America.
References
Allen J. A. 1904. Mammals from southern Mexico and Central and South
America. Bulletin of the American Museum of Natural History 20, 29-80.
Da Silva L. G., de Oliveira T. G., Kasper C. B., Cherem J. J., Moraes Jr E. A.,
Paviolo A. & Eizirik E. 2016. Biogeography of polymorphic phenotypes:
Mapping and ecological modelling of coat colour variants in an elusive
Neotropical cat, the jaguarundi (Puma yagouaroundi). Journal of Zoology
299, 295-303.
D’Azara F. 1801. Essais sur l’Histoire Naturelle des Quadrupedes de la Prov-
ince de Paraguay, vol 1. Charles Pougens, Paris.
Berlandier L. 1859. In Baird S.F. Mammals of the boundary. In Report on the
United States and Mexican boundary survey made under the direction of
the Secretary of the Interior by William H. Emory, Major First Cavalry and
United States Commissioner 2, 3-55.
Fischer G. 1814. Zoognosia tabulis synopticis illustrata. In usum praelectionum
Academiae imperialis medico-chirugicae mosquensis edita, vol. 3. Nicolai
Sergeidis Vsevolozsky, Moscow.
Geoffroy St. Hilaire É. 1803. Catalogue des Mammifères du Muséum National
d’Histoire Naturelle, Paris, France.
Hershkovitz P. 1951. Mammals from British Honduras, Mexico, Jamaica and
Haiti. Fieldiana Zoology 31, 547-569.
Holmberg L. E. 1898. La fauna de la República Argentina. In Segundo Censo de
la República Argentina Mayo 10 de 1895. Tomo 1. Territorio. Taller Tipográ-
fico de la Penitenciaría Nacional, Buenos Aires, pp. 477-602.
Mearns E. A. 1901. Two new cats of the eyra group from North America. Pro-
ceedings of the Biological Society of Washington 14, 149-151.
Ruiz-García M. & Pinedo-Castro M. 2013. Population genetics and phylogeo-
graphic analyses of the jaguarundi (Puma yagouaroundi) by means of three
mitochondrial markers: The first molecular study of this species. In Molec-
ular population genetics, evolutionary biology and biological conservation
on Neotropical carnivores. Ruiz-García M. & Shostell J. M. (Eds). Nova,
New York, pp. 245-288.
Severtzov A. N. 1858. Notice sur la classification multisériale des Carnivores,
spécialement des Félidés, et les études de zoologie générale qui s’y rat-
tachent. Revue et magasin de zoologie pure et appliqué, Ser. 2, 10, 385-
393.
Thomas O. 1898. On new mammals from Western Mexico and Lower Califor-
nia. The Annals and Magazine of Natural History (7th series) 1, 40-46.
Thomas O. 1914. On various South-American mammals. The Annals and Mag-
azine of Natural History (8th series) 13, 345-363.
Wozencraft W. C. 2005. Order Carnivora. In Mammal species of the world. A
taxonomic and geographic reference (3rd ed.). Wilson D. E. & Reeder D. M.
(Eds). Johns Hopkins University Press, Baltimore, pp. 532-628.
Subspecies Morphology Molecular Biogeography Certainty Comments
Herpailurus yagouaroundi yagouaroundi ++ ++ ++
Herpailurus yagouaroundi ameghinoi + - o
Herpailurus yagouaroundi cacomitli + - o
Herpailurus yagouaroundi eyra + - o
Herpailurus yagouaroundi fossata + - o
Herpailurus yagouaroundi melantho + - o
Herpailurus yagouaroundi panamensis + - o
Herpailurus yagouaroundi tolteca + - o
A new taxonomy of the Felidae
33
puma
Genus Puma Jardine, 1834; 266.
This is a monotypic genus, but may be revised to include Herpailurus.
Puma concolor
E: Puma, cougar, mountain lion; F: Puma, couguar; G: Puma, Silber-
löwe; Sp: Puma, léon americano, léon bayo, léon colorado, onza ber-
meja.
Traditionally this widespread species has been split into as many
as 32 subspecies, most of which were of doubtful validity (Young &
Goldman 1946). Culver et al. (2000) carried out a phylogeographical
study of pumas throughout their range based on both contemporary
and museum samples and analysis of three mitochondrial gene se-
quences (16S rRNA, ATPase-8, NADH-5) and composite microsatellite
genotypes (10 feline loci).
On the basis of this study, six phylogeographical groups were claimed
and designated as subspecies:
Puma concolor concolor (Linnaeus, 1771; 522).
Locality from where the subspecies was first described: Brassilia; re-
stricted by Goldman (in Goldman & Young 1946) to Cayenne region,
French Guiana.
Type: None designated.
Range: N and W South America.
Puma concolor puma (Molina, 1782; 295).
Locality from where the subspecies was first described: Chile; restrict-
ed to “in the vicinity of Santiago” by Nelson & Goldman (1929).
Type: None designated.
Range: S South America.
Puma concolor couguar (Kerr, 1792; 151).
Type locality: Pennsylvania, Virginia, Carolina and Georgia, in North
America; restricted to Pennsylvania by Nelson & Goldman (1929).
Holotype: Based on Cougar de Pensilvanie of Buffon (1776; 222).
Range: North America.
Puma concolor capricornensis (Goldman, 1946; 246).
Type locality: Piracicaba, about 80 miles northwest of Sao Paulo, Bra-
zil.
Holotype: USNM 100118 adult male skull.
Range: NE South America.
Puma concolor costaricensis (Merriam, 1901; 596).
Type locality: Boquete, Chiriqui, Panama.
Holotype: MCZ 10118 female skin and skull.
Range: Costa Rica and Panama.
Puma concolor cabrerae (Pocock, 1940; 308).
Type locality: La Rioja, province of La Rioja, northern Argentina, alti-
tude 968 metres.
Holotype: BMNH 1874.8.4.2 adult male skull.
Range: SE South America.
Wozencraft (2005) listed these subspecies, except for capricornensis,
but added anthonyi, without explanation. This appears to be an error.
Discussion
A more recent study of mtDNA in pumas from throughout their range,
although with lower sample sizes, supports only two main geographi-
cal groupings with North American populations having colonised since
c. 8,000 years b.p. (Caragiulo et al. 2014).
On this basis, we tentatively recognise two subspecies within Puma
concolor:
Puma concolor concolor (Linnaeus, 1771).
Distribution: South America, possibly excluding W of Andes in north.
Puma concolor couguar (Kerr, 1792).
Distribution: North and Central America, possibly N South America W
of Andes.
References
Buffon G.-L. L. 1776. Cougar de Pensilvanie. In Histoire naturelle générale et
particulière. Servant de suite à l’histoire des animaux quadrupeds. Supple-
ment, Tome troisième. Buffon, G.-L. L., Comte de and Daubenton, L.-J.-M.
(Eds). Imprimerie Royale, Paris, pp. 222-223.
Caragiulo A., Dias-Freedman I., Clark J. A., Rabinowitz S. & Amato G. (2014).
Mitochondrial DNA sequence variation and phylogeography of Neotropic
pumas. Mitochondrial DNA 25, 304-312.
Culver M., Johnson W. E., Pecon-Slattery J. & O’Brien S. J. 2000. Genomic
ancestry of the American puma (Puma concolor). Journal of Heredity 91,
186-197.
Goldman E. A. 1946. Classification of the races of the puma. In The puma.
Mysterious American cat, pp. 175-302. Young S. P. & Goldman E. A. (Eds).
The American Wildlife Institute, Washington D.C.
Jardine Sir W. 1834. Naturalists’ library, Mammalia, volume 2. The Felinae.
Lizars, and Stirling and Kenney, Edinburgh.
Kerr R. 1792. The Animal Kingdom or zoological system of the celebrated Sir
Charles Linnaeus. Class I. Mammalia: Containing a complete systematic
description, arrangement, and nomenclature, of all the known species and
varieties of the mammalia, or animals which give suck to their young, be-
ing a translation of that part of the Systema Naturae as lately published
with great improvements by Professor Gmelin of Goettingen together with
numerous additions from more recent zoological writers and illustrated
with copper plates. Printed for A. Strahan, Edinburgh, T. Cadell, London,
and W. Creech, Edinburgh.
Linnaeus C. 1771. Mantissa plantarum altera. Generum editionis VI et spe-
cierum editionis II. Regni animalis appendix. Holmiae: Laurentii Salvii.
A puma in California (S. Kennerknecht/pumapix).
CATnews Special Issue 11 Winter 2017
34
marbled cat
Bay Cat lineage
Genus Pardofelis Severtzov, 1858; 387.
Pardofelis as defined here is usually regarded as a monotypic genus
with the species, P. marmorata (Wozencraft 2005).
Pardofelis marmorata (Martin, 1837; 108).
E: Marbled cat; F: Chat marbré; G: Marmorkatze; Sp: Gato jaspeado.
Type locality: Java or Sumatra; restricted to Sumatra by Robinson &
Kloss (1919; 261).
Holotype: BMNH 1855.12.24.25 young male, skin and skull, from
Sumatra
Distribution: Sumatra, Borneo, SE Asia as far N as Yunnan, N Burma,
Assam to Nepal (Corbet & Hill 1992).
P. marmorata is usually divided into two distinct subspecies:
P. m. marmorata
Type: As above.
Distribution: Sumatra, Borneo, Mainland SE Asia excluding N Burma
to India.
Distinguishing characters: Greyer with large distinct blotches.
P. m. charltonii (Gray, 1846; 211).
Type locality: Darjiling (= Darjeeling), North India.
Holotype: Female skin BMNH 1846.3.4.6 and skull BMNH 1846.3.17.23.
Distribution: Nepal to Assam, Bangladesh and N Burma.
Distinguishing characters: Rich ochreous brown, limited blotch-like
markings.
Distribution of tentative subspecies of puma. Borders between
subspecies are speculative.
Merriam C. H. 1901. Preliminary revision of the pumas (Felis concolor group).
Proceedings of the Washington Academy of Sciences 3, 577-600.
Molina G. I. 1782. Saggio sulla storia naturale del Chilli. Stamperia di S. Tom-
maso d’Aquino, Bologna.
Nelson E. W. & Goldman E. A. 1929. List of the pumas with three described as
new. Journal of Mammalogy 10, 345-350.
Pocock R. I. 1940. Description of a new race of puma (Puma concolor), with a
note on an abnormal tooth growth in the genus. Annals and Magazine of
Natural History (11th series) 6, 307-313.
Wozencraft W. C. 2005. Order Carnivora. In Mammal species of the world. A
taxonomic and geographic reference (3rd ed.). Wilson D. E. & Reeder D. M.
(Eds). Johns Hopkins University Press, Baltimore, pp. 532-628.
Young S. P. & Goldman E. A. 1946. The puma. Mysterious American cat. The
American Wildlife Institute, Washington D.C.
Subspecies Morphology Molecular Biogeography Certainty Comments
P. c. concolor + ++ ++
P. c. couguar + ++ ++
P. c. anthonyi + + o Probably junior synonym of P. c. concolor
P. c. capricornensis + + o Probably junior synonym of P. c. concolor
P. c. costaricensis + + o Probably junior synonym of P. c. couguar
P. c. cabrerae + + o Probably junior synonym of P. c. concolor
P. c. puma + + o Probably junior synonym of P. c. concolor
A marbled cat in Borneo (S. Kennerknecht/Panthera).
P. c. couguar
P. c. concolor
A new taxonomy of the Felidae
35
marbled cat
Discussion
A preliminary analysis confirmed two basic pelage patterns, but their
distributions are different from those above, with the greyer southern
form restricted to the Sunda Islands and the Malay Peninsula as far
north as the Isthmus of Kra, and the ochreous northern form ranging
from throughout mainland SE Asia north of the Isthmus of Kra to Ne-
pal. Preliminary analysis suggests that the pelages of these two forms
are consistently distinct and geographically separated (A. Kitchener &
E. Meijaard, pers. obs.; see also Brongersma 1935; 33).
A recent molecular study, based on mtDNA, X-linked and Y-linked
nuclear genes supports this view (Luo et al. 2014). Eleven samples
from animals in Indochina showed a divergence time of about two
million years compared with three Sunda animals, which were from
the Malay Peninsula. Further molecular and morphological research
is required to confirm the results of these preliminary studies. Given
the wider geographical distribution of the northern form, the earliest
available name is Felis longicaudata based on a dried specimen col-
lected by Diard probably from Cochinchina, from which the skeleton
was extracted, but only the skull was figured (Blainville 1843).
The analysis of pelage patterns also suggested that there could be
differentiation between Sumatra (greyer) and Borneo (browner) popu-
lations (see also Brongersma 1935; 33), which may be recognised as
distinct subspecies. If so, the Bornean subspecies would require a for-
mal scientific description as a new subspecies.
The following taxonomic arrangement is tentative and awaits a
more in-depth molecular and morphological study, which may show
that there are two distinct species and a possible new subspecies
on Borneo.
Pardofelis marmorata marmorata (Martin, 1837).
Distribution: Borneo, Sumatra, Malay Peninsula S of Isthmus of Kra
and S Thailand.
Distinguishing characters: Greyer with large distinct blotches.
Pardofelis marmorata longicaudata (Blainville, 1843; 186, pl. X).
Type locality: de l’Inde (Cochinchine?).
Holotype: MNHN-ZM-AC-A3424 articulated skeleton.
Distribution: Nepal to Assam, Bangladesh, SE Asia N of the Isthmus
of Kra.
Distinguishing characters: Rich to pale ochreous brown, limited blotch-
like markings.
References
Blainville de H. M. D. 1843. Ostéographie ou description iconographique com-
parée du squelette et du système dentaire des cinques classes d’animaux
vertébrés récents et fossils pour servir de base a la zoologie et la géologie.
Mammifères. Carnassiers. Vol. 2. Arthus Bertrand, Paris.
Brongersma L. D. 1935. Notes on some recent and fossil cats, chiefly from the
Malay Archipelago. Zoologische mededeelingen Rijksmuseum Leiden 18,
1-93.
Corbet G. B. & Hill J. E. 1992. The mammals of the Indomalayan region. Oxford:
Oxford University Press.
Gray J. E. 1846. New species of Mammalia. Annals and Magazine of Natural
History 18, 211-212.
Luo S.-J., Zhang Y., Johnson W. E., Miao L., Martelli P., Antunes A., Smith J.
L. D. & O’Brien S. J. 2014. Sympatric Asian felid phylogeography reveals
a major Indochinese-Sundaic divergence. Molecular Ecology 23, 2072-
2092.
Martin W. 1837. A new species of the genus Felis. Proceedings of the Zoologi-
cal Society of London 4, 107-108.1
Pocock R. I. 1932. The marbled, Pardofelis marmorata, cat and some other Ori-
ental species, with a definition of a new genus of the Felidae. Proceedings
of the Zoological Society of London 102, 741-766.
Robinson H. C. & Kloss C. B. 1919. On a collection of mammals from the Ben-
coolen and Palembang residencies, South West Sumatra. Journal of the
Federated Malay States Museum 7, 257-291.
Severtzov N. A. 1858. Notice sur la classification multisériale des Carnivores,
spécialement des Félidés, et les études de zoologie générale qui s’y rat-
tachent. Revue et magasin de zoologie pure et appliqué, Ser. 2, 10, 385-
393.
Wozencraft W. C. 2005. Order Carnivora. In Mammal species of the world. A
taxonomic and geographic reference (3rd ed.). Wilson D. E. & Reeder D. M.
