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Biological Journal of the Linnean Society, 2004, 83, 47–63. With 7 figures
© 2004 The Linnean Society of London, Biological Journal of the Linnean Society, 2004, 83, 47 – 63 47
Blackwell Science, LtdOxford, UKBIJBiological Journal of the Linnean Society0024-4066The Linnean Society of London, 2004? 2004
831
4763
Original Article
WILDCAT EVOLUTION AND CONSERVATION
N. YAMAGUCHI
ET AL.
*Corresponding author. E-mail: nobuyuki.yamaguchi@
zoo.ox.ac.uk
Craniological differentiation between European wildcats
(Felis silvestris silvestris), African wildcats (F. s. lybica)
and Asian wildcats (F. s. ornata): implications for their
evolution and conservation
NOBUYUKI YAMAGUCHI1*, CARLOS A. DRISCOLL1,3, ANDREW C. KITCHENER2,
JENNIFER M. WARD2 and DAVID W. MACDONALD1
1Wildlife Conservation Research Unit, Department of Zoology, South Parks Road, Oxford OX1 3PS, UK
2National Museums of Scotland, Chambers Street, Edinburgh EH1 1JF, UK
3Laboratory of Genomic Diversity, National Cancer Institute, Frederick, MD 21702–1201, USA
Received 30 July 2003; accepted for publication 19 January 2004
Intraspecific diversification of the wildcat (Felis silvestris), including the European wildcat (F. s. silvestris), the Asian
wildcat (F. s. ornata) and the African wildcat (F. s. lybica), was examined based on 39 cranial morphology variables.
The samples of free-ranging cats originated from Britain, Europe, Central Asia and southern Africa, consisting of
both nominal wildcat specimens (referred to henceforth as ‘wildcats’) and nominal non-wildcat specimens (‘non-wild-
cats’) based on museum labels. The skull morphology of ‘wildcats’ from Britain and Europe is clearly different from
that of ‘wildcats’ of Central Asia and southern Africa. The latter are characterized especially by their proportionately
larger cheek teeth. On the basis of principal component, discriminant function and canonical variate analyses, the
skull morphology of British ‘non-wildcats’ is less distinct than is that of British ‘wildcats’ from the skull morphologies
of ‘wildcats’ of Central Asia and southern Africa. On the other hand, the skull morphology of southern African ‘non-
wildcats’ is as distinct from those of ‘wildcats’ of Britain and Europe as is that of southern African ‘wildcats’. We sug-
gest that the evolution of the modern wildcat probably consisted of at least three different distribution expansions
punctuated by two differentiation events: the exodus from Europe during the late Pleistocene, coinciding with the
emergence of the steppe wildcat lineage (phenotype of Asian–African wildcat), followed by its rapid range expansion
in the Old World. The second differentiation event was the emergence of the domestic cat followed by its subsequent
colonization of the entire world with human assistance. Considering the recent evolutionary history of, and mor-
phological divergence in, the wildcat, preventing hybridization between the European wildcat and the domestic cat
is a high conservation priority. © 2004 The Linnean Society of London, Biological Journal of the Linnean Society,
2004, 83, 47–63.
ADDITIONAL KEYWORDS: Felidae – Felis lunensis – F. lybica – F. ornata – F. silvestris – forest cat –
hybridization – morphology – Pleistocene – steppe cat.
INTRODUCTION
The wildcat (Felis silvestris Schreber, 1777) is distrib-
uted widely throughout Europe, Africa and Asia (Now-
ell & Jackson, 1996). The existence of distinguishable
phenotypes across this wide distribution, along with
apparently considerable local variation (Pocock, 1951;
Haltenorth, 1953), has left wildcat taxonomy in a state
of confusion for many years (Guggisberg, 1975). It has
become customary to consider a single species of the
wildcat with three main morphological types: the
European wildcat (F. s. silvestris), the African wildcat
(F. s. lybica Forster, 1780) and the Asian wildcat
(F. s. ornata Gray, 1830) (Guggisberg, 1975; Hemmer,
1978; Kitchener, 1991; Nowell & Jackson, 1996).
According to earlier morphological studies (Pocock,
1951; Roberts, 1951; Haltenorth, 1953; Heptner &
48 N. YAMAGUCHI ET AL.
© 2004 The Linnean Society of London, Biological Journal of the Linnean Society, 2004, 83, 47 – 63
Sludskii, 1972; Guggisberg, 1975), the three wildcat
groups possess external characteristics that are fairly
distinct from each other and ubiquitous within each
group. The European wildcat has been characterized
by its bushy tail, which ends as a broadly rounded
black tip, and long soft fur with relatively conspicuous
stripes, and is distributed in Britain, Europe including
some Mediterranean islands, and part of south-west-
ern Asia (Fig. 1). The African wildcat is widely distrib-
uted in Africa, the Arabian Peninsula, part of south-
western Asia and other Mediterranean islands. It has
a tapering tail, a reddish or rusty-brown tint to the
back of the ears and inconspicuous body stripes com-
pared with the European wildcat. The domestic cat
[F. s. catus Linnaeus, 1758: concerning the use of dif-
ferent scientific names for the wildcat and the domes-
tic cat see Opinion 2027 (Case 3010) (International
Commission on Zoological Nomenclature, 2003)] is
widely believed to have descended from the African
wildcat (and perhaps some Asian wildcat populations)
and historical records suggest that the domestic cat
first appeared in Egypt approximately 4000 years ago.
The Asian wildcat is the least known and inhabits
Eurasia from the Middle East to western India,
through Central Asia to Mongolia and north-western
China. It is similar to the African wildcat, but is dis-
tinguishable by its coat pattern, which consists mainly
of distinct dark spots instead of stripes.
In the last 20 years, advances in biomolecular tech-
niques and the accessibility of powerful statistical
analyses have yielded several important results
regarding the phylogeny of the wildcat. Such studies
have focused mainly on taxonomy-related conserva-
tion problems of the wildcat concerning hybridization
between its local populations and free-ranging domes-
tic cats in Scotland (French, Corbett & Easterbee,
1988; Daniels et al., 1998; Beaumont et al., 2001;
Daniels et al., 2001; Reig, Daniels & Macdonald,
2001), continental Europe (Randi & Ragni, 1986,
1991; Randi et al., 2001; Pierpaoli et al., 2003), or
Figure 1. Distribution of the wildcat. Skulls used for this study originated from four geographically separate regions
consisting of Britain (A), Europe (B), Central Asia (C) and southern Africa (D).
Felis silvestris silvestris F. s. lybica
F. s. ornata
A
B
C
D
WILDCAT EVOLUTION AND CONSERVATION 49
© 2004 The Linnean Society of London, Biological Journal of the Linnean Society, 2004, 83, 47 – 63
southern Africa (Wiseman, O’Ryan & Harley, 2000).
