Mammal Rev. 2010, Volume 40, No. 1, 90–102. Printed in Singapore.

SHORT COMMUNICATION
What is Panthera palaeosinensis? mam_151 90..102

Ji H. MAZÁK* Shanghai Science and Technology Museum, No. 2000 Century

Avenue, Pudong New Area, Shanghai 200127, China. E-mail: jimazak@gmail.com,
jimazak@citiz.net

ABSTRACT
The pantherine cat Panthera palaeosinensis from putative Plio-Pleistocene deposits
in North China is one of the oldest known species of Panthera, but its relationship to
other Pantherinae is still debated. I compare the holotype skull of P. palaeosinensis
with 508 skulls of extant and extinct pantherine cats and show that, when control-
ling for size, the skull morphology of P. palaeosinensis is most similar to that of the
lion Panthera leo or leopard Panthera pardus. Results support the hypothesis that P.
palaeosinensis represents a form closely related either to the early lion or leopard
clade or to the ancestor of the genus Panthera and suggest an Asian origin for
Panthera.

Keywords: Asia, multivariate analysis, Panthera origin, Panthera palaeosinensis, skull

morphology
Mammal Review (2010), 40, 90–102
doi: 10.1111/j.1365-2907.2009.00151.x

INTRODUCTION
The subfamily Pantherinae is a group of large-bodied felids, comprising six living
species (clouded leopard Neofelis nebulosa, snow leopard Uncia uncia, tiger Panthera
tigris, jaguar Panther onca, lion Panthera leo and leopard Panthera pardus) and
several extinct forms (including the Eurasian jaguar Panthera gombaszoegensis, the
European cave lion Panthera leo spelaea and the American lion Panthera leo atrox).
Molecular phylogenies suggest that the origin and radiation of the Pantherinae
can be traced back to 5–10 Ma and probably took place in Asia (Collier & O’Brien
1985, Wayne et al. 1989, Johnson & O’Brien 1997, Johnson et al. 2006), but this
scenario lacks support from the fossil record. Our knowledge of the origin and
evolution of the Pantherinae is still incomplete. The oldest precisely dated Panthera
fossil is from East Africa, is estimated to date from at least 3.46 Ma, and is attributed
to a lion or lion-like Panthera or leopard (Barry 1987, Turner 1990, Turner & Antón
1997, 2004, Werdelin & Lewis 2005). Complete pantherine skulls of Pre-Pleistocene
age are extremely rare in the fossil record.
The Plio-Pleistocene Chinese pantherine cat Felis palaeosinensis (= Panthera palae-
osinensis; Zdansky 1924) is one of the oldest known fossils attributed to Panthera,
but its taxonomic affinities have long been debated. It has been interpreted as a
small primitive tiger (Hemmer 1967, 1987, Hemmer et al. 2001), or a species close to
the ancestor of the tiger (Christiansen 2008b), a primitive or large form of leopard

*Correspondence author.

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 The Panthera palaeosinensis skull 91

(Pei 1934, Teilhard de Chardin & Leroy 1945, Kurtén 1968), or an ancestor of two or
more of today’s Panthera species (Kitchener 1999). The holotype, the only known
almost complete skull, is intermediate in size between large Indian leopards and
Sunda Island tigers (Hemmer 1967, Kitchener 1999). Since cats may vary greatly in
size, owing to complex ecological factors (Kitchener 1991), it is misleading to deter-
mine the affinity of specimens relying primarily on size. Most previous researchers
have focused on size, and on dental or non-metric craniodental characteristics
(Zdansky 1924, Hemmer 1967, Christiansen 2008b), and have not assessed the overall
skull morphology by using morphometric techniques. Thus, no consensus has
emerged about what the skull morphology can tell us about the affinities of P.
palaeosinensis.
Here, I address two specific questions concerning the skull of P. palaeosinensis: (i)
when controlling for size and in comparison with a large comparative sample of
extant and extinct Pantherinae skulls, does P. palaeosinensis fall within the range
of skull morphologies of modern tigers? If not, then which pantherine taxon does
P. palaeosinensis most resemble?; and (ii) can the mosaic of craniodental features,
suggesting P. palaeosinensis is ancestral to Panthera, be confirmed in multivariate
analyses using linear metric characters?