(Eds). Johns Hopkins University Press, Baltimore, pp. 532-628.
Distribution of tentative subspecies of marbled cats. Borders be-
tween subspecies are speculative.
Subspecies Morphology Molecular Biogeography Certainty Comments
Pardofelis marmorata marmorata ++ ++ ++
Pardofelis marmorata longicaudata ++ + ++ Possibly distinct species
P. m. marmorata
P. m. longicaudata
CATnews Special Issue 11 Winter 2017
36
Borneo bay cat, Asiatic golden cat
Genus Catopuma Severtzov, 1858; 387.
The genus Catopuma contains two polychromatic species, the bay cat,
C. badia, from Borneo and the Asiatic golden cat, C. temminckii, from
Sumatra and the mainland of Asia.
Catopuma badia
E: Bay cat, Borneo bay cat; F: Chat bai; G: Borneo Goldkatze; Sp: Gato
rojo de Borneo.
Although sometimes considered to be conspecific with C. temminckii,
C. badia is a much smaller distinct monotypic species confined to Bor-
neo (Wozencraft 2005). Like C. temminckii, it is polymorphic with red-
dish, greyish and mixed pelage colorations and there appears to be no
geographical separation of these colour morphs and so we conclude
that the species is monotypic.
Catopuma badia (Gray, 1874; 322).
Type locality: Sarawak, Borneo.
Holotype: BMNH 56.9.19.16 young male skin and skull.
Distribution: Borneo.
Diagnosis: Compared with C. temminckii, much smaller (head-and-
body length (HB): 73-105 cm C. temminckii; 50-69 cm C. badia (Kitch-
ener 1991) with reddish and greyish morphs.
References
Gray J. E. 1874. Description of a new species of cat (Felis badia) from Sarawak.
Proceedings of the Zoological Society of London 42, 322-323.
Kitchener A. 1991. The natural history of the wild cats. A & C Black, London.
Severtzov M. N. 1858. Notice sur la classification multisériale des Carnivores,
spécialement des Félidés, et les études de zoologie générale qui s’y rat-
tachent. Revue et magasin de zoologie pure et appliqué, Ser. 2, 10, 385-
393.
Wozencraft W. C. 2005. Order Carnivora. In Mammal species of the world. A
taxonomic and geographic reference (3rd ed.). Wilson D. E. & Reeder D. M.
(Eds). Johns Hopkins University Press, Baltimore, pp. 532-628.
Species Morphology Molecular Biogeography Certainty Comments
Catopuma badia ++ ++ ++ Monotypic species
Catopuma temminckii
E: Asiatic golden cat, Temminck’s golden cat; F: Chat de Temminck,
chat doré d’Asie; G: Asiatische Goldkatze; Sp: Gato dorado asiático.
This polymorphic species is usually divided into two to three subspe-
cies (Wozencraft 2005):
Catopuma temminckii temminckii (Vigors and Horsfield, 1827; 451).
Type locality: Sumatra.
Holotype: BMNH 1855.12.24.250 juvenile male skin and skull.
Distribution: Sumatra, Malay Peninsula, Indochina, Burma to Nepal.
Catopuma temminckii dominicanorum (Sclater, 1898; 2, pl. 1).
Type locality: Kuatun, Foochow, China.
Holotype: BMNH 1899.12.27.1 male skin and skull.
Distribution: S China.
Catopuma temminckii tristis (Milne-Edwards, 1872; 223, pl.31d).
Type locality: de l’interieur de la Chine [from the interior of China].
Holotype: MNHN-ZM-MO-1867-535 adult male mounted skin.
Distribution: Tibet, Sichuan and Upper Burma.
Wozencraft (2008) suggested that SE Asian C. temminckii are prob-
ably distinct from those of Sumatra and those from Nepal, S Tibet and
probably NW Yunnan and W Sichuan, thereby recognising two further
subspecies:
Catopuma temminckii bainesi (Sowerby, 1924; 352).
Type locality: Tengueh, S.W. Yunnan China.
Holotype: Skin was in Royal Asiatic Society (North China Branch) Mu-
seum, Shanghai.
Distribution: Yunnan.
Catopuma temminckii moormensis (Hodgson, 1831; 177).
Type locality: Nepal.
Holotype: BMNH GERM 118a skull.
Distribution: Nepal, S Tibet and probably NW Yunnan and W Sichuan,
China.
© S. Kennerknecht/Panthera
© A. Sliwa
A new taxonomy of the Felidae
37
Asiatic golden cat
Discussion
This is a very variable species with a wide range of pelage colorations
and markings, but northern populations seem to be particularly poly-
morphic. There is also a large difference in size between animals from
Sumatra and the Malay Peninsula and those from SE Asia and China.
Luo et al. (2014) carried out a first phylogeographical study on C. tem-
minckii with specimens from China, Indochina and the Malay Penin-
sula. No samples from Sumatra or parts of the western distribution
range were included. Based on mtDNA (spanning cytochrome b, AT-
Pase8 and 16S ribosomal DNA), X-linked gene (PLP), Y-chromosome
haplotypes of intronic regions of three Y-linked genes (DBY, SMCY3
and UTY11) and one Y-linked microsatellite SMCY7-STR C. temminckii
populations showed a relatively recent divergence time with a sepa-
ration between populations from the Malay Peninsula (n=7 plus one
Sunda specimen of unknown locality) and, by inference, Sumatra from
those north of the Isthmus of Kra (n = 36).
Patel et al. (2016) carried out a comprehensive study of whole mito-
chondrial genomes and pelage coloration in C. temminckii. They found
that this species has diversified since around the time of the Toba su-
per-eruption in Sumatra c.74 kya. No geographical structure was found
in the genetic data in mainland Asia, but there was a more or less
distinct clade that included animals from the Malay Peninsula and Su-
matra. This latter clade displays the least polychromatism compared
with mainland populations.
On the basis of this study, we suggest the recognition of two sub-
species:
Catopuma temminckii temminckii (Vigors and Horsfield, 1827).
Distribution: Sumatra and the Malay Peninsula.
Distinguishing characters: Relatively small, typical reddish coloration
(melanistic morphs also).
Catopuma temminckii moormensis (Hodgson, 1831).
Distribution: From Nepal to N Burma, China, Tibet and SE Asia.
Distinguishing characters: Relatively large, pelage very variable rang-
ing from blotches and spots to dark grey, blackish, brown and reddish
morphs.
References
Hodgson B. H. 1831. Some account of a new species of Felis. Gleanings in
Science 3, 177-178.
Luo S.-J., Zhang Y., Johnson W. E., Miao L., Martelli P., Antunes A., Smith J.
L. D. & O’Brien S. J. 2014. Sympatric Asian felid phylogeography reveals a
major Indochinese-Sundaic divergence. Molecular Ecology 23, 2072-2092.
Milne-Edwards A. 1872. Étude pour servir à l’histoire de la faune mamm-
alogique de la Chine. Recherches pour servir à l’histoire des mammifères
comprenant des considérations sur la classification de ces animaux. Mas-
son, Paris, pp. 67-229.
Patel R. P., Förster D. W., Kitchener A. C., Rayan M. D., Mohamed S. W., Wer-
ner L., Lenz D., Pfestorf H., Kramer-Schadt S., Radchuk V., Fickel J. & Wilt-
ing A. 2016. Two species of southeast Asian cats in the genus Catopuma
with diverging histories: An island endemic forest specialist and a wide-
spread habitat generalist. Royal Society Open Science 3: 160350.
Pocock R. I. 1932. The marbled cat (Pardofelis marmorata) and some other
Oriental species, with the definition of a new genus of the Felidae. Pro-
ceedings of the Zoological Society of London 102, 741-766.
Sclater P. L. 1898. Report on the additions to the Society’s menagerie. Proceed-
ings of the Zoological Society 66, 1-2, pl. I.
Sowerby A. de C. 1924. A new cat from west China. China Journal of Science
and Arts 2, 352-353.
Vigors N. A. & Horsfield T. 1827. Descriptions of two species of the genus
Felis, in the collections of the Zoological Society. Zoological Journal 3,
449-451.
Wozencraft W. C. 2005. Order Carnivora. In Mammal species of the world. A
taxonomic and geographic reference (3rd ed.). Wilson D. E. & Reeder D. M.
(Eds). Johns Hopkins University Press, Baltimore, pp. 532-628.
Wozencraft W. C. 2008. Order Carnivora – carnivores. In A guide to the mam-
mals of China. Smith, A.T. & Xie Y. (Eds). Princeton University Press, Prince-
ton, pp. 388-448.
Distribution of tentative subspecies of Asiatic golden cat. Borders
between subspecies are speculative.
Subspecies Morphology Molecular Biogeography Certainty Comments
Catopuma temminckii temminckii ++ ++ ++
Catopuma temminckii dominicanorum + o o Colour variant of moormensis
Catopuma temminckii tristis ++ o o Colour variant of moormensis
Catopuma temminckii bainesi + o o Colour variant of moormensis
Catopuma temminckii moormensis + + ++
C. t. moormensis
C. t. temminckii
CATnews Special Issue 11 Winter 2017
38
bobcat
Lynx lineage
Genus Lynx Kerr, 1792; 157.
Monophyletic group with four species, recognised from morphology
(Nowak 1999) and DNA-based analyses (Johnson et al. 2004).
Lynx rufus
E: Bobcat; F: Lynx roux, lynx bai; G: Rotluchs, Luchskatze; Sp: Lince
rojo.
Wozencraft (2005) recognised twelve subspecies of Lynx rufus fol-
lowing Larivière and Walton (1997), Hall (1981) and Anderson (1987):
Lynx rufus rufus (Schreber, 1777a; pl. 109B, 1777b; 412).
Locality from where the species was first described: der Provinz Neu
York in America [= the Province of New York in America].
Type: None designated. Based on Pennant’s (1781; 281) Bay Lynx (Al-
len 1920).
Distribution: E and midwestern USA.
Lynx rufus baileyi (Merriam, 1890; 70, pl. 11).
Type locality: Moccasin Spring, Arizona [USA].
Holotype: USNM 186519 adult female skin and skull.
Distribution: SW arid zone from California to W Texas and Utah, and
S to Durango, Mexico.
Lynx rufus californicus (Mearns, 1897; 458).
Type locality: San Diego, California [USA].
Holotype: USNM 1588 adult female skin and skull.
Distribution: Nevada to C and S California.
Lynx rufus escuinapae J. A. Allen, 1903; 614.
Type locality: Escuinapa, Sinaloa, Mexico.
Holotype: AMNH M-14326 adult female skin and skull.
Distribution: C Mexico extending north along W coast to Sonora.
Lynx rufus fasciatus (Rafinesque, 1817; 46).
Locality from where the subspecies was first described: North-West
coast [USA].
Type: None designated. Based on Lewis and Clark’s description of
specimens obtained near the mouth of the Columbia, on Netul River
(now Lewis and Clark River) near Astoria, Oregon on 13 December
1805 (Allen 1814; 96, Bailey 1936, Young 1958; 137).
Distribution: Coastal forests from SW British Columbia, Canada to N
California, USA.
Lynx rufus floridanus (Rafinesque, 1817; 46).
Locality from where the subspecies was first described: Florida, Geor-
gia and Louisiana, restricted to Florida by Stark (1828; 103).
Type: None designated.
Distribution: SE USA.
Lynx rufus gigas (Bangs, 1897; 50, plate II).
Type locality: from fifteen miles back of Bear River, Nova Scotia
[Canada].
Holotype: MCZ 4951 male skin and skull.
Distribution: Maine and adjacent SE Canada, including Nova Scotia.
Lynx rufus oaxacensis Goodwin, 1963; 1.
Type locality: Los Nanches, San Pedro Jilotepec, District of Tehuantep-
ec, Oaxaca, Mexico.
Holotype: AMNH M-189300 female skin and skull.
Distribution: Uplands of C and S Oaxaca, from the districts of Tlaxlaco
and Ixtlan S to the districts of Yautepec and Tehuantepec; not known
to occur E of the Isthmus of Tehuantepec.
Lynx rufus pallescens (Merriam, 1899; 104).
Type locality: south base of Mount Adams, near Trout Lake, Washing-
ton [USA].
Holotype: USNM 76585 adult male skin and skull.
Distribution: Rocky Mountains from British Columbia, Canada to New
Mexico.
Lynx rufus peninsularis (Thomas, 1898; 42).
Type locality: Santa Anita, Lower California, Mexico.
Holotype: BMNH 1898.3.1.51 adult male skin and skull.
Distribution: Baja California, Mexico.
Lynx rufus superiorensis (Peterson and Downing, 1952; 1).
Type locality: McIntyre Township, near Port Arthur, Ontario [Canada].
Holotype: ROM 20947 male skin and skeleton.
Distribution: S Ontario, SE Manitoba to Wisconsin and Minnesota,
USA.
Lynx rufus texensis (Allen, 1895; 188); renaming of Felis maculata
Horsfield & Vigors, 1829; 381, pl. 13; type locality: Mexico (which is
preoccupied by Felis (Lynx) vulgaris maculatus Kerr, 1792; 157) and
Lynx rufus var. maculatus Audubon and Bachman, 1851; 295.
Type locality: Castroville, on the head waters of the Medina, in Texas
[USA].
Holotype of maculata: BMNH 1856.12.24.275 skin and skull.
Distribution: W Louisiana to E Texas and NE Mexico.
Discussion
Young (1958) identified 12 subspecies based on pelage coloration.
Hall & Kelson (1959) listed 11 subspecies. Samson (1979) confirmed
11 of Young’s 12 subspecies (L. r. oaxacensis was not included), us-
ing multivariate analyses of cranial characters. Hall (1981) refined
boundaries for all 12 subspecies. Werdelin (1981) found that skulls
of floridanus are morphometrically distinct from those of pallescens,
baileyi and californicus, but no other eastern subspecies were sam-
pled in this study. At the same time Read (1981) came to a much
© S. Kennerknecht/pumapix
A new taxonomy of the Felidae
39
bobcat
different conclusion, suggesting that there were far fewer valid sub-
species.
Recently, studies on the phylogeography and population history of
bobcats on a continental scale with genetic analyses have been per-
formed (Croteau 2009, Reding 2011). Reding (2011) analysed 1700
samples with 15 microsatellites and 1 KB of mtDNA sequence. The
primary signature involves a longitudinal cline with a transition or
suture zone along the Great Plains in the central USA. This diver-
gence was evident in both marker types. Significantly negative FS
values and unimodal mismatch distributions support a scenario of
post-glacial expansion from two disjunct Pleistocene refugia, which
were probably separated by the aridification of the Great Plains dur-
ing interglacial periods. Under the conservation criterion of recipro-
cal monophly on a DNA sequences-based tree (Moritz 1994), the
findings of Reding (2011) support two historically independent units
representing eastern and western bobcats. There were some unique
haplotypes found in the few Mexican samples that were analysed.
To clarify the status of Mexican bobcats, more rigorous sampling is
required.
Croteau (2009) came to a very similar conclusion with a much smaller
sample size. She identified, based on mtDNA analyses, two phylo-
geographical groups, western versus mid-western/eastern bobcats.
Loveless et al. (2016) modelled the geographical distribution of the
bobcat during the LGM and today and confirmed an east-west divi-
sion with the Great Plains being devoid of bobcats during the late
Pleistocene and acting as an ecological barrier even today along with
the Rocky Mountains. Skull morphometrics supported two glacial ref-
ugia with longer, thinner skulls in the northwest and shorter, broader
skulls in the southeast (Loveless et al. 2016).