These recent studies have also suggested, largely
independently of the earlier taxonomic work, that
European wildcats, African wildcats and domestic cats
are phylogenetically very close, such that they belong
to a single polytypic species. These data suggest that
the European and African wildcats may have diverged
from each other as recently as 20 000 years ago, and
that the domestic cat is phylogenetically closer to the
African than it is to the European wildcat (Collier &
O’Brien, 1985; Randi & Ragni, 1986, 1991; Johnson &
O’Brien, 1997). However, because most of these stud-
ies are based on regional populations either in Scot-
land or in Italy, there has been no elucidation of the
intraspecific phylogeny of the wildcat, especially con-
cerning the position of the Asian wildcat, which until
now has escaped study owing to the paucity of data on
many quantitative parameters. Furthermore, a biolog-
ically coherent approach to the conservation of extant
wildcats necessitates understanding their evolution-
ary history, especially when current taxonomic dis-
tinctions, whether based on traditional morphology or
molecular biology, are proving elusive (French et al.,
1988; Kitchener, 1998; Daniels et al., 2001).
In this paper, we aim to explore and discuss the phy-
logenetic relationships amongst European, African
and Asian wildcats based on their skull morphology,
against a backdrop of the probable recent evolutionary
history of these taxa. All samples studied here origi-
nated from populations separated either by sea or by
great geographical distances, so that we consider
there has not been recent regular gene flow, which
may result in similar phenotypes, between any com-
bination of the sampled populations. Thus, we have
eliminated the potential confounding effects concern-
ing possible recent population mixtures and isolations,
of which there is currently only a poor understanding.
MATERIAL AND METHODS
SPECIMENS
The morphological investigation was undertaken
using 218 skulls of free-ranging cats from Britain and
Europe (originating from Germany, France, Spain,
Italy, Hungary, Austria, Switzerland, Slovenia and
Romania), 145 from southern Africa (South Africa,
Namibia, Botswana, Zimbabwe, Malawi, Mozambique
and Zambia), and 103 from Central Asia (Uzbekistan,
Kazakhstan and Kyrgyzstan) (Fig. 1) from museum
collections. The skulls of the European cats were
examined by two of the authors (N.Y. & J.M.W.) and
the others by one author (N.Y.). Only adult and sub-
adult skulls, assessed from the fusion of skull sutures
(Daniels et al., 1998), were included in the analyses.
Although the very existence of pristine wildcats and
possible identification of their characteristics have
been the subject of extensive debate (Daniels et al.,
1998; Kitchener, 1998; Daniels et al., 2001; Reig et al.,
2001; Yamaguchi et al., in press), cats were nominally
classified as either ‘wildcat’ or ‘non-wildcat’ based on
museum labels. Being mindful of the debate over what
really constitutes a true wildcat (Yamaguchi et al., in
press), we are careful throughout the text, to use quo-
tation marks, such as ‘wildcat’, when we refer to the
nominal categories labelled in the museum collections.
Because these specimens were identified at various
times throughout a period of more than 100 years, it is
impossible to know exactly on what basis these dis-
tinctions were made; however, they were likely deter-
mined from external morphology, especially pelage
(e.g. Pocock, 1951; Haltenorth, 1953; Heptner & Slud-
skii, 1972; Smithers, 1983; Kitchener, 1998).
Pooling all regional free-ranging cats, which very
probably include ubiquitous domestic cats and their
hybrids, may mask any morphological differences
between wildcats from different geographical regions.
Although defining a wildcat based purely on external
morphology may be problematic, morphological diver-
sity appears to reflect distinct gene pools both in Brit-
ish and European (Beaumont et al., 2001; Randi et al.,
2001) and in southern African (Wiseman et al., 2000)
populations. For example, strict possession of the coat
coloration and markings classically taken to charac-
terize wildcats appears to be sufficient to place an
individual in the non-domestic genetic group in the
Scottish population (Beaumont et al., 2001). There-
fore, we assumed that, to a certain extent, basing
analyses on nominal ‘wildcats’ alone should reduce the
possibility of regional morphological differences being
masked.
SKULL PARAMETERS
The following five skull characteristics, which have
been traditionally used to distinguish European wild-
cats from domestic cats (Pocock, 1951; Kitchener,
1995), were scored (1–3) for each specimen (Fig. 2):
1. shape of the anterior end of the nasals;
2. extent of a pit at the posterior end of the nasals;
3. shape of the parietal suture;
4. length of the nasals relative to the maxillae;
5. whether the mandibles tip over on a horizontal
surface.
The scores of two observers using ten randomly
selected skulls agreed totally for characters -3 and -5,
disagreed by a score of 1 in one case each for charac-
ters -1 and -4 and in two cases for character -2; they
never disagreed by a score of 2 for any character. All
five scores were summed as the total skull score and
used in analyses.
50 N. YAMAGUCHI ET AL.
© 2004 The Linnean Society of London, Biological Journal of the Linnean Society, 2004, 83, 47 – 63
Forty-four numerical measurements (modified
from French et al., 1988) of the cranium and mandi-
ble were taken. The cranial volume was measured
using either steel shot approximately 1 mm in
diameter or glass beads approximately 2 mm in
diameter. The volume measured with the smaller
steel shot was consistently very slightly greater
than that measured with the glass beads. There-
fore, we converted the former to the latter using
the equation:
volume (glass beads) = 1.017 ¥ volume
(steel shot) - 1.214,
based on regression coefficients calculated for volu-
metric measurements for both methods on ten
randomly selected skulls (R2 = 0.998, t = 70.63,
Figure 2. Diagram indicating how to score the five skull characteristics.
Score
Character
321
1
4
2
3
5
WILDCAT EVOLUTION AND CONSERVATION 51
© 2004 The Linnean Society of London, Biological Journal of the Linnean Society, 2004, 83, 47 – 63
P < 0.0001). All other parameters were measured to
the nearest 0.02 mm using metal calipers. To test for
measurement errors both by an individual observer
and between observers, six skulls were randomly
selected and each measurement was taken six times
on each skull. The coefficient of variation for each
variable was calculated (Lynch et al., 1996). Vari-
ables with a mean coefficient of variation of more
than 2% were excluded from further analyses. As a
result 31 numerical variables were retained for anal-
ysis (Fig. 3 and Appendix 1). There were significant
differences between the two observers in the maxi-
mum length of the nasals (paired t-test, N = 6 skulls,
d.f. = 5, t = 3.57, P = 0.016) and the depth of the
mandible behind M1 (t = -2.71, P = 0.042) although
much less consistently compared with that in the
cranial volume, so these were also removed from fur-
ther analyses.