METHODS
Samples and craniodental variables
The skulls used in this study were: a cast of the holotype of P. palaeosinensis, 643
adult extant Pantherinae (Neofelis, Uncia and Panthera) skulls and four almost
complete crania of the extinct European cave lion P. (l.) spelaea. The majority of
the samples were measured by the late Dr Vratislav Mazák (V. M.) and donated
kindly to the author by Prof C. P. Groves, and the remainder was measured by the
author (J. H. M). Ideally, other fossil taxa of Panthera [the Eurasian jaguar P. gom-
baszoegensis, or American lion P. (l.) atrox] would have been included, but suffi-
ciently complete specimens were not available for this study.
The holotype of P. palaeosinensis is an almost complete skull (cranium and man-
dible) with distortion in the right upper canine and anterior nasal aperture. Since the
left zygomatic arch is missing, the bizygomatic width was estimated by doubling the
measurement from the right zygomatic arch to the midline.
Twenty craniodental measurements, representing the overall external skull mor-
phology and some function-related cranial regions, were recorded for each skull
(Fig. 1). I reduced the 20 original measurements to 17, which were available for
maximum sample sizes with no statistically significant difference between observ-
ers [a previous investigation, using paired t-tests, had shown that there was no
statistically significant difference (P < 0.05) in measurements between V. M. and J.
H. M, except the Basal length II.; Mazák 2008 doi:10.1016/j.mambio.2008.06.003].
The 17 parameters were grouped into: overall length and width of skull (greatest
skull length, condylobasal length, basal length, bizygomatic width, mandible
length and height), facial region (rostral breadth, infra-orbital breadth, inter-
orbital breadth, nasal length), braincase region (post-orbital constriction), occipital
region (mastoidal breadth, occipital height) and masticatory region (upper carnas-
sial length, upper tooth row length, lower carnassial length, lower tooth row
length). Only specimens for which the complete set of measurements was available
were used in the statistical analysis, reducing the total number of specimens from

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92 J. H. Mazák

Fig. 1. Holotype of P. palaeosinensis, illustrating the 17 craniodental variables used in this paper.
The data given in brackets are the measurements, in mm, taken from the cast of the holotype.

648 to 504, consisting of the holotype of P. palaeosinensis, plus 14 clouded leop-

ards (N. nebulosa), 12 snow leopards (U. uncia), 211 tigers (P. tigris), 29 jaguars (P.
onca), 198 lions (P. leo) and 40 leopards (P. pardus).
In order to include four specimens of P. (l.) spelaea in some analyses, I further
reduced the 17 variables to nine, which were available for all specimens. Ideally, the

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 The Panthera palaeosinensis skull 93

sexes should have been separated, because there is marked sexual dimorphism in
skull morphology in pantherine cats (Pocock 1939, Gittleman & Van Valkenburgh
1997, Mazák 2004). However, the holotype of P. palaeosinensis has never been sexed,
so the sexes are combined in order to maximize the sample sizes for comparison.

Shape variables
Since cats from the subfamily Pantherinae differ greatly in size [mean greatest skull
length varies from 174.1 mm (n = 19) in clouded leopards to 334.1 mm (n = 249) in
lions], I used shape variables (size-adjusted data) to compare the external skull
morphology of P. palaeosinensis with those of its relatives (Jungers et al. 1995,
Mazák 2008 doi:10.1016/j.mambio.2008.06.003). Ideally, the geometric mean would
have been calculated using all 17 raw variables, but using a relatively high number
of measurements (ⱖ12) introduces a danger of confounding the analysis (Coleman
2008). Therefore, I selected six size-related variables (greatest skull length, condylo-
basal length, basal length, bizygomatic width, mandible length and height) as
proxies for general size to calculating the geometric mean as follows:

GM = [n∏ i =1 Xi ]
1n

where GM is geometric mean, [n PI=1 X i]1/n is the Nth root of the product of N
measurements. Shape variables were then calculated by dividing all raw measure-
ments by the geometric mean of the six size-related raw variables and log-10 trans-
forming the data. When P. (l.) spelaea was included, with the reduced number of
variables, the shape variables were calculated by dividing each measurement by the
geometric mean of all nine variables.