We recognise the following subspecies:
Lynx rufus rufus (Schreber, 1777), including L. r. rufus, L. r. superio-
rensis, L. r. floridanus, L. r. gigas.
Distribution: E of the Great Plains, North America.
Lynx rufus fasciatus (Rafinesque, 1817) including L. r. pallescens,
L. r. baileyi, L. r. fasciatus, L. r. californicus, L. r. peninsularis, L. r. tex-
ensis.
Distribution: W of the Great Plains, North America.
The status of bobcats in Mexico (L. r. esquinapae and L. r. oaxacensis)
needs to be clarified.
References
Allen J. A. 1895. On the names of mammals given by Kerr in his ‘Animal King-
dom’, published in 1792. Bulletin of the American Museum of Natural
History 7, 179-192.
Allen J. A. 1903. A new deer and a new lynx from the State of Sinaloa, Mexi-
co. Bulletin of the American Museum of Natural History 19, 613-615.
Allen J. A. 1920. Note of Gueldenstaedt’s names of certain species of Felidae.
Journal of Mammalogy 1, 90-91.
Distribution of tentative subspecies of bobcat. Borders between
subspecies are speculative.
Species Morphology Genetics Biogeography Certainty Comments
Lynx rufus rufus ++ ++ ++
Lynx rufus baileyi + - -
Lynx rufus californicus + - -
Lynx rufus escuinapae + + ? Needs further research
Lynx rufus fasciatus ++ ++ ++ Distinctive western genetic clade
Lynx rufus floridanus + - -
Lynx rufus gigas + - -
Lynx rufus oaxacensis + + ? Needs further research
Lynx rufus pallescens + - -
Lynx rufus peninsularis + ? -
Lynx rufus superiorensis + - -
Lynx rufus texensis + - -
L. r. fasciatus
L. r. rufus
L. r. escuinapae?
L. r. oaxacensis?
CATnews Special Issue 11 Winter 2017
40
bobcat
Allen P. (Ed.) 1814. History of the expedition under the command of Captains
Lewis and Clark, to the sources of the Missouri thence across the Rocky
Mountains ad down the River Columbia to the Paciifc Ocean performed dur-
ing the years 1804-5-6 by order of the Government of the United States, vol.
2, p. 96. Philadelphia and New York: Bradford and Inskeep, and Inskeep.
Anderson E. M. 1987. A critical review and annotated bibliography of litera-
ture on the bobcat. Colorado Division of Wildlife, Special Report No. 62,
Colorado, USA.
Audubon J. J. & Bachman J. 1851; 295. The Quadrupeds of North America,
Vol. II. Audubon, New York.
Bailey V. 1936. The mammals and life zones of Oregon. North American Fauna
No. 55.
Bangs O. 1897. Notes on the lynxes of eastern North America, with descrip-
tions of two new species. Proceedings of the Biological Society of Wash-
ington 11, 47-51.
Croteau E. K. 2009. Population genetics and phylogeography of bocats (Lynx
rufus) using microsatellites and mitochondrial DNA. PhD Thesis, Southern
Illinois University, Carbondale, 134 pp.
Goodwin G. G. 1963. A new subspecies of bobcat (Lynx rufus) from Oaxaca,
Mexico. American Museum Novitates No. 2139, 1-7.
Hall E. R. 1981. The mammals of North America. 2nd edition. John Wiley and
Sons, New York.
Hall E. R. & Kelson K. R. 1959. The mammals of North America. Ronald Press,
New York.
Larivière S. & Walton L. R. 1997. Lynx rufus. Mammalian Species No.563, 1-8.
Loveless A. M., Reding D. M., Kapfer P. M. & Papes M. 2016. Combining eco-
logical niche modelling and morphology to assess the range-wide popu-
lation genetic structure of bobcats (Lynx rufus). Biological Journal of the
Linnean Society 117, 842-857.
Mearns E. A. 1898. Preliminary diagnoses of new mammals of the genera
Lynx, Urocyon, Spilogale and Mephitis from the Mexican Boundary Line.
Proceedings of the United States National Museum 20, 457-461.
Merriam C. H. 1890; 79 pl 11. Results of a biological survey of the San Fran-
cisco Mountain region and desert of the Little Colorado in Arizona. North
American Fauna 3, 78-86.
Merriam C. H. 1899; 104. Mammals of Shasta. North American Fauna 16, 87-
107.
Moritz C. 1994. Defining evolutionarily significant units for conservation.
Trends in Ecology and Evolution 9, 373-375.
Peterson R. L. & Downing S. C. 1952. Notes on the bobcats (Lynx rufus) of
eastern North America with the description of a new race. Contributions of
the Royal Ontario Museum Division of Zoology and Palaeontolgy 33, 1-23.
Pennant T. 1781. History of quadrupeds. Vol I. B. White, London.
Rafinesque C. S. 1817. Descriptions of seven new genera of North American
quadrupeds. The American Monthly Magazine and Critical Review 2, 44-46.
Read J. A. 1983. Geographic variation in the bobcat (Felis rufus) in the south-
central United States. MSc Texas A&M University, 115 pp.
Reding D. M. 2011. Patterns and processes of spatial genetic structure in a
mobile and continuously distributed species, the bobcat (Lynx rufus). PhD
Thesis, Iowa State University, Ames, Iowa. 195 pp.
Samson F. B. 1979. Multivariate analysis of cranial characters among bob-
cats with a preliminary discussion of the number of subspecies. Bobcat
Research Conference. National Wildlife Federation Science and Technical
Series 6, 80-86.
Schreber J. C. D. 1777a. Die Säugthiere in Abbildungen nach der Natur mit
Beschreibungen 3(25); pl.109B.
Schreber J. C. D. 1777b. Die Säugthiere in Abbildungen nach der Natur mit
Beschreibungen 3(24), 412.
Stark J. 1828. Elements of natural history, adapted to the present state of
the science, containing the generic characters of nearly the whole Animal
Kingdom and descriptions of the principal species. Blackwood, Edinburgh.
Thomas O. 1898. On new mammals from western Mexico and Lower Cali-
fornia. The Annals and Magazine of Natural History (7th series) 1, 40- 46.
Werdelin L. 1981. The evolution of lynxes. Annales Zoologici Fennici 18, 37-71.
Wozencraft W. C. 2005. Order Carnivora. In Mammal species of the world. A
taxonomic and geographic reference (3rd ed.). Wilson D. E. & Reeder D. M.
(Eds). Johns Hopkins University Press, Baltimore, pp. 532-628.
Young S. P. 1958. The bobcat of North America: Its history, life habits, eco-
nomic status and control, with list of currently recognized subspecies. The
Wildlife Management Institute, Washington D.C.
A bobcat in California (S. Kennerknecht/pumamix).
A new taxonomy of the Felidae
41
Canada lynx
Lynx canadensis
E: Canada lynx; F: Lynx du Canada, loup-cervier; G: Kanadaluchs; Sp:
Lince de Canada.
Wozencraft (2005) recognised three subspecies of Lynx canadensis:
Lynx canadensis canadensis Kerr, 1792; 157.
Locality from where the species was first described: Canada; restrict-
ed by Miller (1912) to Eastern Canada.
Type: None designated.
Distribution: Mainland Canada.
Lynx canadensis mollipilosus Stone, 1900; 48.
Type locality: Wainwright Inlet, Pt. Barrow [Alaska].
Holotype: ANSP 141 male skin and skull.
Distribution: Alaska.
Distinguishing characters: Browner and less grey than true Lynx
canadensis, with a very dense, soft, woolly pelage. Skull decidedly
narrower, higher and more arched than L. canadensis, and much more
constricted across the frontals and between the orbits, the postorbi-
tal processes are conspicuously more slender. Measurements: Total
length 1,040 mm; tail vertebra 130 mm; hind foot ca. 260 mm (Stone
1900).
Lynx canadensis subsolanus Bangs, 1897; 49.
Type locality: Codroy, Newfoundland [Canada].
Holotype: MCZ 1190 male skin and skull.
Distribution: Newfoundland, Canada.
Distinguishing characters: Differs from L. c. canadensis in darker and
richer colour, and some cranial characters.
Discussion
Van Zyll de Jong (1975) measured skulls of the Canada lynx from
across the range, including Newfoundland. He found that zygomatic
width, mastoid width, interorbital width, postorbital width and width
between the postorbital processes were significantly greater relative
to condylobasal length in L. c. subsolanus. But there was an overlap in
many other measures. He concluded that the separation was of rela-
tively recent date and that the subspecies status was questionable,
which means that the species would be monotypic.
Based on genetics, no evidence for isolation in different glacial refugia
within North America was found (Rueness et al. 2003). It seems that
the Canada lynx is genetically structured following ecological differen-
tiation due to large-scale climatic factors (Stenseth et al. 1999). Can-
ada lynx are known to disperse over very large distances, even up to
1,100 km (Poole 1997). This creates a high level of gene flow (Schwartz
et al. 2002). This was confirmed by Row et al. (2012). They analysed
samples from across mainland North America and Newfoundland and
found only little genetic differentiation among mainland Canada lynx,
but large differentiation between the mainland and Newfoundland.
This is to be expected for an island population. There is currently a
study going on which also looks into slower mutating genetic markers
and includes an attempt to estimate divergence time with different
markers (J. Row, pers. comm.).
There has been a lot of speculation about the colonization of New-
foundland by Canada lynx. Cameron (1958) said that they arrived early
in the post-glacial period, and Dodds (1960) concluded that they ar-
rived as late as 1861 in Newfoundland. Cameron’s hypothesis is very
unlikely to be true as lynx arrived late in North America and did not
make it south of the large ice sheet during the Pleistocene (Guthrie
1990, Pielou 1991) and so cannot have arrived on Newfoundland early
in the post-glacial period.
Therefore we conclude that Lynx canadensis is a monotypic species:
Lynx canadensis Kerr, 1792.
Distribution: Canada, Alaska and northern USA.
References
Bangs O. 1897. Notes on the lynxes of eastern North America, with descrip-
tions of two new species. Proceedings of the Biological Society of Wash-
ington 11, 47-51.
Cameron A.W. 1958. Mammals of the islands in the Gulf of St. Lawrence.
Superintendent of Publications, Queen’s Printer, Ottawa.
Dodds D. G. 1960. The economics, biology and management of the snowshoe
hare in Newfoundland. Dissertation, Cornell University.
Guthrie R. D. 1990. Frozen fauna of the mammoth steppe. University of Chi-
cago Press, Chicago.
Johnson W. E., Godoy J. A., Palomares F., Delibes M., Fernandes M., Revilla E.
& O’Brien S. J. 2004. Phylogenetic and phylogeographic analysis of Iberian
Lynx populations. Journal of Heredity 95, 19-28.
Kerr R. 1792. The Animal Kingdom or zoological system of the celebrated Sir
Charles Linnaeus. Class I. Mammalia: Containing a complete systematic
description, arrangement, and nomenclature, of all the known species and
varieties of the mammalia, or animals which give suck to their young, be-
ing a translation of that part of the Systema Naturae as lately published
with great improvements by Professor Gmelin of Goettingen together with
numerous additions from more recent zoological writers and illustrated
Species Morphology Genetics Biogeography Certainty Comments
Lynx canadensis canadensis ++ ++ ++
Lynx canadensis mollipilosus + - -
Lynx canadensis subsolanus + - + Probable human introduction
© highwaywilding.org
CATnews Special Issue 11 Winter 2017
42
Eurasian lynx
with copper plates. Printed for A. Strahan, and T. Cadell, London, and W.
Creech, Edinburgh.
Miller G. S. Jr. 1912. List of North American land mammals in the United
States National Museum, 1911. Bulletin of the United States National
Museum 79, 1-455.
Nowak R. M. 1999. Walker’s Mammals of the World, 6th edition. Johns Hop-
kins University Press, Baltimore.
Pielou E. C. 1991. After the Ice Age – the return of life to glaciated North
America. University of Chicago Press, Chicago.
Poole K. G. 1997. Dispersal patterns of lynx in the Northwest Territories. Jour-
nal of Wildlife Management 61, 497-505.
Row J. R., Gomez C., Koen E. L., Bowman J., Murray D. L. & Wilson P. J. 2012.
Dispersal promotes high gene flow among Canada lynx populations across
mainland North America. Conservation Genetics 13, 1259-1268.
Rueness E. K., Stenseth N. C., O’Donoghue M., Boutin S., Ellegren H. & Jakob-
sen K. S. 2003. Ecological and genetic spatial structuring in the Canadian
lynx. Nature 425, 69-72.
Schwartz M. K., Mills L. S., McKelvey K. S., Ruggiero L. F. & Allendorf F. W.
2002. DNA reveals high dispersal synchronizing the population dynamics
of Canada lynx. Nature 415, 520-522.
Stenseth N. C., Chan K.-S., Tong H., Boonstra R., Boutin S., Krebs C. J., Post E.,
O’Donoghue M., Yoccoz N. G., Forchhammer M. C. & Hurrell J. W. 1999.
Common dynamic structure of Canada lynx populations within three cli-
matic regions. Science 285, 1071-1073.
Stone W. 1900. Report on the birds and mammals collected by the McIlhenny
Expedition to Pt. Barrow, Alaska. Proceedings of the Academy of Natural
Sciences of Philadelphia 52, 4-49.
Van Zyll de Jong C. G. 1975. Differentiation of the Canada lynx, Felis (Lynx)
canadensis subsolana, in Newfoundland. Canadian Journal of Zoology 50,
699-705.
Wozencraft W. C. 2005. Order Carnivora. In Mammal species of the world. A
taxonomic and geographic reference (3rd ed.). Wilson D. E. & Reeder D. M.
(Eds). Johns Hopkins University Press, Baltimore, pp. 532-628.
Lynx lynx
E: Eurasian lynx; F: Lynx commun, lynx boréal; G: Luchs; Sp: Lince.
Wozencraft (2005) recognised five subspecies of Lynx lynx:
Lynx lynx lynx (Linnaeus, 1758; 43).
Locality from where the species was first described: Europae sylvis &
desertis; restricted by Thomas (1911; 136) to Wennersborg [= Väners-
borg], S. Sweden.
Type: None designated.
Distribution: From W Europe through the boreal forests of Scandinavia
and Russia.
Lynx lynx isabellinus (Blyth, 1847; 1178).
Type locality: Tibet.
Holotype: An imperfect skin collected by Blyth.
Distribution: Central Asian mountains (Pamir, Hindukush, Tian Shan),
Tibetan Plateau and south slopes of the Himalayas.
Distinguishing characters: Fur dense and soft, colour monochromatic,
grayish or white-brownish without spots or only faintly visible spots.
Size comparable to European lynx and therefore smaller than the other
lynx in Asia. The border between L. l. isabellinus and L. l. wardi is not
clear. It seems to be around the Irtysh Valley.
Lynx lynx kozlovi Fetisov, 1950; 21.
Type locality: Barun-Burinkhan, Selenginskiy region, Buryatskaya
ASSR, USSR [= Russia].
Holotype: Adult male skull No.80 in collection of Zoological Museum,
Irkutsk State University.
Distribution: C Siberia, from the Yenissei River to Lake Baikal.
Distinguishing characters: Somewhat smaller than Altai lynx, fur very
dense. Winter coat extremely diverse in general colour and degree of
spottiness.