In addition to these variables, five derived variables
were calculated: (1) cranial index by dividing the
greatest length of the skull by cranial volume
(Schauenberg, 1969); (2) broadness of the nuzzle by
dividing the distance between the infraorbital fora-
mena by lateral length of snout; (3) ratio of palatal
breadth to Pm2-M1; (4) ratio of postorbital constriction
to interorbital breadth; (5) ratio of the distance
Figure 3. Skull variables measured during the study. The numbers correspond to those in the text.
12
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
2021
22
23
24
25
26
27
28
52 N. YAMAGUCHI ET AL.
© 2004 The Linnean Society of London, Biological Journal of the Linnean Society, 2004, 83, 47 – 63
between pogonion and coronoid process to that
between pogonion and angular process.
STATISTICAL ANALYSES
All statistical analyses were carried out using the
Statistica statistics package (Statsoft, Tulsa, USA).
A principal component analysis (PCA) was carried
out using all free-ranging cats together mainly to
reduce the numbers of variables for the subsequent
analyses. Specimens with any missing value were
removed from the analysis, and hence from the sub-
sequent analyses too, reducing the sample size to 80
British cats (39 ‘wildcats’ and 41 ‘non-wildcats’), 22
European cats (19, 3), 58 Central Asian cats (57, 1)
and 89 southern African cats (80, 9). Because of the
resultant small sample sizes for both European and
Central Asian ‘non-wildcats’, these were excluded
from the later analyses. Discriminant function and
canonical variate analyses were carried out to inves-
tigate if the four geographically separated groups
could be distinguished based on skull morphology
using all principal components with eigenvalues
greater than 1 (Tabachnick & Fidell, 2001). The four
groups a priori consisted of cats originating in Brit-
ain, Europe, Central Asia and southern Africa. Fur-
thermore, to enable future use of standard
measurements for the classification of skulls, each
variable for ‘wildcats’ originating from the four geo-
graphical areas was also analysed and statistically
significant differences were detected using Kruskal–
Wallis tests and ANOVAs.
RESULTS
A PCA based on total skull score, 29 measured vari-
ables and five derived variables resulted in six princi-
pal components (Table 1). The first component (PC1)
was probably related to the overall size of the skull
along the anteroposterior axis, PC2 to overall breadth
of the skull along the mediolateral axis, PC3 to cranial
capacity, PC4 to characteristics concerning the middle
part of the skull, including teeth, along the anteropos-
terior axis, PC5 to characteristics concerning the mid-
dle part of the skull along the mediolateral axis, and
PC6 to the anterior part of the skull. The PCA quite
clearly separated British and European ‘wildcats’ from
those originating from Central Asia and southern
Africa without any attempt to discriminate those
groups (Fig. 4). However, the range of PC scores for
British ‘non-wildcats’ spread well beyond those of Brit-
ish ‘wildcats’ and overlapped with those of Central
Asian and southern African ‘wildcats’ (Fig. 4). In con-
trast, the range of PC scores for southern African ‘non-
wildcats’ overlapped extensively with those of south-
ern African ‘wildcats’ (Fig. 4).
A discriminant function analysis (DFA) successfully
classified most (c. 91%) ‘wildcat’ skulls into one of the
four geographical groups from which each skull origi-
nated, on the basis of the six PCs (Table 2). The canon-
ical variate analysis (CVA), which produced three
canonical variates, showed good separation between
the skulls of British and European ‘wildcats’ and those
from Central Asia and southern Africa (Fig. 5). A sum-
mary of the overall similarity relationships amongst
the four groups was obtained from the squared Mahal-
anobis distance (D2) between the centroid of each
group on the basis of the three canonical variates
extracted in the analysis. The UPGMA (unweighted
pair-group method using arithmetic average) tree
built from the pairwise D2 similarity matrix placed
British and European ‘wildcats’ closest together
whereas Central Asian and southern African ‘wildcats’
were closest to each other (Fig. 6A).
The separation between the four geographically
separated populations became less clear when ‘wild-
cats’ and ‘non-wildcats’ were pooled (Table 3). Also, a
CVA which extracted three CVs did not show a clear
separation between the two larger geographical
groups compared with the result using only ‘wild-
cats’ (Fig. 7). Furthermore, when free-ranging cats
were pooled, the mean D2 between British–Euro-
pean and Central Asian–southern African cats
became less compared with those based only on
‘wildcats’ (Fig. 6B). When southern African ‘non-
wildcats’ were analysed with ‘wildcats’ from Britain,
Europe and Central Asia, the tree was relatively
similar to that of ‘wildcats’ only (Fig. 6C). However,
when British ‘non-wildcats’ were analysed in the
same way, the shape of the tree changed dramati-
cally and the average D2 between the two larger
groups became smaller (Fig. 6D).
The most highly significant differences between
the four groups of ‘wildcats’ were nasal shape, rela-
tive nasal length, total skull score, distance between
infra-orbital foramena divided by lateral snout, pala-
tal breadth divided by Pm2-Pm4, Pm 2-M1, Pm2-Pm4,
auditory bulla length, mandibular Pm3-M1 and man-
Table 1. Results of principal component analysis based on
35 variables with 251 valid cases
Principal
component Eigenvalue
%
explained
%
cumulative
1 16.25 46.43 46.43
2 4.48 12.81 59.24
3 4.07 11.62 70.86
4 1.71 4.70 75.76
5 1.36 3.87 79.63
6 1.09 3.13 82.76
WILDCAT EVOLUTION AND CONSERVATION 53
© 2004 The Linnean Society of London, Biological Journal of the Linnean Society, 2004, 83, 47 – 63
dibular Pm4 breadth (Appendices 2 and 3). In
general, British and European ‘wildcats’ were char-
acterized by proportionately shorter cheek tooth
rows with smaller teeth and a proportionally
broader muzzle compared with Central Asian and
southern African ‘wildcats’. These groupings were
based on the raw variables that showed the most
highly significant differences between the four
groups and were consistent with the results obtained
by PCA, DFA and CVA.
Table 2. Classification matrix obtained by discriminant function analysis, based on the six extracted principal compo-
nents, concerning the probabilities of classifying each cat correctly into one of the four geographically separate populations;
only ‘wildcats’ were included in the analysis
Observed classification
Predicted classification
British European Central Asian Southern African
British 39 0 0 0
European 4 15 0 0
Central Asian 0 0 51 6
Southern African 0 0 8 72
Figure 4. Bivariate plot of the first two principal components of each free-ranging cat, which account for 46.4% and 12.8%,
respectively, of the variance, based on the 35 skull variables. B, Britain; E, Europe; CA, Central Asia and SA, southern
Africa.