Shape analysis
A covariance-based principal component analysis (PCA) was carried out to reveal the
internal variance structure of shape variables and to visualize the overall pattern of
variation in skull morphology of all pantherine samples. Only principal components
(PCs) with eigenvalues greater than 1 were extracted, after which, they were rotated
by varimax criterion. A stepwise discriminant function analysis (DFA) was applied to
test to which pantherine group P. palaeosinensis would be assigned; the pairwise F
statistic tests on the matrix of squared generalized distances (D2) constructed from
group centroids (the means for each taxon on each linear function) were used to
evaluate whether the distance of P. palaeosinensis from the group centroids of
extant and extinct pantherine cats significantly exceeded the distance within the
pantherine cats from their centroid. DFA also produced a jack-knifed classification
that assigns the group association of individual specimens, from which I tested
whether P. palaeosinensis could be distinguished from or included with other taxa.
The differences between specimens were also evaluated using Euclidean distances,
on the basis of the PCs extracted from the PCA. Euclidean distances were calculated
as the mean distances of all members of a group from their own centroid, which
were then rescaled to range from 0 to 1. Using Euclidean distances based on PCA
scores rather than on the shape variables themselves meant that I could provide a
measure of shape differences defined by the variation between samples.
Unweighted pair-group average (Unweighted Pair Group Method with Arithmetic
Mean, UPGMA) cluster analyses, based on the mean squared Euclidean distances

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94 J. H. Mazák

between samples (Sneath & Sokal 1973), were performed to investigate the grouping
patterns among the pantherine cats and to provide a phenetic scale for determining
craniodental similarities or dissimilarities between the species.

RESULTS AND DISCUSSION

In an analysis of 17 log-transformed shape variables, the three-dimensional plot of
principal components (PCs) 1–3 shows P. palaeosinensis in a tight cluster, which
includes all extant Pantherinae. In plots of PCs 2–4, P. palaeosinensis is clearly within
the range of lion skull morphologies, although one tiger and one leopard are nearby
(Fig. 2a, b). This analysis indicates that P. palaeosinensis shares similarities in skull
morphology with all extant Pantherinae, in accordance with previous suggestions
that P. palaeosinensis shows a mixture of craniodental traits of more than one extant
species of Panthera (Zdansky 1924, Hemmer 1967, Christiansen 2008b). Among
modern pantherine cats, the snow leopard and clouded leopard are the most distinct
(Fig. 2a, b), while all Panthera show a high degree of similarity with each other in the
absence of size factors, although differing in some minor details (see Supporting
Information, Fig. S1). A stepwise DFA shows that P. palaeosinensis is located between
the group centroids of lion and leopard (Fig. 3), while a jack-knifed classification
classified P. palaeosinensis with the lion (Supporting Information Table S6). When
comparing differences using a pairwise F statistic based on squared generalized
distances (D2), P. palaeosinensis differs significantly from all modern pantherines, but
shows the shortest distance to the lion (see Table S7); this is consistent with the
results from Euclidean distances (Table 1). The UPGMA cluster analyses, based on
mean squared Euclidean distances between samples for the four PCs, yield three
major clusters: snow leopard, clouded leopard and Panthera; P. palaeosinensis is
clustered with the lion (Fig. 4a).
When the upper Pleistocene European cave lion [P. (l.) spelaea] is included in the
PCA, P. palaeosinensis falls at the boundary between the tiger/cave lion/lion and
leopard sub-cluster (Fig. 2c). The four specimens of P. (l.) spelaea are fully within the
range of the modern lion sample, while the clouded leopard and snow leopard are
shifted away from the Panthera cluster. The jack-knifed classification of stepwise DFA
yields P. palaeosinensis as again classified with the lion (Supporting Information
Table S8). The Euclidean distance based on three PCs shows P. palaeosinensis closest
to the lion and leopard (Table 2). However, the UPGMA tree reveals an interesting
pattern: P. palaeosinensis is clustered with the leopard, while the tiger, cave lion and
lion are clustered together, with the latter two being very closely nested (Fig. 4b).
The results of this study demonstrate that: (i) when shape is considered in the
absence of size, P. palaeosinensis does not show a close affinity to the tiger. Instead,
it appears closest to the lion or leopard (Fig. 5), although it bears a high degree of
similarity with all extant Pantherinae; and (ii) three major morphological patterns of
the skull in the Pantherinae emerge: clouded leopard, snow leopard and Panthera
(tiger, jaguar, lion, leopard and extinct European cave lion). This grouping is consis-
tent with previous findings based on classical description, cladistics and geometric
morphometric analysis (Pocock 1939, Hemmer 1966, 1981, Christiansen 2008a). This
increases confidence that the taxonomic affinities of P. palaeosinensis are correctly
elucidated by the present analysis.
What do the results of this study imply for the taxonomic affinity of P. palaeosin-
ensis and what are the implications for the origin and evolution of the Pantherinae?