Lynx lynx neglectus Stroganov, 1962; 408.
Heptner (1969; 1260) renamed Felis neglecta as Felis stroganovi,
because this name was preoccupied by Felis neglecta Gray, 1838;
27 (holotype BMNH 1838.4.16.325) = Caracal aurata. However, now
that this species is in the genus Lynx, the original name is available.
Type locality: Glazkovka, Suputinksiy Nature Reserve, Primorsk Terri-
tory, USSR [= Russia].
Holotype: ZMMU S-41310 adult male skull.
Distribution: Russian Far East, Ussuri and Amur territories, North Ko-
rea, NE China (Manchuria).
Distinguishing characters: Characterized mainly by size and to a lesser
extent craniological features. It occupies an intermediate position be-
tween the Baikal lynx and the Siberian lynx.
The subspecies status of stroganovi was challenged by Matjuschkin
(1978).
Lynx lynx sardiniae Mola, 1908b; 48.
Type locality: Nuoro, Sardinia.
Syntypes: Zoology Institute, Sassari University two mounted skins;
both now lost.
Distribution: Sardinia.
In 1908 Mola published two short articles in the Bollettino della So-
cietà Zoologica Italiana describing, firstly, a problematico incrocio di
Felidi (Mola 1908a), then a Lince della Sardegna (Mola 1908b). Both
descriptions were clearly and doubtlessly related to two Sardinian
wildcats (Felis lybica lybica). In November 1979, during his survey for
the preparation of the monograph on Felis lybica in Italy (Ragni 1981),
B. Ragni saw and examined one of Mola’s two mounted specimens in
© KORA
A new taxonomy of the Felidae
43
Eurasian lynx
the zoological collection of the Zoology Institute of Sassari University;
the animal was in a poor state of preservation, but was obviously Felis
lybica and not Lynx lynx (see photo). The other specimen was lost and
the cited one was destroyed a few years after his visit (G. Delitala,
curator of zoological collections, Sassari University, pers. comm.). Both
specimens were without skulls. A few years after Mola’s “discovery”
Prof. Alessandro Ghigi, a famous Italian zoologist, assessed the major
lapsus of the Sardinian student, affirming that the described Sardinian
lynx was, effectively, a Sardinian wildcat (Ghigi 1911).
The Sardinian wildcat, described by Mola (1908) as a lynx (Photo
G. M. Dalitalia).
Discussion
However, there is no broad consensus on the number of recognised
subspecies of Lynx lynx and their geographical distributions. Other
subspecies that have been recognised in recent years include:
Lynx lynx balcanicus Bureš, 1941; 51; junior synonym L. l. martinoi
Mirić, 1978.
Type locality: Šara Mts., the Republic of Macedonia.
Holotype: Skull in Zoo Skopji.
Distribution: The Balkan lynx is distributed in the south-west Balkans.
Albania, Macedonia, Kosovo, Montenegro and potentially Greece are
countries that are sharing this scattered and fragmented population.
Lynx lynx carpathicus Heptner in Heptner and Sludskii, 1972; 408.
Originally described by Kratochvíl & Štollmann, 1963; 315 in Štollmann
(1963) as Lynx lynx orientalis carpathicus, which is not available as a
valid subspecific name.
Type locality: Turie, Žilina District, Low Tatras, [Slovakia].
Holotype: No.131-62 adult male skull and skin, kept in Povážie mu-
seum, Žilina, Slovakia.
Distribution: Carpathian Mountains.
Distinguishing characters: Significantly larger skull size than other lynx
in Europe (Štollmann 1963, Hell 1980). Reddish and highly spotted fur
with little variability.
Lynx lynx dinniki Satunin, 1915; 391; Renaming of Lynx pardina
orientalis Satunin, 1905; 165, 323, preoccupied by Felis orientalis
Schlegel, 1857; 23 = Panthera pardus orientalis.
Type locality: Designated by Heptner & Sludskii (1972; 385) as “Psebai,
north-western parts of Greater Caucasus Range”, [Psebai, Krasnodar
Terrirory, Russia].
Lectotype: Skin from collection of N.Ya.Dinnik (Dinnik 1914) selected
by Heptner & Sludskii, 1972; 385.
Distribution: Caucasus Mountains S to Turkey, Iraq and Iran, formerly
also in the Kopet-Dag, Turkmenistan.
Distinguishing characters: Intermediate in size between Carpathian
and Balkan lynx. Fur short and sparse, which leaves a very slender
impression. Reddish fur with bright pattern of spots and stripes pre-
dominate. It was formerly considered to be a member of the species
Lynx pardina based on the spotted coat pattern.
Lynx lynx wardi (Lydekker, 1904; 576).
Type locality: Altai Mountains.
Holotype: BMNH 1904.10.14.1 skin.
Distribution: Altai Mountains (Russia, Kazakhstan, China and Mongo-
lia).
Distinguishing characters: Much larger in size than the neighbouring L.
l. kozlovi, L. l. lynx and L. l. isabellinus.
Lynx lynx wrangeli Ognev, 1928; 22.
Type locality: Valley of River Adycha, Hotan-Haia, Verkhoyansk Dis-
trict, Yakutia, Russia.
Holotype: ZIN 12692 male skull.
Distribution: from the Yenissei River E to the Pacific, S to the Stanovoy
Mountains.
Distinguishing characters: Skull larger than in any other group or sub-
species with the zygomata markedly broader, frontal region flat and
without longitudinal depression. Fur colour light and monochromatic
almost entirely without spots, only a few spots on the legs.
Lynx lynx melinus Kerr, 1792; 157.
Locality from where subspecies first described: on the banks of the
Volga, below Casan [= Kazan, Russia]
Type: None designated. Based on Pennant (1781; 279).
Distribution: Finland, European Russia, W Siberia (Ratkiewicz et al.
2014).
Despite the number of supposed subspecies a comprehensive phylo-
geography and morphological analysis has not been carried out, al-
though some regional studies have been conducted.
L. l. balcanicus is significantly smaller than other European lynx (Mirićć
1978), and has less dense fur with shorter hair (Mirićć1981). Prelimi-
nary genetic analyses have shown that the Balkan lynx is different
from other European lynx (Gugolz et al. 2008).
There has been much debate about the taxonomic status of L. l. car-
pathicus. While Vasiliu & Decei (1964) and Hemmer (1993) did not
recognise this subspecies, Matjuschkin (1978) and Heptner & Sludskii
(1972) had no doubt about its distinctiveness, which they considered
as one of the most distinctive forms in Eurasia. Hemmer (1993) argued
that the name carpathicus was not available as the subspecies had
been incorrectly described. There is increasing evidence for extra-
Mediterranean ice-age refugia in Europe (Schmitt 2007), including
the Carpathian Mountains, which were a forest refugium (Burga &
Perret 1998), which was very likely occupied by lynx. Recent genetic
analyses have shown that Carpathian lynx are isolated from the other
lynx populations and have low variability in the mtDNA control region
(Breitenmoser-Würsten et al. 2003, Gugolz et al. 2008).
Matjuschkin (1978) and Heptner & Sludskii (1972) considered L. l. din-
niki as a very morphologically distinct form, which has been geographi-
cally isolated for a long time. During the last Ice Age lynx lived south of
the Caucasus Mountains, which were covered by ice. Sea levels were
much lower, including that of the Black Sea, so that the Dardanelles,
Bosphorus and Marmara, were dry, creating a land bridge between
CATnews Special Issue 11 Winter 2017
44
Eurasian lynx
Asia Minor and the Balkans (Hewitt 1999). The geographical distribu-
tion of L. l. dinniki may have reached southeastern Europe and prelimi-
nary genetic analyses have shown that L. l. balcanicus and L. l. dinniki
share haplotypes (Gugolz et al. 2008) and are perhaps consubspecific.
The taxonomic status of the Asian lynxes is still poorly understood.
L. l. wrangeli is the largest form and shows the greatest sexual di-
morphism. Only 12% of L. l. wardi lack a metaconid on m1, whereas
75% of L. l. isabellinus individuals lack a metaconid on m1, which is
much higher than in any other population. The taxonomic status of
lynx in the Altai, the Baikal region and the Amur region needs further
investigation and clarification. C. Groves (pers. comm.) considers L. l.
neglectus as distinct because this area of the Russian Far East and
northern Manchuria is bioegographically distinct.
Rueness et al. (2014) have carried out a phylogeographical analysis
of lynx throughout Eurasia excluding the Himalayas. Based on both
mtDNA and microsatellites, they identified three main clades (west-
ern, eastern and southern), which do not appear to correspond to pu-
tative subspecies. There is clearly more research required with more
extensive sampling in order to elucidate the phylogeography and sub-
specific variation of Lynx lynx.
On the basis of current evidence we propose the following six subspe-
cies, although this number may be further reduced in the future:
Lynx lynx lynx (Linnaeus, 1758), including melinus.
Distribution: Scandinavia, Finland, Baltic States, Belarus, European
part of Russia E to the Yenissei River.
Lynx lynx balcanicus (Bureš, 1941).
Distribution: The Balkans and possibly Greece; possibly a synonym of
L. l. dinniki.
Lynx lynx carpathicus Heptner, 1972.
Distribution: E and C Europe.
Lynx lynx dinniki Satunin, 1915.
Distribution: The Caucasus, Asia Minor, Iran, Iraq.
Lynx lynx isabellinus (Blyth, 1847), including kamensis.
Distribution: C Asia including the Himalayas and Tibet.
Lynx lynx wrangeli Ognev, 1928, including kozlovi, neglectus, wardi?
Distribution: E of the Yenissei River to China??
References
Blyth E. 1847. Report of Curator, Zoological Department, September 1847.
Journal of the Asiatic Society of Bengal 16, 1176-1181.
Bureš I. 1941. Risove v Makedonija. Priroda 42, 51ć52.
Burga C. A. & Perret R. 1998. Vegetation und Klima der Schweiz seit dem
jüngeren Eiszeitalter. Ott Verlag, Thun.
Breitenmoser-Würsten Ch., Binns M., Johnson W., Breitenmoser U., Gaillard
C. & Obexer-Ruff G. 2003. Population and conservation genetics of two re-
introduced lynx (Lynx lynx) populations in Switzerland – a molecular evalu-
ation 30 years after the first translocations. Mammalian Biology Special
issue 68, 15.
Dinnik N. Ya. 1914. Zveri Kavkaza. T.2, Khishchnye. [Mammals of Caucasus.
pt.2, Carnivora]. Tipografiya K. P. Kozlovskogo, Tiflis.
Fetisov A. S. 1950. [A new subspecies of lynx (Lynx lynx kozlovi subsp. n.)
from eastern Siberia]. Izvestiya Biologo-Geograficheskogo Nauchno-
Issledovatel’skogo Instituta pri Irkutskom Gosudarstvennom Universitete
imeni A. A. Zhdanova, Irkutsk, 12, 21-22.
Ghigi A. 1911. Ricerche faunistiche e sistematiche sui Mammiferi d’Italia che
formano oggetto di caccia. Natura, Milano, II, 7-48.
Species Morphology Genetics Biogeography Certainty Comments
Lynx lynx lynx ++ ++ ++
Lynx lynx balcanicus + ++ + Possibly part of dinniki
Lynx lynx carpathicus ++ ++ +
Lynx lynx dinniki ++ ++ ++
Lynx lynx isabellinus ++ o ++
Lynx lynx kozlovi + o o
Lynx lynx neglectus + o ?+ May be distinct subspecies?
Lynx lynx wardi + o o
Lynx lynx wrangeli ++ + ++
Lynx lynx melinus + o o
Lynx lynx sardiniae - - - Misidentified; actually Felis lybica
Distribution of tentative subspecies of Eurasian lynx. Borders be-
tween subspecies are speculative.
L. l. lynx L. l. wrangeli
L. l. isabellinus
L. l. dinniki
L. l. carpathicus
L. l. balcanicus
A new taxonomy of the Felidae
45
Iberian lynx
Gray J. E. 1838. On some new species of quadrupeds and shells. Annals of
Natural History or Magazine of Zoology, Botany and Geology 1, 27-30.
Gugolz D., Bernasconi M. V., Breitenmoser-Würsten Ch. & Wandeler P. 2008.
Historical DNA reveals the phylogenetic position of the extinct Alpine lynx.
Journal of Zoology 275, 201-208.
Hell P. 1980. Ergebnisse der Luchsforschung in der CSSR: II.Teil - ueber
die Variabilitaet der Schaedel- und Koerpermasse des Luchses Felis
lynx in den Westkarpaten. Beiträge zur Jagd- und Wildforschung 11,
260-275.
Hemmer H. 1993. Felis (Lynx) lynx Linnnaeus, 1758 – Luchs, Nordluchs. In
Handbuch der Säugetiere Europas 5/II (Raub säuger 2). Niethammer J. &
Krapp F. (Eds). AULA, Wiesbaden, pp.1119-1167
Heptner V. G. 1969. On systematics and nomenclature of Palearctic cats. Zoo-
logicheskii Zhurnal 48, 1258-1260.
Heptner V. G. & Sludskii A. A. 1972. Mammals of the Soviet Union. Vol. II,
part 2. Carnivora (hyaenas and cats). Vysshaya Shkola, Moscow. [English
translation published in 1992, Smithsonian Institution Libraries, Washing-
ton D.C.]
Hewitt G. M. 1999. Post-glacial re-colonization of European biota. Biological
Journal of the Linnean Society 68, 87-112.
Horsfield T. & Vigors N. A. 1829. Observations on some of the Mammalia con-
tained in the collection of the Zoological Society. The Zoological Journal
4, 380- 384.
Linnaeus C. 1758. Systema Naturae per regna tria naturae, secundum classis,
ordines, genera, species cum characteribus, differentiis, synonymis, locis.
10th edition, vol. 1. Laurentii Salvii, Holmiae.
Lydekker R. 1904. The coloration of the lynxes. The Field 104, 576.
Matjuschkin E. N. 1978. Der Luchs. A. Ziemsen, Wittenberg Lutherstadt.
Mirić D. 1978. Lynx lynx martinoi ssp. nova – neue Luchsunterart von der Bal-
kanhalbinsel. Bulletin of the Museum Hist. Nat. 33, 29-36.
Mirić D. 1981. The lynx populations of the Balkan Peninsula. Serbian Academy
for Sciences and Art, Belgrade.
Mola P. 1908a. Considerazioni sopra un problematico incrocio di Felidi. Bollet-
tino della Società Zoologica Italiana, Roma, Sez. 2, 9, 42-45.
Mola P. 1908b. Ancora della Lince della Sardegna. Bollettino della Società
Zoologica Italiana, Roma, Sez. 2, 9, 46-48.
Ognev S. I. 1928. Rysi (Lynx). Okhotnik, 5-6.
Pennant T. 1781. History of quadrupeds. Vol I. B. White, London.
Ragni B. 1981. Gatto selvatico. Felis silvestris Schreber, 1777. In Distribuzione
e biologia di 22 specie di Mammiferi in Italia. Pavan M. (a cura di) C.N.R.,
Rome, pp. 105-113.
Rueness E. K., Naidenko S., Trosvik P. & Stenseth N. C. 2014. Large-scale ge-
netic structuring of a widely distributed carnivore – the Eurasian lynx (Lynx
lynx). PLoS ONE 9(4): e93675.