-2.5 -2.0 -1.5 -1.0 -0.5 0 0.5 1.0 1.5 2.0 2.5
2.5
2.0
1.5
1.0
0.5
0
-0.5
-1.0
-1.5
-2.0
-2.5
B wild
B non-wild
E wild
E non-wild
CA wild
CA non-wild
SA wild
SA non-wild
Principal component 1
Principal component 2
54 N. YAMAGUCHI ET AL.
© 2004 The Linnean Society of London, Biological Journal of the Linnean Society, 2004, 83, 47 – 63
DISCUSSION
MORPHOLOGICAL SIMILARITIES AMONGST THE
FOUR GROUPS
DFA is designed to develop classification functions to
best classify each specimen by following a priori
groupings, so that it will usually result in fairly good
discrimination between the groups. However, the
results of PCA and raw data comparisons, both of
which were not influenced by a priori groupings, were
consistent with those obtained by the DFA, suggesting
that the results are not merely due to a priori
groupings.
The results suggest that consistent similarities
and differences exist amongst the four geographi-
cally separated ‘wildcat’ populations. All analyses
placed the skulls of British and European ‘wildcats’
closest together, and distinct from those of the Cen-
tral Asian and southern African ‘wildcats’, which
were also similar to each other. The D2 between the
centroid of each group suggest that the extent of
morphological similarity between Central Asian and
southern African ‘wildcats’ is comparable to that
between British and European ‘wildcats’ (Fig. 6).
The geographical distances over land are c. 6500 km
between European and Central Asian populations,
and c. 12 000 km between Central Asian and south-
ern African ones and between European and south-
ern African ones. In addition, Britain has been
separated from the European continent for the last
c. 9500–7500 years based on the estimated world-
wide rise in sea-levels as documented by 14C-dated
borings of corals in the Bahamas, Tahiti and New
Guinea (Yalden, 1999). We have to underline the
fact that there will always be a risk of misinterpre-
tation when phenotypic characteristics are used to
reconstruct an intraspecific phylogeny. Neverthe-
less, the results may suggest that the Central Asian
‘wildcat’ population has had a stronger connection
with the southern African ‘wildcat’ population than
it has with the geographically closer European one.
Similarly, in spite of more than 7000 years of
Figure 5. Bivariate plot of the first two canonical variates, which explain 82.7% and 9.7%, respectively, of all discrimina-
tory power, based on the six principal components. This plot shows overall morphological similarities amongst the ‘wild’
cat skulls originating in the four geographically separate areas. B, Britain; E, Europe; CA, Central Asia and SA, southern
Africa.
B wild
E wild
CA wild
SA wild
Canonical variate 1
Canonical variate 2
5
4
3
2
1
0
-1
-2
-3
-4
-5.0 -2.5 0 2.5 5.0
WILDCAT EVOLUTION AND CONSERVATION 55
© 2004 The Linnean Society of London, Biological Journal of the Linnean Society, 2004, 83, 47 – 63
separation, the skull morphology of European ‘wild-
cats’ still remains more similar to that of British
‘wildcats’ than it does to that of Central Asian ‘wild-
cats’, despite the potential for gene flow between
these two groups via south-west Asia.
We must be cautious in applying results that are
based on only four regional populations more widely to
the intraspecific phylogeny of the three taxonomic
groups of wildcat. Furthermore, we have not
addressed the possibility that regionally-specific selec-
tion pressures have affected skull morphology. For
example, the similarities between Central Asian and
southern African wildcats may be due to morphologi-
cal convergence caused by similar selection pressures,
although we cannot test this speculation. Neverthe-
less, one of the most likely interpretations of our
results is that the three groups of wildcat are not
equally distinct from each other with respect to their
phylogenetic relationships, such that the skulls of
European wildcats appear to be very clearly distinct
from those of African and Asian wildcats. These
results are also in agreement with earlier taxonomic
studies, which, although not based on quantitative
analyses, have suggested a close taxonomic relation-
ship between African and Asian wildcats (e.g. Pocock,
1951). Regarding diagnostic characters, the European
wildcat possesses a proportionally broader anterior
part of the skull with a different shape of the nasals
compared with the Asian and African wildcats, while
the latter two have proportionally and absolutely
larger teeth, longer cheek tooth rows and larger audi-
tory bullae.
Figure 6. UPGMA trees constructed from the matrices of pairwise unbiased squared Mahalanobis distance ( D2) between
the centroid of each group on the bases of the canonical variates extracted in the analysis. Tree (A) includes only ‘wildcats’,
(B) all free-ranging cats from Britain and southern Africa and ‘wildcats’ from Europe and Central Asia, (C) southern
African ‘non-wildcats’ and ‘wildcats’ from the other regions, and (D) British ‘non-wildcats’ and ‘wildcats’ from the other regions.
Squared Mahalanobis Distance
Britain W
Europe W
Central Asia W
Southern Africa W
Britain ALL
Europe W
Central Asia W
Southern Africa ALL
A
B
C
D
Britain W
Europe W
Central Asia W
Southern Africa NW
Britain NW
Europe W
Central Asia W
Southern Africa W
30 24 18 12 6 0
56 N. YAMAGUCHI ET AL.
© 2004 The Linnean Society of London, Biological Journal of the Linnean Society, 2004, 83, 47 – 63
WILDCATS AND WILD-LIVING CATS
The results demonstrate that the morphological dif-
ferences between the skulls of all wild-living cats from
the four regions are not as clear as are those amongst
the different geographical groups of ‘wildcats’, with or
without a priori grouping. Interestingly, this appears
to be caused by British ‘non-wildcats’ but not by those
from southern Africa.
Currently, there is no uncontroversial definition of a
wildcat, either morphological or genetic (Nowell &
Jackson, 1996; Daniels et al., 1998; Beaumont et al.,
Figure 7. Bivariate plot of the first two canonical variates, which explain 78.7% and 9.6%, respectively, of all discrimina-
tory power, based on the six principal components. This plot shows overall morphological similarities amongst the free-
ranging cat skulls originating in the four geographically separate areas. B, Britain; E, Europe; CA, Central Asia and SA,
southern Africa.
B cats
E wild
CA wild
SA cats
Canonical variate 1
Canonical variate 2
5
4
3
2
1
0
-1
-2
-3
-4
-5.0 -2.5 0 2.5 5.0
Table 3. Classification matrix obtained by discriminant function analysis, based on the six extracted principal compo-
nents, concerning the probabilities of classifying each cat correctly into one of the four geographically separate populations;
analysis included ‘wildcats’ from Europe and Central Asia, and both ‘wildcats’ and ‘non-wildcats’ from Britain and southern
Africa
Observed classification
Predicted classification
British European Central Asian Southern African
British 75 0 2 3
European 8 9 2 0
Central Asian 2 0 47 8
Southern African 0 0 10 79
WILDCAT EVOLUTION AND CONSERVATION 57
© 2004 The Linnean Society of London, Biological Journal of the Linnean Society, 2004, 83, 47 – 63
2001; Daniels et al., 2001; Reig et al., 2001), and we
discuss this issue fully elsewhere (Yamaguchi et al., in
press). However, strict possession of the classical wild-
cat pelage is sufficient to place an individual in the
non-domestic genetic group in the Scottish population,
which is unlikely to have very recent domestic cat
ancestry (Beaumont et al., 2001; Daniels, 2001; Randi
et al., 2001). Therefore, the ‘non-wildcat’ category
probably contains a larger proportion of cats that are
domestic cats or with recent domestic ancestry com-
pared with the ‘wildcat’ category. Obviously, one can-
not assume that each museum possesses an unbiased
collection; wild-living cats with obvious domestic coat
phenotypes may not have been added to some collec-
tions in the first place. Also, as the sample size of
southern African ‘non-wildcats’ was small compared
with that of British ‘non-wildcats’, we need to be care-
ful in making generalizations based on these results.