© 2010 The Author. Journal compilation © 2010 Mammal Society, Mammal Review, 40, 90–102
 The Panthera palaeosinensis skull 95

(a)

2
PC 3

0 Pantherinae

P.pardus
-2
P.leo

P.onca
4
2 P.tigris

0 4
3 P.uncia
2
PC 2 -2 1
0 N.nebulosa
-4 -1 PC 1
-2
P.palaeosinensis

(b)

2 Pantherinae
PC 3
P.pardus
0
P.leo

-2 P.onca
3
2 P.tigris
1
4 0
2 P.uncia
0 -1
-2 PC 4
-2 N.nebulosa
PC 2 -4 -3

P.palaeosinensis

Fig. 2. Three-dimensional (3D) plots of principal component analysis for the comparison of
Panthera palaeosinensis with extant and extinct pantherines. (a) In a 3D plot of principal
component (PC) 1–3, P. palaeosinensis forms a tight cluster with tigers, lions and jaguars. Within
the extant Pantherinae, the snow leopard stands as the most distinct; (b) In a 3D plot of PC 2–4, P.
palaeosinensis falls within the range of lions, although a tiger is nearby. The clouded leopard is
clearly shifted away from the others; (c) When the extinct European cave lion is included with a
reduction in the number of variables, P. palaeosinensis falls at the boundary between lion and
leopard sub-clusters.

© 2010 The Author. Journal compilation © 2010 Mammal Society, Mammal Review, 40, 90–102
96 J. H. Mazák

(c)

PC 3 0 Pantherinae

-1 P.(l.)spelaea

-2 P.pardus

-3 P.leo

P.onca
4
3
P.tigris
2
1 6 U.uncia
4
PC 2 0 2
-1
0 N.nebulosa
-2 -2 PC 1
P.palaeosinensis

Fig. 2. (Continued)

First, there is no support for regarding P. palaeosinensis as a tiger or ancestor of the

tiger (Hemmer 1967, 1987, Christiansen 2008b), but instead, there is support for the
hypothesis that it represents an ancestor of two or more extant Panthera (Kitchener
1999) or an early form of leopard (Teilhard de Chardin & Leroy 1945, Kurtén 1968).
However, it is difficult to determine the essential relationship of P. palaeosinensis
with its relatives, because it possesses a mixture of craniodental traits from several
extant Pantherinae, the only complete skull of the species is the holotype (Zdansky
1924), and analysis has so far been focused on dental characteristics or non-metric
craniodental features (Hemmer 1967, Christiansen 2008b). There are two possible
hypotheses for interpreting the high degree of similarity in craniodental morphology
among the extant or recently extinct Pantherinae: (i) the pattern of craniodental
morphology is functionally well adapted for the predatory lifestyle of pantherine
cats, so that the degree of variation of this pattern is mechanically constrained,
whatever the range in body size, which is closely related to the sizes of typical prey;
or (ii) the Pantherinae share a relatively recent history and rapid evolutionary radia-
tion, as suggested by genetic analyses (Wayne et al. 1989, Johnson & O’Brien 1997),
thus explaining the high degree of similarity in craniodental traits. Within these tight
constraints, P. palaeosinensis shows a dentition that is close to the tiger’s (Hemmer
1967, Christiansen 2008b), while its overall skull morphology is most similar to that
of the lion or leopard, and this mosaic of characteristics suggests that it may repre-
sent a small form either closely related to the early lion or leopard clade, or that it
is close to the ancestry of the entire genus Panthera. Molecular phylogenies suggest
that pantherine cats probably evolved within the last 5 or 10 million years (Collier &
O’Brien 1985, Johnson et al. 2006) with an Asian origin and radiation (Johnson et al.
2006). A cladistical analysis based on various skeletal and anatomical characters, as

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 The Panthera palaeosinensis skull 97

Canonical Discriminant Functions

10

6
Pantherinae
4
DF 2 (25.5 %)

Group Centroids

P.pardus
2
P.leo

0 P.onca

P.tigris
-2
U.uncia

-4
N.nebulosa

-6 P.palaeosinensis

-8 -6 -4 -2 0 2 4 6

DF 1 (46.8 %)
Fig. 3. Plot of discriminant function (DF) 1–2 of stepwise discriminant function analysis for the
comparison of Panthera palaeosinensis with extant pantherines. Panthera palaeosinensis falls at
the boundary between the centroids of the lion and leopard samples.