Satunin K. A. 1905. Die Säugetiere des Talyschgebietes und der Mughan-
steppe. Mlekopitayushchie Talysha i Mugani. Mitteilungen des Kaukasis-
chen Museums - Izvěstiya Kavkazskago Muzeya 2, 87-402.
Satunin K. A. 1915. Mammalia Caucasica, tome 1 (Chiroptera, Insectivora et
Carnivora). Mémoires du Musée du Caucase, series A, I, 1-410.
Schmitt T. 2007. Molecular biogeography of Europe: Pleistocene cycles and
postglacial trends. Frontiers in Zoology 4, 11.
Štollmann A. 1963. Príspevok k pozunaiu rysa ostrovida Lynx lynx (L.) v
Českolovenských Karpátoch. Zoologické Listy 12, 301-316.
Stroganov S. U. 1962. Zveri Sibiri: Khishchnye. Moskva: Izd-vo Akademii nauk
SSSR.
Vasiliu G. D. & Decei, P. (1964). Über den Luchs Lynx lynx in den rumänischen
Karpaten. Säugetierkundliche Mitteilungen 12, 155-183.
Wozencraft W. C. 2005. Order Carnivora. In Mammal species of the world. A
taxonomic and geographic reference (3rd ed.). Wilson D. E. & Reeder D. M.
(Eds). Johns Hopkins University Press, Baltimore, pp. 532-628.
Lynx pardinus
E: Iberian lynx; F: Lynx pardelle, lynx d’Espagne; G: Pardelluchs; Sp:
Lince iberico.
Lynx pardinus (Temminck, 1827; 116).
Type locality: Portugal, puisque le commerce reçoit des peaux pre-
pares de Lisbonne, et que M. le baron de Vionénil tua, en 1818, sur les
bords du Tage, à dix lieues de Lisbonne [= on the banks of the River
Tagus {Portugal}, ten leagues from Lisbon].
Holotype: MNHN-ZM-MO-2001-325 mounted skin (skull inside; Al-
maça 1992).
Distribution: Iberian Peninsula.
Discussion
This is a monotypic species, which was formerly included in Lynx lynx
(Johnson et al. 2004). No subspecies are recognised.
References
Almaça C. 1992. Name, authorship, type specimen, and type locality of the
Iberian lynx. Mammalia 56, 659-662.
Johnson W. E., Godoy J. A., Palomares F., Delibes M., Fernandes M., Revilla E.
& O’Brien S. J. 2004. Phylogenetic and phylogeographic analysis of Iberian
lynx populations. Journal of Heredity 95, 19-28.
Temminck C. J. 1827. Monographies de mammalogie, ou description de quel-
ques genres de mammifères, dont les espèces ont été observées dans les
différens musées de l’Europe, vol. 1. C. C. Vander Hoek, Leiden.
Species Morphology Molecular Biogeography Certainty Comments
Lynx pardinus ++ ++ ++ Distinct monotypic species
© A. Sliwa
CATnews Special Issue 11 Winter 2017
46
Andean cat
Ocelot lineage
Genus Leopardus Gray, 1842; 260.
This genus comprises eight species that represent an adaptive radia-
tion in South and Central America. Characterised by a chromosome
count of 2n = 36 compared with 2n = 38 in other felids. Hybridisation
between species has been documented (Trigo et al. 2008, 2013).
Leopardus jacobita
E: Andean mountain cat; F: Chat des Andes; G: Bergkatze; Sp: Chin-
chay, gato andino, gato lince.
This species is regarded as monotypic (Wozencraft 2005):
Leopardus jacobita (Cornalia, 1865; 1).
Type locality: Bolivia, circa Potosi et Humacuaca in montibus sat
elevatis; further defined by Cabrera (1958; 297) as “Sur del depar-
tamento boliviano de Potosi, cerca de la frontera argentina, entre
Potosi y Humahuaca” [= southern Bolivian province of Potosi, near
the Argentinian border, between Potosi and Humahuaca].
Holotype: MSNM Ma 143 mounted skin; specimen lost to bombing in
1943 (Mann 1945; Gippoliti 2005).
Distribution: S Peru, NE Chile, SW Bolivia and NW Argentina.
Discussion
Genetic analysis of mitochondrial DNA (459 bp control region, 789
bp ND5, ATP-8 and 16S mitochondrial genes) and 11 nuclear micro-
satellites revealed two Evolutionary Significant Units ESUs that are
separated between latitudes 26° and 35° S (Cossío et al. 2012), with
the possibility that the northern ESU can be split into two Manage-
ment Units. With a greatly enhanced number of samples Ruiz-Garcia
et al. (2013) found that four populations of L. jacobita (northern Peru;
southern Peru – northern Bolivia; southern Bolivia – northern Ar-
gentina; Mendoza, Argentina) were more genetically isolated from
each other on the basis of microsatellites than between putative
subspecies of L. colocolo. However, morphological samples were not
sufficient to confirm whether these populations represent distinct
subspecies. Further research is urgently required, owing to the en-
dangered status of this species.
Therefore, we continue to recognise L. jacobita as a monotypic spe-
cies until further evidence is available:
Leopardus jacobita (Cornalia, 1865).
Distribution: Andes from S Argentina and Chile to Boliva and C Peru;
distrubution fragemented.
References
Cabrera A. 1958. Catalogo del los mamiferos de America del Sur. Revista
del Museo Argentino de Ciencias Naturales “Bernardino Rivadavia”e
Instituto Nacional de Investigacion de las Ciéncias Naturales, Ciéncias
Zoológicas 4, 1-307.
Cornalia E. 1865. Descrizione di una nuova specie del genere: Felis. Felis jac-
obita (Corn.). Memorie della Societá Italiana di Scienze Naturali 1, 1-9.
Cossíos E. D., Walker R. S., Lucherini M., Ruiz-García M. & Angers B. 2012.
Population structure and conservation of a high-altitude specialist, the An-
dean cat Leopardus jacobita. Endangered Species Research 16, 283-294.
Gippoliti S. 2005. Historical museology meets tropical biodiversity conserva-
tion. Biodiversity and Conservation 14, 3127-3134.
Mann G. 1945. Mamíferos de Tarapacá. Observaciones realizadas durante
una expedición al norte de Chile. Biológica (Santiago) 2, 23-98.
Ruiz-García M., Cossíos D., Lucherini M., Yáñez J., Pinedo-Castro M. & An-
gers B. 2013. Population genetics and spatial structure in two Andean
cats (the pampas cat, Leopardus pajeros, and the Andean mountain cat,
L. jacobita) by means of nuclear and mitochondrial markers and some
notes on biometrical markers. In Molecular population genetics, evolu-
tionary biology and biological conservation on Neotropical carnivores,
Ruiz-García M. & Shostell J. M. (Eds). Nova, New York , pp.187-244.
Trigo T. C., Freitas T. R. O., Kunzler G., Cardoso L., Silva J. C. R., Johnson W.
E., O’Brien S. J., Bonatto S. L. & Eizirik E. 2008. Inter-species hybridiza-
tion among Neotropical cats of the genus Leopardus, and evidence for an
introgressive hybrid zone between L. geoffroyi and L. tigrinus in southern
Brazil. Molecular Ecology, 17, 4317-4333.
Trigo T. C., Schneider A., de Oliveira T. G., Lehugeur L. M., Silveira L., Freitas
T. R. O. & Eizirik E. 2013. Molecular data reveal complex hybridization
and a cryptic species of Neotropical wild cat. Current Biology 23, 2528-
2533.
Wozencraft W. C. 2005. Order Carnivora. In Mammal species of the world. A
taxonomic and geographic reference (3rd ed.). Wilson D. E. & Reeder D.
M. (Eds). Johns Hopkins University Press, Baltimore, pp. 532-628.
Species Morphology Molecular Biogeography Certainty Comments
Leopardus jacobita ++ ++ ++ May comprise four or more subspecies
or even species
© J. Sanderson
A new taxonomy of the Felidae
47
ocelot
Leopardus pardalis
E: Ocelot; F: Ocelot; G: Ozelot; Sp: Ocelote, gato onza, gato tigre, man-
igordo, pumilio, tigrillo, cunaguaro.
Wozencraft (2005) listed the following subspecies:
Leopardus pardalis pardalis (Linnaeus, 1758; 42).
Locality from where species was first described: America, restricted to
State of Vera Cruz, Mexico by Allen (1919).
Type: None designated.
Distribution: Mexico to Panama.
Leopards pardalis aequatorialis (Mearns, 1903; 246).
Type locality: Paramba, northern Ecuador.
Holotype: USNM 113267 adult female skin and skull.
Distribution: NE South America.
Leopardus pardalis albescens (Pucheran, 1855; 149).
Type locality: Arkansas [USA].
Holotype: MNHN-ZM-MO-2001-323 male mounted skin.
Distribution: E Texas, USA S to Tamaulipas, Mexico.
Leopardus pardalis melanurus (Ball, 1844; 128).
Type locality: Unknown, but probably British Guiana [= Guyana] (Pocock
1941b; 328-333).
Holotype: BMNH 1855.12.24.251 adult probably male skin and skull.
Distribution: Venezuela, Guyana, Surinam and probably Trinidad.
Leopardus pardalis mitis (F. G. Cuvier, 1820; 221).
Type locality: Unknown, but restricted to Rio de Janeiro [Brazil] by Jar-
dine (1834; 194).
Syntypes: MNHN-ZM-MO-2001-319 and MNHN-ZM-MO-2001-320
mounted skins.
Distribution: C and E Brazil, Paraguay and N Argentina.
Leopardus pardalis pseudopardalis (Boitard, 1842; 263).
Type locality: Mexique et la baie de Campèche?, emended to Carta-
gena [Colombia] by Cabrera (1958; 284).
Holotype: MNHN-ZM-AC-A1753 juvenile part skull.
Distribution: N Colombia and Venezuela.
Leopardus pardalis pusaeus (Thomas, 1914; 347).
Type locality: Congon, 15 miles W. of Guayaquil, Ecuador.
Holotype: BMNH 1899.8.1.29 male skin and skull.
Distribution: SE Ecuador to NE Peru.
Leopardus pardalis sonoriensis (Goldman, 1925; 123).
Type locality: Camoa, Rio Mayo, Sonora, Mexico.
Holotype: USNM 96216 adult male skin and skull.
Distribution: Arizona, USA and Sonora, Mexico.
Leopardus pardalis steinbachi (Pocock, 1941a; 235).
Type locality: Buenavista, Santa Cruz, Bolivia.
Holotype: BMNH 1928.2.9.15 adult male skin and skull.
Distribution: C Bolivia.
Discussion
Wozencraft (2005) followed the arrangement by Cabrera (1958) and
Murray & Gardner (1997). Eizirik et al. (1998) examined the mitochon-
drial control region in ocelots throughout their range and identified
four phylogeographical groups in Central America, northwestern South
America, northeastern South America and southern South America
south of the Amazon.
Ruiz-Garcia et al. (2013) examined craniometric variation and micro-
satellite diversity in ocelots from throughout their range, except for the
southern most part of the range, most samples were from Colombia.
Microsatellite differentiation identified three groups; Texas, Central
America and South America (excluding eastern Brazil), whereas crani-
ometric variation revealed that animals from Bolivia were significantly
smaller.
Nascimento (2010) carried out a traditional morphological study of
ocelots and concluded that there are two species. The Central Ameri-
can ocelot, Leopardus pardalis, is smaller and has a greyer pelage than
the South American ocelot, Leopardus mitis, which is larger and has
a brighter, yellower pelage. Nascimento (2010) suggests that these
two species are sympatric in Nicaragua and Costa Rica, supporting
their distinction as separate species. However, given high individual
morphological variability within populations, this situation needs to
be examined in more detail including more detailed morphological and
molecular analyses.
Following Eizirik et al. (1998), up to four subspecies can be differen-
tiated:
Leopardus pardalis pardalis
Leopardus pardalis ssp. [traditionally part of L. p. mitis]
Leopardus pardalis pseudopardalis
Leopardus pardalis mitis
Following Ruiz-Garcia et al. (2013), up to four different subspecies can
be differentiated:
Leopardus pardalis pardalis
Leopardus pardalis albescens
Leopardus pardalis steinbachi
Leopardus pardalis mitis
However, the geographical delimitation of different genetic clades is
not completely congruent between these two molecular studies. The
morphological differentiation between Central and South American
forms is clear and supported partly by molecular data as well as a
clear biogeographical barrier, the Andes.
© P. Meier
CATnews Special Issue 11 Winter 2017
48
ocelot
Provisionally, two subspecies are recognised:
Leopardus pardalis pardalis (Linnaeus, 1758).
Distribution: from Texas and Arizona south to Costa Rica.
Distinguishing characters: Smaller and greyer than mitis.
Leopardus pardalis mitis (Cuvier, 1820).
Distribution: South America as far south as northern Argentina; limit of
range with respect to L. p. pardalis is unclear.
Distinguishing characters: Larger and yellower than pardalis.
References
Allen J. A. 1919. Notes on the synonymy and nomenclature of the smaller
spotted cats of tropical America. Bulletin of the American Museum of
Natural History 41, 341-419.
Ball R. 1844. Description of the Felis melanura. Proceedings of the Zoological
Society of London 12, 128-129.
Boitard M. 1842. Jardin des Plantes. Description et moeurs des mammifères
de la Ménagerie et du Muséum d’Histoire Naturelle. J. J. Dubochet, Paris.
Cabrera A. 1958. Catalogo del los mamiferos de America del Sur. Revista del
Museo Argentino de Ciencias Naturales “Bernardino Rivadavia” e Instituto
Nacional de Investigacion de las Ciéncias Naturales, Ciéncias Zoológicas
4, 1-307.
Cuvier F. G. 1820. Le chati femelle. In Histoire naturelle des mammifères. Geof-
froy St.-Hilaire E. & Cuvier F. (Eds), 1, XVIII: 1-3, Pl. 54.
Eizirik E., Bonatto S. L., Johnson W. E., Crawshaw Jr. P. G., Vié J. C., Brousset
D. M., O’Brien S. J. & Salzano F. M. 1998. Phylogeographic patterns and
evolution of the mitochondrial DNA control region in two Neotropical cats
(Mammalia, Felidae). Journal of Molecular Evolution 47, 613-624.
Goldman E. A. 1925. Two new ocelots from Mexico. Journal of Mammalogy
24, 122-124.
Jardine Sir W. 1834. The Naturalist’s Library. Mammalia, Volume II, the Feli-
nae. Lizars, Edinburgh.
Linnaeus C. 1758. Systema Naturae per regna tria naturae, secundum classis,
ordines, genera, species cum characteribus, differentiis, synonymis, locis.
10th edition, vol. 1. Laurentii Salvii, Holmiae.
Mearns E. A. 1903. The ocelot cats. Proceedings of the United States National
Museum 25, 237-249.
Murray J. L. & Gardner G. L. 1997. Leopardus pardalis. Mammalian Species
No. 548, 1-10.
Nascimento F. O. do 2010. Revisão taxonômica gênero do Leopardus Gray,
1842 (Carnivora, Felidae). PhD thesis, University of Sao Paulo, Brazil.
Pocock R. I. 1941a. Some new geographical races of Leopardus, commonly
known as ocelots and margays. Annals and Magazine of Natural History
(11th series) 8, 234-239.
Pocock R. I. 1941b. The races of the ocelot and margay. Field Museum of Natu-
ral History 27, 319-369.