Also, the criteria for defining the non-wildcat group in
southern Africa may have been different to those used
for the British non-wildcats, which could affect the
composition of a priori groups (e.g. museum labels). In
other words, the two non-wildcat groups may not be
considered to be necessarily equivalent. However, the
results do suggest that introgression with domestic
cats may have had greater effects on the skull mor-
phology of British wildcats than on that of southern
African wildcats. The high degree of morphological
similarity between southern African ‘wildcats’ and
‘non-wildcats’ is consistent with the suggested origin
of the domestic cat. Therefore, one of the possible
effects of the supposed worldwide introgression by the
domestic cat (Nowell & Jackson, 1996) may be to shift
the skull morphology of wild-living cats in Britain and
Europe towards that of the more homogeneous wide-
spread African (and probably Asian) wild-living cats.
EVOLUTION OF THE WILDCAT
The modern wildcat (F. silvestris) probably descended
from Martelli’s wild cat (F. (s.) lunensis Martelli, 1906)
which is known from Europe and may date back to as
early as the late Pliocene c. 2 Mya (Kurtén, 1965b,
1968; Kitchener, 1991). Fossil remains suggest that
the transition from Martelli’s wild cat to the modern
wildcat may have occurred during the middle Pleis-
tocene, possibly by oxygen isotope stage (OIS) 11
(c. 0.45–0.35 Mya) (Kurtén, 1965b; García, Arsuaga &
Torres, 1997; Mannion, 1999). In comparison to this
long and contiguous fossil record in Europe, wildcat
fossils are recorded only from the late Pleistocene
onwards (less than c. 130 000 years ago) in Africa
(both north and south of the Sahara) and the Middle
East (Kurtén, 1965a, b; Savage, 1978; Klein, 1986;
Kowalski & Rzebik-Kowalska, 1991; García et al.,
1997), although we are unaware of any published ref-
erences concerning fossil records of the wildcat from
Asia. If we interpret this lack of a fossil record as the
absence of the wildcat in both Africa and the Middle
East, it is possible that the wildcat may have
expanded its range suddenly, rapidly and recently,
during the late Pleistocene. Based on the absolute
dates of fossil sites in the Palestine region and South
Africa (Kurtén, 1965a; Klein, 1986), this rapid expan-
sion may have occurred even as recently as in the last
c. 50 000 years. This timing of range expansion coin-
cides well, on a geological time scale, with the sup-
posed divergence at c. 20 000 years ago between
European wildcats and African wildcats, based on
allozyme electrophoresis data from animals originat-
ing in Italy, Sicily and Sardinia (Randi & Ragni, 1991).
Based on Kurtén (1965b), Klein (1986) and Savage
(1978), Randi & Ragni (1991) suggested that through-
out Asia and Africa the European wildcat phenotype
was replaced by the African wildcat phenotype.
Although we do not necessarily reject their hypothe-
sis, especially concerning wildcat colonization in Asia,
we could not find evidence suggesting a large wave of
character replacement from those original references.
We re-analysed the data (mandibular and dental mea-
surements) published in Kurtén (1965a, b), and care-
fully checked what he suggested in these texts. This
re-evaluation of the work of Kurtén (1965a, b) sug-
gested that late Pleistocene Palestine, less than c.
50 000 years ago, was inhabited by wildcats character-
ized by their proportionately larger teeth, compared
with present-day wildcats in central and northern
Europe and Britain, and possibly late Pleistocene
wildcats of these regions as well. The proportionately
larger teeth in present-day Central Asian–southern
African wildcats (compared with those of European
wildcats) may be explained if these two share a com-
mon ancestor possessing that character in the geolog-
ically recent past. This suggests that Asian and
African wildcats are both derived from a large-toothed
wildcat such as the one that inhabited Palestine dur-
ing the late Pleistocene. For the African wildcat this
scenario may be even more realistic. The wildcat that
had been in Europe could not spread into Africa with-
out crossing the Middle East and then the narrow
Sinai Peninsula (or another narrow land bridge which
may have existed between the Arabian Peninsula and
Africa) even during the late Pleistocene glacial max-
ima (Stringer, 2000). As a result, it must have experi-
enced a bottleneck (with probable associated genetic
and phenotypic consequences) before it colonized the
entirety of Africa from the Middle East. However,
there may have been more than one migration route
between Europe and the Asian steppe. Interestingly,
the current Central Asian wildcat is reported to be
unable to cope with low temperatures (Heptner &
Sludskii, 1972). Its winter coat does not attain the lev-
58 N. YAMAGUCHI ET AL.
© 2004 The Linnean Society of London, Biological Journal of the Linnean Society, 2004, 83, 47 – 63
els of density, length and luxuriance seen in British,
central European and Caucasian populations of the
European wildcat in spite of severe winter air temper-
atures as low as -40 ∞C in the northern part of its dis-
tribution (Heptner & Sludskii, 1972; A. C. Kitchener,
pers. observ.). This evidence may suggest that the cur-
rent Central Asian wildcat has recently originated
from a warmer region. If so, on the basis of the current
distribution of the species (Fig. 1), it may have come
from the Middle East, where intergradation occurs
today between African and Asian wildcat phenotypes
in the northern Arabian Peninsula and south-west
Asia (Harrison & Bates, 1991). This hypothesis, colo-
nization through the Middle East, would explain well
the high level of similarity in skull morphology
between Central Asian and southern African ‘wildcats’
observed in this study.
Therefore, the evolution of the modern wildcat prob-
ably consisted of at least three different range expan-
sions punctuated by two differentiation events.
Firstly, during the late Pleistocene (possibly by c.
50 000 years ago) it moved out of Europe which had
been the centre for wildcat evolution for nearly two
million years, and this may have coincided with the
emergence of the steppe wildcat phenotype, which col-
onized the Middle East, i.e. the exodus from Europe.