Table 1. Euclidean distances based on principal component scores for palaeosinensis and extant
Pantherinae

N Palaeosinensis Nebulosa Uncia Tigris Onca Leo Pardus

Palaeosinensis 1 0.000
Nebulosa 14 0.346 0.000
Uncia 12 0.458 1.000 0.000
Tigris 211 0.255 0.629 0.683 0.000
Onca 29 0.364 0.718 0.482 0.379 0.000
Leo 198 0.000 0.437 0.572 0.124 0.397 0.000
Pardus 40 0.133 0.362 0.611 0.267 0.367 0.278 0.000

well as biogeography, suggested an eastern Asian radiation of the Pantherinae

(Hemmer 1981). These theories are not, however, supported by palaeontological
evidence. The first occurrences of lion and leopard are from the Laetoli Beds in East
Africa, dating from before 3.46 Ma (Turner 1990, Turner & Antón 1997, 2004, Wer-
delin & Lewis 2005). When taking these African specimens into account, the mosaic
pattern of craniodental characteristics in P. palaeosinensis places it as more plesio-
morphic than the apparently earlier Laetoli specimens (Turner 1990, Turner & Antón

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98 J. H. Mazák

(a)

(b)

Fig. 4. Unweighted pair-group average cluster analysis based on squared Euclidean distance from
principal component analysis scores for the comparison of Panthera palaeosinensis with extant
and extinct pantherines. (a) Three major clusters are formed: snow leopard, clouded leopard and
all Panthera, and P. palaeosinensis clusters with the lion; (b) When the extinct cave lion is
included, P. palaeosinensis clusters with the leopard, while the modern lion and cave lion are
grouped in a sub-cluster.

1997), and this potentially supports an Asian origin of the Pantherinae. The out-
standing problem is now to clarify the palaeontological context of the holotype of P.
palaeosinensis [although a maxilla dated between 2.55 and 2.15 Ma was assigned to
P. palaeosinensis (Qiu, Deng & Banyue 2004), its identity needs to be confirmed].
Although P. palaeosinensis has been widely claimed to be of Plio-Pleistocene age (Pei
1934, Teilhard de Chardin & Leroy 1945), according to the original description, the
specimen was found in deposits of Hipparion fauna, which would suggest an Early
Pliocene or even Late Miocene origin (Zdansky 1924, Teilhard de Chardin & Leroy
1942). If this earlier age could be confirmed by further study, then the Asian origin
of Panthera would be strongly supported.
In summary, the morphometric analyses on skull shape variables reported here
(and Supporting Information Figs S2–S3, Tables S1–S9) suggest that P. palaeosinensis
has a close relationship to early lions or leopards, or is close to the ancestry of the
entire genus Panthera, which is in agreement with previous scenarios (Kitchener

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 The Panthera palaeosinensis skull 99

Table 2. Euclidean distance based on principal component scores for palaeosinensis and extant
and fossil Pantherinae

N Palaeosinensis Nebulosa Uncia Tigris Onca Leo Pardus Spelaea

Palaeosinensis 1 0.000
Nebulosa 14 0.452 0.000
Uncia 12 0.426 1.000 0.000
Tigris 211 0.234 0.457 0.580 0.000
Onca 29 0.369 0.586 0.537 0.165 0.000
Leo 198 0.132 0.433 0.574 0.063 0.348 0.000
Pardus 40 0.147 0.272 0.616 0.317 0.352 0.305 0.000
Spelaea 4 0.258 0.386 0.701 0.051 0.356 0.000 0.357 0.000

Fig. 5. Sequential steps in the

reconstruction of the head of Panthera
palaeosinensis based on skull
proportions and modern Panthera facial
musculature show that P. palaeosinensis
is lion-like or leopard-like.

© 2010 The Author. Journal compilation © 2010 Mammal Society, Mammal Review, 40, 90–102
100 J. H. Mazák

1999) and potentially supports an Asian origin for Panthera (Hemmer 1981, Johnson
et al. 2006). Further study, including the dating of the holotype, is required to
confirm these results.