Pucheran J. 1855. Description du chat bai et du chat albescent; et remarques
sur les caractères et sur la distribution géographique de plusieurs autre
chats, pp. 137-155. In Mammifères. Geoffroy St Hilaire I. Voyage autour
du monde sur la frégate la Vénus commandée par Abel du Petit-Thouars.
Zoologie. G & J. Baudry, Paris, pp. 1-176.
Ruiz-García M., Corrales C. & Pineda-Castro M. 2013. Craniometric and micro-
satellite genetic differentiation among putative ocelot subspecies (Leop-
ardus pardalis). In Molecular population genetics, evolutionary biology
and biological conservation on Neotropical carnivores. Ruiz-García M. &
Shostell J. M. (Eds). Nova, New York, pp. 289-332.
Thomas O. 1914. On various South-American mammals. Annals and Magazine
of Natural History (8th series) 13, 345-363.
Wozencraft W.C. 2005. Order Carnivora. In Mammal species of the world. A
taxonomic and geographic reference (3rd ed.). Wilson D. E. & Reeder D. M.
(Eds). Johns Hopkins University Press, Baltimore, pp. 532-628.
Subspecies Morphology Molecular Biogeography Certainty Comments
Leopardus pardalis pardalis ++ ++ ++
Leopardus pardalis albescens + + o
Leopardus pardalis sonoriensis o o o
Leopardus pardalis mitis ++ ++ ++ mtDNA suggests might be two
subspecies
Leopardus pardalis melanurus + - o
Leopardus pardalis aequatorialis + - o
Leopardus pardalis pseudopardalis + ++ o
Leopardus pardalis pusaeus + o o
Leopardus pardalis steinbachi + o o
Distribution of tentative subspecies of ocelot. Borders between
subspecies are speculative.
L. p. pardalis
L. p. mitis
A new taxonomy of the Felidae
49
margay
Leopardus wiedii
E: Margay; F: Margay; G: Langschwanzkatze, Margay; Sp: Tigrillo,
margay, caucal, gato tigre.
Wozencraft (2005) recognised 11 subspecies, following de Oliveira
(1998):
Leopardus wiedii wiedii (Schinz, 1821; 235).
Type locality: Brasilien [= Brazil]; Brasil, restringida al Morro de Arará,
sobre el río Mucurí, estado de Baía [= Brazil, restricted to the Morro de
Arara, on the Mucuri river, state of Baia] by Cabrera (1958; 290) and to
northern Espirito Santo, Brazil by Allen (1919; 357), both of which are
based on Wied (1826; 378).
Lectotype: RMNH.MAM 17695 (original register no. c) skin (Avila-
Pires 1965).
Distribution: S Brazil W to S Colombia.
Leopardus wiedii amazonicus (Cabrera, 1917; 28).
Type locality: Tabatinga [Amazonas, Brasil].
Holotype: MNCN 774 adult female skin and skull.
Distribution: Amazonas, Brazil.
Leopardus wiedii boliviae Pocock, 1941; 237.
Type locality: Buena Vista, Santa Cruz, Bolivia, 300 m. alt.
Holotype: BMNH 1926.1.5.4 adult male skin and skull.
Distribution: Bolivia to N Argentina.
Leopardus wiedii pirrensis (Goldman, 1914; 4).
Type locality: Cana (altitude 2,000 feet), eastern Panama.
Holotype: USNM 179162 adult female skin and skull.
Distribution: Panama.
Leopardus wiedii vigens (Thomas, 1904; 192).
Type locality: Igarapé-Assu, near Pará [Brazil]. Alt. 50 m.
Holotype: BMNH 1904.7.4.43 & 1904.7.4.43a adult male skin and skull.
Distribution: Lower Amazon, Brazil, Guianas.
Leopardus wiedii glauculus (Thomas, 1903; 235).
Type locality: Beltran, Jalisco, Mexico.
Holotype: BMNH 1890.1.4.1 adult female skin and skull.
Distribution: Jalisco, Sinaloa and N Yucatan, Mexico.
Leopardus wiedii nicaraguae (Allen, 1919; 357).
Type locality: Volcan de Chinandego, Nicaragua.
Holotype: AMNH M-28957 adult male skin and skull.
Distribution: Nicaragua.
Leopardus wiedii oaxacensis (Nelson and Goldman, 1931; 303);
pre-dated by Felis mexicana Saussure 1860; 3, but name preoccupied
by F. mexicana Desmarest 1816.
Type locality: Cerro San Felipe, Oaxaca, Mexico (altitude 10,000 feet).
Holotype: USNM 68169 adult male skin and skull.
Distribution: High mountains of Oaxaca, Mexico.
Leopardus wiedii salvinia Pocock, 1941; 239.
Type locality: Vera Paz, Guatemala.
Holotype: BMNH 1875.2.27.1 adult male skin and skull.
Distribution: Guatemala and perhaps Belize.
Leopardus wiedii yucatanicus (Nelson and Goldman, 1931; 304).
Type locality: Merida, Yucatan.
Holotype: USNM 8612 subadult female skin and skull.
Distribution: Yucatan Peninsula and N Chiapas, Mexico.
Leopardus wiedii cooperi (Goldman, 1943; 384).
Type locality: Eagle Pass, Texas.
Holotype: USNM 25 adult male skin.
Distribution: Texas, USA and NE Mexico.
Discussion
A study of mtDNA among margays by Eizirik et al. (1998) revealed
three phylogeographical groups; South America, south of the Amazon;
South America, north of the Amazon; and Central America. However,
a more recent analysis of skins and skulls by Nascimento (2010) failed
to find any significant geographical variation, although Central Ameri-
can margays appear to be smaller and greyer compared with South
American animals. A more comprehensive molecular and morphologi-
cal study is required. Until then, we recognise three subspecies as an
interim conservative arrangement based on Eizirik et al. (1998):
Leopardus wiedii wiedii (Schinz, 1821).
Distribution: South America S of the Amazon.
Leopardus wiedii vigens (Thomas, 1904).
Distribution: South America N of the Amazon.
Leopardus wiedii glauculus (Thomas, 1903).
Distribution: C America.
References
Allen J. A. 1919. Notes on the synonymy and nomenclature of the smaller
spotted cats of tropical America. Bulletin of the American Museum of
Natural History 41, 341-419.
de Avila-Pires F. D. 1965. The type specimens of Brazilian mammals collected
by Prince Maximilian zu Wied. American Museum Novitates No. 2209,
1-21.
Cabrera Á. 1917. Mamíferos del Viaje al Pacífico verificado de 1862 a 1865 por
una Comisión de Naturalistas enviada por el Gobierno Español. Trabajos
del Museo Nacional de Ciencias Naturales. Serie Zoológica 31, 1-62.
Cabrera A. 1958. Catalogo del los mamiferos de America del Sur. Revista del
Museo Argentino de Ciencias Naturales “Bernardino Rivadavia”e Instituto
Nacional de Investigacion de las Ciéncias Naturales, Ciéncias Zoológicas
4, 1-307.
© A. Sliwa
CATnews Special Issue 11 Winter 2017
50
margay
Eizirik E., Bonatto S. L., Johnson W. E., Crawshaw Jr. P. G., Vié J. C., Brousset
D. M., O’Brien S. J. & Salzano F. M. 1998. Phylogeographic patterns and
evolution of the mitochondrial DNA control region in two Neotropical cats
(Mammalia, Felidae). Journal of Molecular Evolution 47, 613-624.
Goldman E. A. 1914. Descriptions of five new mammals from Panama. Smith-
sonian Miscellaneous Collections 63, 1-7.
Goldman E. A. 1943. The races of ocelot and margay in Middle America. Jour-
nal of Mammalogy 24, 372-385.
Nascimento do F. O. 2010. Revisão taxonômica gênero do Leopardus Gray,
1842 (Carnivora, Felidae). PhD thesis University of Sao Paulo, Brazil.
Nelson E. W. & Goldman E. A. 1931. New carnivores and rodents from Mexico.
Journal of Mammalogy 12, 302-306.
Oliveira de T. G. 1998. Leopardus wiedii. Mammalian Species No. 579, 1-6.
Pocock R. I. 1941. Some new geographical races of Leopardus, commonly
known as ocelots and margays. Annals and Magazine of Natural History
(11th series) 8, 234-239.
de Saussure H. 1860; 3. Note sur quleques mammifès du Mexique. Revue et
Magasin de Zoologie pure et appliqué, 2nd series, 12, 3-11.
Schinz H. R. 1821. Das Thierreich eingetheilt nach dem Bau der Thiere als
Grundlage ihrer Naturgeschichte und der vergleichenden Anatomie von
den Herrn Ritter von Cuvier. Erster Band, Säugethiere und Vögel. J. G.
Gotta‘sche Buchhandlung, Stuttgart und Tübingen.
Thomas O. 1903. Notes on Neotropical mammals of the genera Felis, Hapale,
Oryzomys, Akodon and Ctenomys, with descriptions of new species. An-
nals and Magazine of Natural History (7th series) 12, 234-243.
Thomas O. 1904. New Callithrix, Midas, Felis, Rhipidomys and Proechimys
from Brazil and Ecuador. The Annals and Magazine of Natural History (7th
series) 14, 188-196.
Distribution of tentative subspecies of margay. Borders between
subspecies are speculative.
Wied zu M. 1826. Beiträge zur Naturgeschichte von Brasilien. Band II. Gr. H. S.
priv. Landes-Industrie-Comptoirs, Weimar.
Wozencraft W. C. 2005. Order Carnivora. In Mammal species of the world. A
taxonomic and geographic reference (3rd ed.). Wilson D. E. & Reeder D. M.
(Eds). Johns Hopkins University Press, Baltimore, pp. 532-628.
Subspecies Morphology Molecular Biogeography Certainty Comments
Leopardus wiedii wiedii ++ ++ ++
Leopardus wiedii amazonicus + o o
Leopardus wiedii boliviae + o o
Leopardus wiedii pirrensis + o o
Leopardus wiedii vigens + + +
Leopardus wiedii glauculus + ++ ++
Leopardus wiedii nicaraguae + o o
Leopardus wiedii oaxacensis + o o
Leopardus wiedii salvinia + o o
Leopardus wiedii yucatanicus + o o
Leopardus wiedii cooperi + o o
L. w. wiedii
L. w. vigens
L. w. glauculus
A new taxonomy of the Felidae
51
pampas cat
Leopardus colocola
E: Pampas cat; F: Chat des pampas; G: Pampaskatze; Sp: Gato pajero,
gato de los pajonales, gato de pajonal, osio.
The taxonomic history of pampas cats is quite complex, owing to previ-
ous misidentifications. For example, there has been confusion with the
Andean mountain cat, L. jacobita, and Hamilton Smith (1827; 479-480)
gave the name L. colocolo to a cat from Guyana (García-Perea 1994),
which cannot now be identified.
In 1782, Molina (p. 295) twice mentioned the cat “Il Colocolo” but he
also twice used the specific name Felis colocola (on p. 295 and in the
list of new species described in Saggio sulla storia naturali del Chili on
p. 341). We suspect that colocola was the name that he intended for
the species and that it was not a spelling or typing error by Molina or
the printer. In his later publication (1788: 322), Molina maintained the
spelling colocola, but by the time of the English translation of this work
in 1808, the name appeared as Felis colocolo in the text (pp. 206-207),
but remained as colocola in the list of new species (p. 239). In Edition
2 of the Saggio (1810: 245) the specific name appears as colocolo.
It seems that Molina’s name colocola has been “corrected” to colocolo
by subsequent authors. As noted above, there is nothing in the original
1782 work to indicate that colocola was a mistake and therefore that
colocolo can be accepted as a “justified emendation”. Article 33.2.2 of
the Code notes “The correction of an incorrect original spelling ... is a
“justified emendation”, and the name thus corrected retains the au-
thorship and date of the original spelling”. This Article is followed by
Article 33.2.3: “Any other emendation is an “unjustified emendation”;
the name thus emended is available and it has its own author and
date and is a junior objective synonym of the name in its original spell-
ing; it enters into homonymy and can be used as a substitute name,
but Article 33.2.3.1 states that when an unjustified emendation is in
prevailing usage and is attributed to the original author and date it is
deemed to be a justified emendation”. Therefore colocolo could be
maintained as being the correct spelling with the original authorship
and date (Molina 1782). However, this is dependent on interpretation
of “prevailing usage”. A quick search on Google Scholar reveals at
least 19 references which use the name colocola dating from 1782
to 2013, although interestingly these were mostly published in South
American countries. Members of the CCTF have agreed to revert to
Molina’s original spelling of colocola for this species.
Until the mid-1990s there was a consensus that the pampas cat com-
prised a single species, but García-Perea (1994) carried out a compre-
hensive review of skull and pelage characters on 86 specimens from
throughout the species’ range. This review concluded that the pampas
cat actually comprises three species and 10 subspecies as follows:
Leopardus colocolo colocolo (Molina, 1782; 295).
Locality from where the species was first described: “Chili” [= Chile];
restricted to “province of Valparaiso”, Chile by Osgood (1943; 79).
Type: None designated.
Distribution: C Chile from Coquimbo to Concepción.
Leopardus colocolo wolffsohni (García-Perea, 1994; 30).
Type locality: Río Camarones, provinicia Tarapacá, between 2000 and
4000 m, Chile.
Holotype: USNM 391853 skin and skull.
Distribution: Tarapacá Province, N Chile on W slopes of Andes.
Leopardus pajeros pajeros (Desmarest, 1816; 114).
Locality from where the species was first described: “las pampas de
Buenos Ayres, entre los 35 y 36 grados” (Azara 1802; 160), Argentina.
Type: None designated. Based on account by Azara (1802; 160) of the
“gato pajero”.
Distribution: La Pampa Province, C Argengtina.
Distinguishing characters: Larger and yellower than pardalis.
Leopardus pajeros budini (Pocock, 1941; 263).
Type locality: Mount Sola, 2500 m, in Salta, northern Argentine [= Ar-
gentina].
Holotype: BMNH 1934.11.4.5 adult male skin and skull.
Distribution: Mountains of NW Argentina on E side of Andes.
Leopardus pajeros crespoi (Cabrera, 1957; 71).
Type locality: Aguaray, provincia de Salta, [Argentina].
Holotype: MACN 36.230 female skin and skull.
Distribution: Known only from type locality.
Leopardus pajeros crucinus (Thomas, 1901; 247).
Type locality: Santa Cruz, Argentina.
Holotype: BMNH 1855.12.24.261 skin and skull.
Distribution: S half of Argentina and Chilean Patagonia.
Leopardus pajeros garleppi (Matschie, 1912; 259).
Type locality: von Cuzco in Südost-Peru, im Gebiet des Apurimac, der
durch den Ucayali zum oberen Amazonas abwässert [from Cuzco in SE
Peru, in the region of Apurimac, {Peru}, from the Ucuyali to the Upper
Amazon].
Holotype: ZMB Mam 21244 adult male skin and skull.
Distribution: Highland steppes of E side of Peruvian Andes.
Leopardus pajeros steinbachi (Pocock, 1941; 264).
Type locality: Tiraque, Cochabamba, western Bolivia, 4000 m.
Holotype: BMNH 1934.9.2.31 adult female skin and skull.
Distribution: Highland steppes of E slopes of Bolivian Andes.
Leopardus pajeros thomasi (Lönnberg, 1913; 7).
Type locality: Near Quito, [Ecuador].
Holotype: NRM A621386 male skin and skull.
Distribution: Highland steppes of E side of Peruvian Andes.