Secondly, the late Pleistocene Middle Eastern wildcat
quickly spread eastward to Asia and southward to
Africa possibly within the order of a few 10 000 years,
i.e. the steppe wildcat wave. However, as Asian and
African wildcats possess consistently distinct coat pat-
terns (e.g. Pocock, 1951), this stage may have involved
more than one wave of expansion or a series of expan-
sions and contractions possibly affected by the late
Pleistocene glacial–interglacial cycles. Thirdly, the
domestic cat was derived from one or more Middle
Eastern/north African steppe wildcat populations by c.
4000 years ago, followed by its colonization of the
entire world with human assistance, i.e. the domestic
cat wave.
IMPLICATIONS FOR CONSERVATION
Our results suggest that the wildcat comprises two
major lineages, i.e. the steppe wildcat and forest wild-
cat lineages, as suggested by Heptner & Sludskii
(1972). Arguably the most serious current threat to
wildcats is introgressive hybridization with sympatric
domestic cats (Nowell & Jackson, 1996; Daniels, 2001).
The domestic cat colonization wave has resulted
mostly from human activities, so that if indigenous
wildcat populations are to continue to exist relatively
unaffected by human-caused disturbances, this prob-
lem must be tackled. However, we may not easily be
able to distinguish between steppe wildcats and free-
ranging domestic cats surviving in the drier habitats
of north-eastern Africa, the Arabian Peninsula and
south-west Asia, from where the first domestic cats
may have originated. It is even possible that some of
the extant populations of the region may derive in
large part from early forms of feral domestic cat. The
further that steppe wildcat populations occur from the
centre of domestication, the more crucial the potential
impact of hybridization might become. However,
although we can only speculate, the higher degree of
similarity in the skull morphology between ‘non-
wildcats’ from southern Africa and ‘wildcats’ from Cen-
tral Asia and southern Africa suggests that hybridiza-
tion with domestic cats would have less marked effects
in steppe wildcat populations than it would in forest
wildcat populations. Our results, with the re-evalua-
tion of the work by Kurtén (1965a, b), suggest that the
steppe wildcat may not have colonized Europe, at least
not on a large scale. Instead, it expanded into Africa
and Asia. Therefore, bringing domesticated steppe
wildcats into Europe and Britain, and allowing them
to range freely, has created an interface between the
two strands of wildcat evolution for the first time in
their evolutionary history. Given its apparently signifi-
cant impact on the skull morphology of the forest
wildcat, minimizing introgressive hybridization
between forest wildcats and domestic cats should be
regarded as a high conservation priority.
ACKNOWLEDGEMENTS
We thank the Nigel Easterbee Memorial Fund for
financial support, together with grants to D.W.M. from
UFAW, Care for the Rare (Justerini & Brooks) and the
PTES. We also thank Gus Mills and Mike Daniels for
useful comments, Paul Johnson for statistical advice,
Susan Leitch for helping to input data into the com-
puter, and P. Jenkins at the British Museum of Natu-
ral History, London, R. Angermann at Museum für
Naturkunde der Humboldt-Universität, Berlin, R.
Hutterer at Zoologisches Forschungsinstitut und
Museum Alexander Koenig, Bonn, G. Storch at Fors-
chungsinstitut und Naturmuseum Senckenberg,
Frankfurt, L. Peregovits at Hungarian Natural His-
tory Museum, Budapest, L. Szemethy at Department
of Wildlife Biology & Management, University of Agri-
cultural Sciences, Gödöllö, E. Randi at Instituto Nazi-
onale per la Fauna Selvetica, Bologna, B. Herzig at
Naturhistorisches Museum Wien, A. Oakeley at
Naturhistrisches Museum, Basel, A. Rol at Zoölogisch
Museum, University of Amsterdam, C. Smeenk at
Nationaal Natuurhistorisch Museum, Leiden,
F. Uribe at Museu de Zoologia, Barcelona, J. Barreiros
at Museo Nacional de Ciencias Naturales, Madrid, J.
Cuisin at Muséum National d’Histoire Naturelle,
Paris, France, W. Cotteril at Natural History Museum,
Bulawayo, D. MacFadyen at Transvaal Museum, Pre-
WILDCAT EVOLUTION AND CONSERVATION 59
© 2004 The Linnean Society of London, Biological Journal of the Linnean Society, 2004, 83, 47 – 63
toria, F. Kigozi at Amathole Museum, King William’s
Town, A. Esipov & E. Bykova at Institute of Zoology,
Tashkent, and V. Gromov & V. Kascheev at Institute of
Zoology, Almaty, for their kind support for the access
to their collections.
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APPENDIX 1
Measurements of crania and mandibles. Numbers cor-
respond measurements shown in Figure 3.
(1) greatest length of skull
(2) condylobasal length
(3) facial length
(4) lateral length of snout
(5) length between Pm2 and M1
(6) length between Pm2 and Pm4
(7) greatest length of Pm4
(8) greatest breadth of Pm4
(9) anteroposterior diameter of the auditory bulla
(10) mastoid breadth
(11) greatest breadth of the occipital condyles
(12) greatest breadth of the foramen magnum
(13) greatest width of the braincase
(14) zygomatic breadth
(15) frontal breadth
(16) least breadth between the orbits
(17) greatest palatal breadth
(18) rostrum breadth: greatest breadth between the
canine alveoli
(19) least breadth of the postorbital constriction
(20) breadth between the infraorbital foramina
(21) minimum length of the nasals
(22) maximum length of the nasals
(23) width of cranial suture
(24) maximum distance between pogonion and coro-
noid process
(25) maximum distance between pogonion and angu-
lar process
(26) length between mandibular Pm3 and M1
(27) depth of the mandible behind M1
(28) height of ramus
(29) maximum width of mandibular condyles (not
shown in Fig. 2)
(30) Maximum width of mandibular Pm4 (not shown
in Fig. 2)
(31) cranial volume (not shown in Fig. 2)
APPENDIX 2
Differences in skull characteristic scores amongst the four geographical groups of ‘wildcats’. Statistically significant
differences were detected by Kruskal–Wallis tests (d.f. = 3 for all tests).