ACKNOWLEDGEMENTS
I am grateful to Prof Colin P. Groves (Canberra), who kindly donated the original data
of skull measurements of Pantherinae taken by the late Dr Vratislav Mazák to me,
and to Velizar Simeonovskii (Chicago) for making the facial reconstruction of P.
palaeosinensis. Thanks go to all colleagues who have helped in data collecting,
especially to Prof Drs Helmut Hemmer (Mainz), Per Christiansen (Copenhagen) and
Alexei Abramov (Saint Petersburg). I am indebted in particular to Drs Andrew Kitch-
ener (Edinburgh) and Lars Werdelin (Stockholm) for reading a preliminary draft of
this paper and providing perspicacious comments which improved the paper greatly.

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Submitted 2 April 2009; returned for revision 8 June 2009; revision accepted 3 September 2009
Editor: KH

SUPPORTING INFORMATION
Additional Supporting Information may be found in the online version of this article:
Fig. S1. 3D plot of PCA for comparing palaeosinensis with extant Pantherinae. (a) on
PC1, palaeosinensis is grouped in a tight cluster which contains lion/tiger/jaguar,
while the snow leopard is separated completely from all others; leopard and clouded
leopard are also somewhat separated from the lion/tiger/jaguar cluster, but with
clear overlap; (b) on PC2, palaeosinensis is grouped within the lion/tiger/leopard
cluster, the jaguar is somewhat separated from others by its proportionally well-
developed occiput; (c) on PC3, the snow leopard is distinctly shifted away from all
others; (d) on PC4, the tiger is separated somewhat from all others by its propor-
tionally long nasal and wider rostrum, and palaeosinensis is grouped with the lion,
although a jaguar is close nearby. In summary, although palaeosinensis shares simi-
larity with all extant Panthera, it is most close to lion.
Fig. S2. When the Late European cave lion is included, PCA extracts three PCs. On PC
1, three main clusters are formed: clouded leopard, leopard/tiger/lion/jaguar and
snow leopard, the snow leopard being the most distinct, while palaeosinensis is
grouped within the Panthera. On PC3, the leopard is separated from the main
Panthera cluster to some extent, while palaeosinensis is localized at the boundary
between lion and leopard.
Fig. S3. Plot of stepwise DFA for comparing palaeosinensis with extant Pantherinae
and extinct late Pleistocene European cave lion P. (l) spelaea. With the reduction in
number of variables, each group shows a relatively wide range, and separation
between groups is less marked. Panthera palaeosinensis plots at the boundary
between the lion and tiger clusters.

© 2010 The Author. Journal compilation © 2010 Mammal Society, Mammal Review, 40, 90–102
102 J. H. Mazák

Table S1. Description and abbreviations of the craniodental measurements used in

this study.
Table S2. Results of principal component analysis based on 17 shape variables mea-
sured from palaeosinensis and 504 extant Pantherinae samples.
Table S3. Factor loadings based on 17 shape variables on palaeosinensis and 504
extant Pantherinae samples (sorted by variable size).
Table S4. Results of principal component analysis based on 17 shape variables on
palaeosinensis and 504 extant Pantherinae samples.
Table S5. Factor loadings based on 17 shape variables on palaeosinensis, 4 late
Pleistocene European cave lion P. (l) spelaea and 504 extant Pantherinae samples
(sorted by variable size).
Table S6. Jack-knifed classification matrix of stepwise DFA on palaeosinensis and 504
extant Pantherinae samples.
Table S7. Pairwise F statistics based on generalized squared distance (D2) between
palaeosinensis and modern Pantherinae.
Table S8. Jack-knifed classification matrix of stepwise DFA on palaeosinensis, 4 late
Pleistocene European cave lions P. (l.) spelaea and 504 extant Pantherinae samples.
Table S9. Pairwise F statistics based on generalized squared distance (D2) between
palaeosinensis and extant Pantherinae and late Pleistocene European cave lion.
As a service to our authors and readers, this journal provides supporting information
supplied by the authors. Such materials may be re-organized for online delivery, but
are not copy-edited or typeset. Technical support issues arising from supporting
information (other than missing files) should be addressed to the authors.

© 2010 The Author. Journal compilation © 2010 Mammal Society, Mammal Review, 40, 90–102