Pampas cat, northern Argentina (Photo A. Seguin).
CATnews Special Issue 11 Winter 2017
52
pampas cat
Leopardus braccatus braccatus (Cope, 1889; 144).
Type locality: The province of Rio Grande do Sul, or in Matto Grosso;
restricted to “Chapada dos Guimaraes, Matto Grosso”, Brazil by Allen
(1919; 378).
Holotype: AMNH MO-354 adult male skin and skull.
Distribution: SW Brazil and Paraguay.
Leopardus braccatus munoai (Ximénez, 1961; 3).
Type locality: Arroyo Perdido, Departmento de Soriano, [Uruguay].
Holotype: MNHNM 884 female skin and skull.
Distribution: S Brazil, Uruguay.
Discussion
Undoubtedly, there is considerable morphological variation in this tax-
on, but so far genetic studies do not appear to support conclusively the
differentiation of more than one species. Molecular data do indicate
the existence of genetic differentiation (i.e. phylogeographical struc-
ture) among present-day populations, but these partitions are rather
recent, at the same level as intra-specific partitions observed in other
felids. Therefore, based on evolutionary depth (i.e. time of divergence),
there is no support for species-level partitions. However, it is still pos-
sible that the group comprises a complex of very recently diverged spe-
cies, whose current genetic connectivity and ecological differentiation
is still not conclusively settled. Here is a summary of recent studies:
Johnson et al. (1999) examined the phylogeography of a limited sample
of pampas cats using mtDNA. Genetic divergence among geographical
groups was significant but shallow in terms of evolutionary time, and
thus they considered that it did not support the recognition of three
species, although three subspecies could be supported:
Leopardus colocolo colocolo incl. pajeros, budini, crespoi.
Leopardus colocolo garleppi incl. thomasi.
Leopardus colocolo braccatus incl. munoai.
Nascimento (2010) recognised six species based on studies of skins
and skulls from throughout the range:
Leopardus colocolo
Leopardus pajeros
Leopardus braccatus
Leopardus garleppi
Leopardus budini
Leopardus munoai
Ruiz-García et al. (2013) investigated microsatellite and mtDNA diver-
sity in the largest sample of pampas cats to be investigated so far, but
even so, some putative subspecies were not sampled. Their results
support those of Cossíos et al. (2009) and can be summarised briefly
as follows:
The pampas cat is a single species comprised of several subspecies,
including:
Leopardus colocolo colocolo
Leopardus colocolo pajeros incl. crucinus
Leopardus colocolo budini
Leopardus colocolo garleppi incl. wolffsohni
Leopardus colocolo braccatus
?Leopardus colocolo steinbachi
?Leopardus colocolo thomasi
There is significant morphological and genetic variation in L. colo-
cola sensu lato, but it is difficult to evaluate what this variation
means taxonomically. While molecular studies suggest that there is
only one species of pampas cat, there is a fundamental difference
in rhinarium structure between some forms, which might indicate a
species difference between western Chilean populations (a small
rhinarium) compared with that of others (large ovate rhinarium).
Perhaps introgression with other Leopardus species has influenced
rhinarium morphology in some pampas cats, although currently there
is no evidence of introgression of L. tigrinus or any other species into
L. colocola.
C. Groves (pers. comm.) has suggested the following provisional clas-
sification:
Leopardus garleppi: While this has a colour pattern not all that differ-
ent from L. wolffsohni, it has the skull features typical for colocola, and
is mtDNA clade A, which is different from all the others whose DNA
is known. The smaller size of thomasi, mentioned by García-Perea
(1994), is not all that cogent as in her Table 1 the skull sizes overlap.
Leopardus steinbachi: This seems to differ, but not greatly, in colour
pattern from garleppi, but has mtDNA clade C, shared with some
northern populations of budini.
Provisional species: Leopardus budini: Colour pattern is distinctive, its
mtDNA is polymorphic, but mostly clade C; this taxon may actually be
of hybrid origin, or indeed, that which García-Perea (1994) thought was
a single taxon might actually be a mixture of two or more, or indeed
that at least the northern end of it could be a hybrid swarm? There is a
possibility that crespoi could be a synonym.
Provisional species: Leopardus pajeros: Again, there is a possibility
that this taxon could actually be, at least in part, a hybrid swarm or a
mixture of two or more: García-Perea (1994) says that it shows colour
pattern types 2B and 2C, but at the same time has its own colour tone;
it has predominantly mtDNA clade D, but with some individuals falling
into clade C and some in clade B.
Leopardus crucinus: This one really is different from any other in colour
pattern and apparently in skull. It has mtDNA clade D.
Leopardus wolffsohni: Distinctive in colour pattern and skull features,
and mtDNA clade C, which also occurs as a minority in neighbouring
budini.
Leopardus colocola: Distinctive in all sorts of features. DNA unknown.
Leopardus braccatus: Classed correctly as a distinct species by García-
Perea (1994).
Leopardus munoai: Colour pattern type is different from braccatus, but
otherwise not said to be very different.
Distribution of tentative subspecies of pampas cat. Borders be-
tween subspecies are speculative.
L. c. garleppi
L. c. wolfsohni
L. c. colocola
L. c. pajeros
L. c. munoai
L. c. braccatus
L. c. budini
A new taxonomy of the Felidae
53
Overall the taxonomy of Leopardus colocola remains confused, although
it is commonly viewed by molecular biologists as comprising a single
species, but the date of the main split between the two main mitochodri-
al clades of northern and southern populations is more than 800kya (Cos-
síos et al. 2009), which would support the recognition of at least two
species based on our species definition above. However, there is a lack
of congruity between morphological and genetic studies, so that further
data and analyses are required to elucidate this confused picture.
We propose the following subspecies based mainly on Cossios et al.
(2009) and Ruiz-Garcia et al. (2013), but recognise that this is likely to
change as more information becomes available, including the possible
recognition of some of these as full species:
Leopardus colocola colocola (Molina, 1782).
Distribution: Central Chile W of Andes.
Leopardus colocola wolffsohni (García-Perea, 1994).
Distribution: Tarapacá province, N Chile W of Andes.
Leopardus colocola pajeros (Desmarest, 1816) including crucinus.
Distribution: C, NC and S Argentina.
Leopardus colocola budini (Pocock, 1941), including steinbachi.
Distribution: NW Argentina and Bolivia E of Andes.
Leopardus colocola garleppi (Matschie, 1912), including thomasi.
Distribution: S Colombia, Ecuador, Peru E of Andes.
Leopardus colocola braccatus (Cope, 1889).
Distribution: SW and C Brazil, Paraguay.
Leopardus colocola munoai (Ximénez, 1961)
Distribution: Uruguay.
References
Allen J. A. 1919. Notes on the synonymy and nomenclature of the smaller
spotted cats of tropical America. Bulletin of the American Museum of
Natural History 41, 341-419.
Azara de F. 1802. Apuntamientos para la Historia Natural de los quadrúpe-
dos del paraguay y Rio de la Plata, vol 1. Imprenta de la Viuda de Ibarra,
Madrid.
Cabrera A. 1957. Dos felidos argentinos inéditos (Mammalia, Carnivora). Ne-
otrópica 3, 70-72.
Cope E. D. 1889. On the mammalia obtained by the naturalist exploring expedi-
tion to southern Brazil. American Naturalist 23, 128-150.
Cossíos D., Lucherini M., Ruiz-García M. & Angers B. 2009. Influence of an-
cient glacial periods on the Andean fauna: the case of the pampas cat
(Leopardus colocolo). BMC Evolutionary Biology 9: 68.
Desmarest A. G. 1816. Chat, Felis, Linn., Briss., Screb. In Nouveau diction-
naire l’histoire naturelle, appliquée aux arts, à l’agriculture, à l’économie
rurale et domestique, à la médecine, etc. Par une société de naturalistes et
d’agriculteurs (nouvelle editon), 6. Deterville, Paris, pp. 73-123.
Garcia-Perea R. 1994. The pampas cat group (Genus Lynchailurus Severtzov,
1858) (Carnivora: Felidae), a systematic and biogeographic review. Ameri-
can Museum Novitates No. 3096, 1-35.
Gray J. E. 1842. Descriptions of some new genera and fifty unrecorded species
of Mammalia. Annals and Magazine of Natural History 10, 255-267.
Hamilton Smith C. 1827. [The colocolo]. In Griffith E. (ed). The Animal Kingdom
arranged in conformity with its organization, by the Baron Cuvier, Member of
the Institute of France etc., etc. etc., with additional descriptions of all the
species hitherto named, and many not before noticed. Volume 2, pp. 479-
480. Whittaker, London.
Johnson W. E., Pecon Slattery J., Eizirik E., Kim J.-H., Menotti Raymond M., Bo-
nacic C., Cambre R., Crawshaw P., Nunes A., Seuanez H. N., Martins Moreira
A., Seymour K. L., Simon F., Swanson W. & O’Brien S. J. 1999. Disparate phy-
logeographic patterns of molecular genetic variation in four closely related
South American small cat species. Molecular Ecology 8, 79-94.
Lönnberg E. 1913. Mammals from Ecuador and related forms. Arkiv för Zoologi
8, 1-36.
Matschie P. 1912. Über Felis jacobita, colocola, und zwei ihnen ähnliche
Katzen. Sitzungsberichte der Gesellschaft Naturforschender Freunde zu
Berlin 4, 255-259.
Molina G. I. 1782. Saggio sulla storia naturale del Chilli. Stamperia di S. Tom-
maso d’Aquino, Bologna.
Molina G.I. 1810. Saggio sulla storia naturali del Chili, 2nd edition. de Fratelli
Masi & Comp, Bologna.
Subspecies Morphology Molecular Biogeography Certainty Comments
Leopardus colocola colocola ++ ++ ++
Leopardus colocola braccatus ++ ++ ++
Leopardus colocola budini + + + Possibly includes steinbachi
Leopardus colocola crespoi + + +
Leopardus colocola crucinus + - + Probable synonym of pajeros
Leopardus colocola garleppi + + + Includes thomasi
Leopardus colocola munoai + ++ + Probably distinct from braccatus
Leopardus colocola pajeros ++ ++ ++ Includes crucinus
Leopardus colocola steinbachi + + + Possible synonym of budini
Leopardus colocola thomasi + o + Probable synonym of garleppi
Leopardus colocola wolfsohni + + ++ Status uncertain
pampas cat
CATnews Special Issue 11 Winter 2017
54
southern tigrina
Leopardus guttulus
E: Southern tigrina; F: Oncille du sud, chat-tigre du sud; G: Südliche
Tigerkatze; Sp: Sp: Tigrillo, tirica, gato tigre, caucel.
Leopardus guttulus (Hensel, 1872; 73).
Type locality: Rio Grande do Sul, Brasilien [= Brazil].
Syntypes: ZMB Mam 21229 and Mam 21231 skulls only.
Distribution: SE and S Brazil, N Argentina and probably Paraguay.
Discussion
Although described originally as a new species, Felis guttula (Hensel
1872), this species was commonly treated as a subspecies of Leopar-
dus tigrinus until 2013 (Wozencraft 2005, Trigo et al. 2013). However,
despite this apparent consensus, others had suggested that Leopardus
tigrinus may well comprise two or even three distinct species (e.g.
Leyhausen 1963). In the fur trade, two kinds of tigrina fur were recog-
nised, the typical kind and the Ceara kind.
Trigo et al. (2013) demonstrated with a comprehensive molecular study
that Leopardus tigrinus and L. guttulus are clearly distinct species. L.
guttulus hybridises with L. geoffroyi in Rio Grande do Sul, which is
the type locality. The syntypes of L. guttulus should be investigated in
order to check whether they are hybrids or not.
Further research is required to establish whether northwestern tigri-
nas comprise a third species, L. pardinoides, or even a fourth species,
L. oncilla, in Costa Rica (Johnson et al. 1999, Trigo et al. 2008, Nas-
cimento 2010, Li et al. 2016). Also the distribution and morphology
of L. guttulus need to be better defined. Until then L. guttulus is re-
cognised as a monotypic species.
Leopardus guttulus (Hensel, 1872; 73).
Distribution: SE and S Brazil, N Argentina and probably Paraguay.
References
Hensel R. 1872. Beiträge zur Kenntniss der Säugethiere Süd-Brasiliens. Physi-
kalische Abhandlungen der Königlichen Akademie der Wissenschaften zu
Berlin, 1-130.
Johnson W. E., Pecon Slattery J., Eizirik E., Kim J.-H., Menotti Raymond M.,
Bonacic C., Cambre R., Crawshaw P., Nunes A., Seuanez H. N., Martins
Moreira A., Seymour K. L., Simon F., Swanson W. & O’Brien S. J. 1999.
Disparate phylogeographic patterns of molecular genetic variation in four
closely related South American small cat species. Molecular Ecology 8,
79-94.
Leyhausen P. 1963. Über südamerikanische Pardelkatzen. Zeitschrift für Tier-
psychologie 20, 627-640.
Li G., Davis B. W., Eizirik E. & Murphy W. J. 2016. Phylogenomic evidence
for ancient hybridization in the genomes of living cats (Felidae). Genome
Research 26, 11-11.
Nascimento do F. O. 2010. Revisão taxonômica gênero do Leopardus Gray,
1842 (Carnivora, Felidae). PhD thesis University of Sao Paulo, Brazil.
Trigo T. C., Freitas T. R. O., Kunzler G., Cardoso L., Silva J. C. R., Johnson W.
E., O’Brien S. J., Bonatto S. L. & Eizirik E. 2008. Inter-species hybridization
among Neotropical cats of the genus Leopardus, and evidence for an intro-
gressive hybrid zone between L. geoffroyi and L. tigrinus in southern Brazil.
Molecular Ecology 17, 4317-4333.
Trigo T. C., Schneider A., Oliveira de T. G., Lehugeur L. M., Silveira L., Freitas
T. R. O. & Eizirik E. 2013. Molecular data reveal complex hybridization and
a cryptic species of Neotropical wild cat. Current Biology 23, 2528-2533.
Wozencraft W. C. 2005. Order Carnivora. In Mammal species of the world. A
taxonomic and geographic reference (3rd ed.). Wilson D. E. & Reeder D. M.
(Eds). Johns Hopkins University Press, Baltimore, pp. 532-628.
Species Morphology Molecular Biogeography Certainty Comments
Leopardus guttulus ++ ++ + Distinct species from L. tigrinus
Molina J. I. 1788. Compendio de la historia geografica, natural y civil del reyno
de Chile, part 1. Don Antonio de Sancha, Madrid.
Molina J.I. 1808. The geographical, natural and civil history of Chili, vol. 1. I.
Riley, Middletown, Connecticut.
Nascimento do F. O. 2010. Revisão taxonômica gênero do Leopardus Gray,
1842 (Carnivora, Felidae). PhD thesis University of Sao Paulo, Brazil.
Osgood W. H. 1943. The mammals of Chile. Zoological Series, Publications of
the Field Museum of Natural History 30, 268 pp.
Pocock R. I. 1941. The examples of the colocolo and of the pampas cat in the
British Museum. Annals and Magazine of Natural History (11th series) 7,
257-274.
Ruiz-García M., Cossíos D., Lucherini M., Yáñez J., Pinedo-Castro M. & Angers
B. 2013. Population genetics and spatial structure in two Andean cats (the
pampas cat, Leopardus pajeros, and the Andean mountain cat, L. jacobita)
by means of nuclear and mitochondrial markers and some notes on bio-
metrical markers. In Molecular population genetics, evolutionary biology
and biological conservation on Neotropical carnivores. Ruiz-García M. &
Shostell J. M. (Eds). Nova, New York , pp.187-244.