Variables
Average (Range, number examined)
HPUK Europe Central Asia Southern Africa
Nasal shape 2.6 (1–3, 81) 2.2 (1–3, 111) 1.3 (1–3, 101) 1.9 (1–3, 139) 133.6 <0.0001
Nasal pit 2.8 (2–3, 83) 2.5 (1–3, 130) 2.3 (1–3, 102) 2.4 (1–3, 139) 23.4 <0.0001
Parietal suture 2.9 (2–3, 85) 2.3 (1–3, 133) 2.5 (1–3, 87) 2.2 (1–3, 142) 56.5 <0.0001
Nasal length 2.5 (1–3, 84) 2.2 (1–3, 132) 1.7 (1–3, 102) 1.5 (1–3, 145) 142.6 <0.0001
Mandible 3 (75) 2.8 (1, 3, 119) 2.8 (1, 3, 89) 2.2 (1, 3, 132) 73.1 <0.0001
Total 13.9 (11–15, 71) 12.3 (5.5–15, 95) 10.7 (7–13, 74) 10.1 (5–14, 121) 175.2 <0.0001
WILDCAT EVOLUTION AND CONSERVATION 61
© 2004 The Linnean Society of London, Biological Journal of the Linnean Society, 2004, 83, 47 – 63
APPENDIX 3
Differences in skull measurements amongst the four geographical groups of ‘wildcats’
Variable
Mean ± Standard Error (Number examined)
FPUK Europe Central Asia Southern Africa
Greatest length
Male 99.54 ± 0.63 (30) 100.20 ± 0.63 (60) 105.26 ± 0.58 (63) 103.55 ± 0.55 (73) 17.96 <0.0001
Female 92.96 ± 0.78 (22) 91.63 ± 0.58 (15) 95.86 ± 0.68 (39) 98.20 ± 0.52 (67) 16.89 <0.0001
Condylobasal length
Male 92.58 ± 0.67 (32) 92.12 ± 0.54 (54) 99.68 ± 0.50 (63) 95.93 ± 0.60 (58) 39.22 <0.0001
Female 85.93 ± 0.58 (25) 84.75 ± 0.57 (15) 91.20 ± 0.64 (39) 91.33 ± 0.50 (55) 25.56 <0.0001
Facial length
Male 37.13 ± 0.37 (32) 38.24 ± 0.31 (59) 39.66 ± 0.32 (62) 37.98 ± 0.29 (56) 9.95 <0.0001
Female 34.01 ± 0.33 (25) 34.96 ± 0.40 (16) 35.67 ± 0.43 (39) 36.17 ± 0.27 (54) 6.39 0.0005
Lateral snout
Male 24.82 ± 0.21 (33) 25.17 ± 0.17 (64) 26.70 ± 0.18 (62) 25.70 ± 0.20 (57) 17.49 <0.0001
Female 22.69 ± 0.18 (25) 22.67 ± 0.18 (17) 24.02 ± 0.24 (39) 24.37 ± 0.20 (55) 13.69 <0.0001
Pm2-M1
Male 22.08 ± 0.12 (32) 22.51 ± 0.13 (51) 24.37 ± 0.16 (58) 24.82 ± 0.16 (57) 74.01 <0.0001
Female 21.08 ± 0.12 (24) 21.41 ± 0.19 (16) 23.03 ± 0.15 (33) 23.83 ± 0.13 (52) 72.46 <0.0001
Pm2-Pm4
Male 21.10 ± 0.13 (33) 21.50 ± 0.11 (53) 23.04 ± 0.16 (58) 23.76 ± 0.16 (57) 62.99 <0.0001
Female 20.06 ± 0.13 (24) 20.36 ± 0.24 (16) 21.81 ± 0.14 (33) 22.72 ± 0.11 (52) 75.19 <0.0001
Pm4 length
Male 11.17 ± 0.07 (34) 11.36 ± 0.08 (62) 12.03 ± 0.09 (63) 11.74 ± 0.08 (58) 18.41 <0.0001
Female 10.47 ± 0.08 (24) 10.72 ± 0.17 (17) 11.47 ± 0.09 (39) 11.32 ± 0.08 (55) 19.55 <0.0001
Pm4 breadth
Male 6.01 ± 0.05 (34) 5.67 ± 0.06 (58) 6.25 ± 0.07 (63) 5.97 ± 0.07 (58) 13.60 <0.0001
Female 5.55 ± 0.06 (25) 5.16 ± 0.09 (16) 5.91 ± 0.06 (39) 5.54 ± 0.06 (54) 15.12 <0.0001
Auditory bulla
Male 20.86 ± 0.17 (34) 20.83 ± 0.13 (56) 22.46 ± 0.15 (62) 23.28 ± 0.17 (58) 57.50 <0.0001
Female 19.93 ± 0.19 (24) 19.59 ± 0.23 (17) 20.75 ± 0.19 (39) 22.43 ± 0.13 (54) 52.83 <0.0001
Mastoid breadth
Male 44.77 ± 0.23 (33) 44.01 ± 0.23 (53) 46.32 ± 0.20 (63) 45.47 ± 0.23 (57) 20.68 <0.0001
Female 42.60 ± 0.25 (25) 41.10 ± 0.38 (16) 42.88 ± 0.27 (39) 43.44 ± 0.23 (54) 8.84 0.0002
Occipital condyles
Male 24.79 ± 0.13 (33) 23.96 ± 0.17 (55) 23.96 ± 0.11 (63) 24.35 ± 0.13 (58) 6.54 0.0003
Female 23.60 ± 0.14 (25) 22.98 ± 0.25 (16) 22.72 ± 0.13 (39) 23.35 ± 0.13 (55) 6.33 0.0005
Statistically significant differences were detected by ANOVA (d.f. = 3 for all tests). Measured variables in mm unless stated.*Measurement was taken by N.Y. only
and the coefficient of variation was 0.36% and d.f. = 2. **Measurement was taken by N.Y. only and the coefficient of variation was 0.71% and d.f. = 2. Additional
derived variables were calculated by dividing the six measured variables with the highest F values by the greatest length of the skull.