Thomas O. 1901. New Neotropical mammals, with a note on the species
of Reithrodon. Annals and Magazine of Natural History (7th series) 8,
246-255.
Wozencraft W. C. 2005. Order Carnivora. In Mammal species of the world. A
taxonomic and geographic reference (3rd ed.). Wilson D. E. & Reeder D. M.
(Eds). Johns Hopkins University Press, Baltimore, pp. 532-628.
Ximénez A. 1961. Nuevo subspecie del gato pajero en el Uruguay. Comunica-
ciones Zoologicas del Museo de Historia Natural de Montevideo 5, 1-5.
© T. G. de Oliveira
A new taxonomy of the Felidae
55
northern tigrina
Leopardus tigrinus
E: Northern tigrina, oncilla; F: Oncille du nord, chat-tigre du nord; G:
Nordliche Ozelotkatze, Nördliche Tigerkatze; Sp: Tigrillo, tirica, gato
tigre, caucel.
Wozencraft (2005) recognised a single species with four subspecies:
Leopardus tigrinus tigrinus (Schreber, 1775; pl.106, 1777; 396).
Type locality: südlichen Amerika, restricted to Cayenne by Allen (1919).
Holotype: Based on le margay in Buffon (1765; 248).
Distribution: NE Brazil and French Guiana to E Venezuela.
Leopardus tigrinus guttulus (Hensel, 1872; 73).
Type locality: Rio Grande do Sul, Brasilien [= Brazil].
Syntypes: ZMB Mam 21229 and Mam 21231 skulls only.
Distribution: SE and S Brazil, N Argentina and probably Paraguay.
Leopardus tigrinus pardinoides (Gray, 1867; 400).
Type locality: India, revised to Bogotá, Colombia by Gray (1874).
Holotype: BMNH 1855.12.24.185 skin and skull.
Distribution: W Venezuela, Colombia and Ecuador.
Leopardus tigrinus oncilla (Thomas, 1903; 237).
Type locality: Volcan de Irazu, Costa Rica.
Holotype: BMNH 1878.7.6.3 skin, skull lost.
Distribution: Costa Rica and ?Panama.
Discussion
The taxonomy of South American small spotted cats was highly unsta-
ble during the 19th and early 20th centuries, with particular confusion
between L. wiedii and L. tigrinus. This issue has mostly been settled,
i.e. little confusion remains regarding the distinctiveness between
these two species. However, there has been an emerging realisation
that L. tigrinus may in fact represent a species complex, whose exact
composition is still not completely clear. This concept is a paradigm
shift relative to the modern consensus supporting a single species of L.
tigrinus, in spite of earlier suggestions that this species actually com-
prised two or three species (Allen 1919, Leyhausen 1963).
Johnson et al. (1999) examined variation in mtDNA (16S rDNA, ATP8
and ND5) in four L. t. oncilla from Costa Rica and 28 L. (t.) guttulus
from SE Brazil. There was a 4.9% mtDNA sequence divergence be-
tween animals from these different populations, suggesting species-
level distinction. The same pattern was again observed by Trigo et al.
(2008), also with mtDNA data. The limited sample size and the use of
only a matrilineal marker (mtDNA) in those studies, precluded a more
confident conclusion that L. t. oncilla was indeed a distinct species. A
recent study (Li et al. 2016), based on genome-wide nuclear markers,
supported the distinctiveness of the single individual from Costa Rica.
This provides strong support for the recognition of L. oncilla as a sepa-
rate species, although additional analyses, including more individuals,
should be performed to further confirm this conclusion and establish
its relationship with neighbouring populations in NW South America.
Based on a morphological analysis of skulls and pelages, Nascimento
(2010) proposed that this species actually comprises four species, L.
tigrinus, L. guttulus, L. pardinoides and L. oncilla.
Trigo et al. (2013) demonstrated with a comprehensive molecular study
that tigrinas occurring in Brazil comprise two species, Leopardus tigri-
nus and L. guttulus, which may not even be sister species. L. guttulus
hybridises with L. geoffroyi in Rio Grande do Sul, which is the type
locality.
Further research is required to establish whether northwestern South
American tigrinas comprise yet another species, L. pardinoides. Li et
al. (2016) suggest that tigrinas from Central America may represent a
distinct species, L. oncilla, but it is unclear whether this would be part
of pardinoides. Also the distribution and morphology of L. guttulus and
L. tigrinus need to be better defined. Tigrinas appear to be absent from
the core of the Amazon, which would pose a significant biogeographi-
cal barrier between these known and potential species. Until then L.
tigrinus is recognised as having two subspecies:
Leopardus tigrinus tigrinus (Schreber, 1775), including pardinoides.
Distribution: N South America possibly as far S as Bolivia and N Ar-
gentina.
Leopardus tigrinus oncilla (Thomas, 1903).
Distribution: Costa Rica and possibly Panama.
References
Buffon Comte de G.-L. L. 1765. Le margay. In Histoire Naturelle, générale
et particuliére avec la description du Cabinet du Roi, Volume 13. Buffon
Comte de G.-L. L. & Daubenton L. J. M. (Eds). De l’Imprimière royale, Paris,
pp. 248-253, pl. 37.
Gray J. E. 1867. Notes on certain species of cats in the collection of the British
Museum. Proceedings of the Zoological Society of London 35, 394-405.
L. t. oncilla
L. t. tigrinus
© Projeto Gatos do Mato - Brazil
CATnews Special Issue 11 Winter 2017
56
Geoffroy’s cat
Gray J. E. 1874. The Bogotá cat (Felis pardinoides, Gray). Annals and Magazine
of Natural History (4th series) 13, 475.
Hensel R. 1872. Beiträge zur Kenntniss der Säugethiere Süd-Brasiliens. Physi-
kalische Abhandlungen der Königlichen Akademie der Wissenschaften zu
Berlin, 1-130.
Johnson W. E., Pecon Slattery J., Eizirik E., Kim J.-H., Menotti Raymond M.,
Bonacic C., Cambre R., Crawshaw P., Nunes A., Seuanez H. N., Martins
Moreira A., Seymour K. L., Simon F., Swanson W. & O’Brien S. J. 1999.
Disparate phylogeographic patterns of molecular genetic variation in four
closely related South American small cat species. Molecular Ecology 8,
79-94.
Leyhausen P. 1963. Über südamerikanische Pardelkatzen. Zeitschrift für Tier-
psychologie 20, 627-640.
Li G., Davis B. W., Eizirik E. & Murphy W. J. 2016. Phylogenomic evidence
for ancient hybridization in the genomes of living cats (Felidae). Genome
Research 26, 11-11.
Nascimento do F. O. 2010. Revisão taxonômica gênero do Leopardus Gray,
1842 (Carnivora, Felidae). PhD thesis University of Sao Paulo, Brazil.
Schreber J. C. D. 1775. Die Säugethiere in Abbildungen nach der Natur mit
Beschreibungen, vol.2 (15). Wolfgang Walther, Erlangen.
Schreber J. C. D. 1777. Die Säugethiere in Abbildungen nach der Natur mit
Beschreibungen, vol. 3: Wolfgang Walther, Erlangen, pp. 396-397.
Thomas O. 1903. Notes on Neotropical mammals of the genera Felis, Hapale,
Oryzomys, Akodon and Ctenomys, with descriptions of new species. An-
nals and Magazine of Natural History (7th series) 12, 234-243.
Trigo T. C., Schneider A., Oliveira de T. G., Lehugeur L. M., Silveira L., Freitas
T. R. O. & Eizirik E. 2013. Molecular data reveal complex hybridization and
a cryptic species of Neotropical wild cat. Current Biology 23, 2528-2533.
Wozencraft W. C. 2005. Order Carnivora. In Mammal species of the world. A
taxonomic and geographic reference (3rd ed.). Wilson D. E. & Reeder D. M.
(Eds). Johns Hopkins University Press, Baltimore, pp. 532-628.
Subspecies Morphology Molecular Biogeography Certainty Comments
Leopardus tigrinus tigrinus ++ ++ + Distinct from L. guttulus
Leopardus tigrinus pardinoides + o + Possibly distinct or even a distinct species
Leopardus tigrinus guttulus ++ ++ + Distinct species from L. tigrinus
Leopardus tigrinus oncilla + ++ ++ Needs further assessment; may be a distinct
species.
Leopardus geoffroyi
E: Geoffroy’s cat; F: Chat de Geoffroy; G: Geoffroy Katze; Klein-
flechkatze, Salzkatze; Sp: Gato de mato, gato montés.
Wozencraft (2005) recognised four subspecies of L. geoffroyi based on
Ximenez (1975):
Leopardus geoffroyi geoffroyi (d’Orbigny and Gervais, 1844; 40).
Type locality: des rives du Rio Negro, en Patagonie [the banks of the
Rio Negro, Patagonia].
Syntypes: MNHN-ZM-MO-2001-298 mounted skin (skull inside),
MNHN 2001-299 mounted skin (skull inside) and MNHN 2001-300
mounted skin (skull inside).
Distribution: C to S Argentina and Chile.
Leopardus geoffroyi salinarum (Thomas, 1903; 239).
Type locality: Cruz del Eje, Cordova, Argentina, altitude 600m.
Holotype: BMNH 1902.2.5.10 female skin and skull.
Distribution: NW Argentina.
Leopardus geoffroyi paraguae (Pocock, 1940; 351).
Type locality: Paraguay.
Holotype: BMNH 1871.3.3.6 adult female skin and skull.
Distribution: Paraguay, Uruguay, NE Argentina and SE Brazil.
Leopardus geoffroyi euxanthus (Pocock, 1940; 352).
Type locality: Tiragui, Cochabamba district in the western Bolivian
highlands, 3,300 m.
Holotype: BMNH 1934.9.2.27 adult male skin and skull.
Distribution: Bolivia and far N of Argentina.
Discussion
Johnson et al. (1999) examined the phylogeography of L. geoffroyi
based on 38 specimens from throughout its range. They examined
variation in mtDNA (16S rDNA, ATP-8 and ND5) and among 20 micro-
satellite loci. No phylogeographical structure was present, so that no
subspecies could be supported, despite some degree of morphological
differentiation.
A recent morphological study of skull morphometrics by Rojo Gómez
& García-Perea (in review) showed that while L. g. salinarum has a
© J. Pereira
A new taxonomy of the Felidae
57
guiña
small distinct skull, there is considerable overlap between L. g. geof-
froyi and L. g. paraguae, suggesting that the latter are consubspecific.
However, this analysis did not remove the effect of size and hence it
is not possible to be certain whether the skull shape of L. g. salinarum
is significantly different from that of L. g. geoffroyi or represents a
scaled-down version as part of clinal variation.
Nascimento (2014) examined geographical variation in skull morpho-
metrics and pelage coloration and markings from all putative sub-
species. He found that there was no evidence for distinct groups,
but only clinal variation with latitude; diagnostic characters for one
putative subspecies occurred in individuals of another putative sub-
species.
Until further genetic data are available, it is probably best to treat
L. geoffroyi as a monotypic species, which displays a morphological
cline in relation to differences in habitat throughout its range.
Leopardus geoffroyi (d’Orbigny and Gervais, 1844).
Distribution: S South America from Bolivia to Patagonia.
References
d’Orbigny A. & Gervais P. 1844. Mammalogie: Nouvelle espèce de Felis. So-
ciètè Philomatique de Paris, 40-41.
Johnson W. E., Pecon Slattery J., Eizirik E., Kim J.-H., Menotti Raymond M.,
Bonacic C., Cambre R., Crawshaw P., Nunes A., Seuanez H. N., Martins
Moreira A., Seymour K. L., Simon F., Swanson W. & O’Brien S. J. 1999.
Disparate phylogeographic patterns of molecular genetic variation in four
closely related South American small cat species. Molecular Ecology 8,
79-94.
Nascimento do F. O. 2014. On the morphological variation and taxonomy of the
Geoffroy’s cat Leopardus geoffroyi (d’Orbigny & Gervais 1844) (Carnivora,
Felidae). Papéis Avulsos de Zoologia, Museu de Zoologia da Universidade
de São Paulo 54, 129-160.
Pocock R. I. 1940. The races of Geoffroy’s cat (Oncifelis geoffroyi). Annals and
Magazine of Natural History (11th series) 6, 350-355.
Thomas O. 1903. Notes on Neotropical mammals of the genera Felis, Hapale,
Oryzomys, Akodon and Ctenomys, with descriptions of new species. An-
nals and Magazine of Natural History (7th series) 7, 234-243.
Wozencraft W. C. 2005. Order Carnivora. In Mammal species of the world. A
taxonomic and geographic reference (3rd ed.). Wilson D. E. & Reeder D. M.
(Eds). Johns Hopkins University Press, Baltimore, pp. 532-628.
Ximenez A. 1975. Felis geoffroyi. Mammalian Species No. 54, 1-4.
Subspecies Morphology Molecular Biogeography Certainty Comments
Leopardus geoffroyi geoffroyi ++ ++ ++
Leopardus geoffroyi salinarum - - -
Leopardus geoffroyi paraguae - - -
Leopardus geoffroyi euxanthus - - -
Leopardus guigna
E: Guiña, kod-kod; F: Guigna, chat du Chili, kodkod; G: Kodkod, Chil-
enische Waldkzatze, Nachtkatze; Sp: Guiña, huiña.
Traditionally two subspecies of Leopardus guigna are recognised
(Wozencraft 2005):
Leopardus guigna guigna (Molina, 1782; 295).
Locality from where the species was first described: Chili; restricted
by Thomas (1903; 240) to Valdivia in Chile based on Philippi (1873).
Type: None designated.
Distribution: S Chile (38° to 48° S) and SW Argentina (39° to 46° S).
Distinguishing characters: Darker and smaller than tigrillo; often mela-
nistic.
Leopardus guigna tigrillo (Schinz, 1844; 470).
Locality from where the subspecies was first described: Chili; restrict-
ed to Talcahuano, Prov. Concepcion, Chile based on Pöppig (1829; 7),
following Cabrera (1958), who pointed out that Osgood’s (1943; 85)
molinae is a junior synonym of tigrillo.
Type: None designated.
Distribution: N and C Chile (30° to 38° S).
Distinguishing characters: Larger and paler than guigna.
Discussion
A molecular study by Napolitano et al. (2014) of mtDNA and micros-
atellites identified a phylogeographical pattern that showed moderate
separation between northern and southern populations, thereby sup-
porting the two recognised subspecies.
Therefore, we recognise two subspecies:
© A. Sliwa
CATnews Special Issue 11 Winter 2017
58
serval
Leopardus guigna guigna (Molina, 1782).
Distribution: S Chile (38° to 48° S) and SW Argentina (39° to 46° S).
Leopardus guigna tigrillo (Schinz, 1844).
Distribution: N and C Chile (30° to 38° S).
References
Molina G. I. 1782. Saggio sulla storia naturale del Chilli. Bologna: Stamperia
di S. Tommaso d’Aquino.
Napolitano C., Johnson W. E., Sanderson J., O’Brien S. J., Hoelzel A. R., Freer
R., Dunstone N., Ritland K., Ritland C. E. & Poulin E. 2014. Phylogeography
and population history of Leopardus guigna, the smallest American felid.
Conservation Genetics 15, 631-653.