62 N. YAMAGUCHI ET AL.
© 2004 The Linnean Society of London, Biological Journal of the Linnean Society, 2004, 83, 47 – 63
Foramen magnum
Male 15.18 ± 0.10 (33) 14.97 ± 0.12 (54) 14.79 ± 0.08 (63) 15.14 ± 0.09 (58) 3.23 0.023
Female 14.79 ± 0.12 (25) 14.60 ± 0.21 (16) 14.36 ± 0.09 (39) 14.63 ± 0.09 (55) 2.42 0.069
Brain case
Male 47.12 ± 0.21 (32) 45.87 ± 0.16 (58) 46.90 ± 0.14 (63) 47.36 ± 0.20 (58) 15.09 <0.0001
Female 45.83 ± 0.26 (22) 44.83 ± 0.35 (16) 45.40 ± 0.19 (39) 46.28 ± 0.17 (55) 7.09 0.0002
Zygomatic breadth
Male 70.48 ± 0.57 (34) 71.26 ± 0.52 (55) 74.90 ± 0.46 (62) 72.60 ± 0.51 (54) 14.13 <0.0001
Female 66.42 ± 0.63 (25) 65.13 ± 0.47 (15) 67.89 ± 0.62 (39) 68.37 ± 0.37 (53) 5.52 0.001
Frontal breadth
Male 49.35 ± 0.44 (32) 52.01 ± 0.38 (55) 53.89 ± 0.40 (59) 53.74 ± 0.44 (51) 20.23 <0.0001
Female 48.23 ± 0.49 (24) 48.89 ± 0.73 (17) 50.02 ± 0.64 (34) 51.85 ± 0.37 (51) 12.33 <0.0001
Interorbital breadth
Male 19.43 ± 0.25 (33) 19.32 ± 0.17 (60) 20.66 ± 0.16 (61) 19.25 ± 0.20 (57) 13.96 <0.0001
Female 18.42 ± 0.21 (25) 17.45 ± 0.25 (17) 18.66 ± 0.18 (39) 18.18 ± 0.15 (55) 5.11 0.002
Palatal breadth
Male 41.10 ± 0.25 (34) 40.44 ± 0.25 (59) 42.65 ± 0.21 (60) 40.73 ± 0.20 (55) 20.87 <0.0001
Female 38.85 ± 0.27 (25) 37.80 ± 0.35 (16) 39.64 ± 0.25 (39) 38.94 ± 0.21 (54) 5.90 0.0008
Rostrum breadth
Male 24.09 ± 0.18 (34) 23.78 ± 0.20 (62) 25.50 ± 0.18 (61) 24.71 ± 0.17 (57) 17.55 <0.0001
Female 22.31 ± 0.20 (25) 21.44 ± 0.22 (16) 23.00 ± 0.24 (39) 23.13 ± 0.17 (55) 9.01 0.0002
Postorbital constriction
Male 33.52 ± 0.26 (32) 32.93 ± 0.27 (61) 33.58 ± 0.21 (63) 35.18 ± 0.27 (55) 15.56 <0.0001
Female 34.29 ± 0.32 (25) 32.86 ± 0.57 (17) 33.81 ± 0.21 (39) 35.28 ± 0.22 (55) 11.77 <0.0001
Between infraorbital foramena
Male 29.22 ± 0.21 (33) 28.17 ± 0.22 (64) 28.92 ± 0.20 (61) 27.96 ± 0.20 (55) 7.11 0.0001
Female 27.08 ± 0.25 (25) 25.79 ± 0.57 (17) 26.55 ± 0.24 (39) 96.59 ± 0.20 (55) 2.85 0.040
Minimum nasal
Male 22.52 ± 0.32 (30) 23.24 ± 0.23 (57) 23.77 ± 0.22 (61) 21.62 ± 0.23 (54) 16.43 <0.0001
Female 20.83 ± 0.25 (23) 21.89 ± 0.35 (17) 22.01 ± 0.34 (39) 20.81 ± 0.20 (53) 5.14 0.002
Cranial suture
Male 15.86 ± 0.75 (32) 11.69 ± 0.69 (60) 10.01 ± 0.67 (63) 14.85 ± 0.65 (56) 14.38 <0.0001
Female 20.88 ± 0.62 (23) 17.94 ± 1.40 (15) 17.67 ± 0.68 (39) 18.47 ± 0.58 (54) 2.76 0.045
Pm3-M1 (mandible)
Male 21.22 ± 0.14 (32) 21.78 ± 0.14 (53) 23.85 ± 0.14 (61) 23.37 ± 0.15 (54) 65.60 <0.0001
Female 20.33 ± 0.14 (23) 20.11 ± 0.28 (15) 22.55 ± 0.15 (38) 22.43 ± 0.15 (54) 45.59 <0.0001
Variable
Mean ± Standard Error (Number examined)
FPUK Europe Central Asia Southern Africa
Statistically significant differences were detected by ANOVA (d.f. = 3 for all tests). Measured variables in mm unless stated.*Measurement was taken by N.Y. only
and the coefficient of variation was 0.36% and d.f. = 2. **Measurement was taken by N.Y. only and the coefficient of variation was 0.71% and d.f. = 2. Additional
derived variables were calculated by dividing the six measured variables with the highest F values by the greatest length of the skull.
APPENDIX 3 Continued
WILDCAT EVOLUTION AND CONSERVATION 63
© 2004 The Linnean Society of London, Biological Journal of the Linnean Society, 2004, 83, 47 – 63
Ramus (mandible)
Male 30.60 ± 0.40 (32) 29.36 ± 0.33 (51) 31.01 ± 0.43 (60) 30.03 ± 0.30 (55) 3.89 0.010
Female 27.38 ± 0.40 (23) 25.57 ± 0.42 (15) 27.19 ± 0.32 (38) 27.77 ± 0.24 (54) 5.63 0.001
WMC (mandible)
Male 14.85 ± 0.15 (29) 15.11 ± 0.19 (53) 16.40 ± 0.17 (60) 15.43 ± 0.20 (55) 13.23 <0.0001
Female 13.57 ± 0.18 (20) 13.40 ± 0.23 (15) 14.65 ± 0.21 (38) 13.91 ± 0.14 (55) 7.44 0.0001
Pm4 breadth (mandible)
Male 3.18 ± 0.04 (31) 3.19 ± 0.03 (52) 3.86 ± 0.03 (60) 3.53 ± 0.03 (54) 98.96 <0.0001
Female 3.03 ± 0.03 (23) 2.97 ± 0.04 (14) 3.68 ± 0.03 (38) 3.40 ± 0.03 (55) 70.21 <0.0001
Angular process (mandible)
Male 66.29 ± 0.59 (32) 66.13 ± 0.50 (54) 70.25 ± 0.48 (61) 67.22 ± 0.48 (55) 28.83 <0.0001
Female 61.49 ± 0.64 (22) 59.63 ± 0.72 (15) 63.03 ± 0.57 (38) 63.38 ± 0.40 (54) 21.94 <0.0001
Coronoid process (mandible)
Male 64.33 ± 0.41 (32) 66.19 ± 0.47 (50) 69.83 ± 0.41 (60) 68.86 ± 0.43 (55) 15.36 <0.0001
Female 60.12 ± 0.57 (23) 60.56 ± 0.56 (15) 63.94 ± 0.51 (38) 65.38 ± 0.43 (53) 6.80 0.0003
Cranial volume (ml)
Male 42.93 ± 0.47 (32) 44.60 ± 0.89 (33) 40.69 ± 0.35 (62) 41.45 ± 0.45 (73) 9.56 <0.0001
Female 41.06 ± 0.69 (21) 41.85 ± 1.43 (14) 36.75 ± 0.37 (38) 38.31 ± 0.43 (65) 11.89 <0.0001
*M1 length (mandible)
Male 8.07 ± 0.09 (19) 9.51 ± 0.08 (59) 9.15 ± 0.07 (53) 47.12 <0.0001
Female 7.55 ± 0.11 (14) 9.14 ± 0.08 (38) 8.81 ± 0.08 (55) 48.73 <0.0001
**M1 breadth (mandible)
Male 3.73 ± 0.05 (19) 4.12 ± 0.03 (59) 3.98 ± 0.04 (53) 15.82 <0.0001
Female 3.40 ± 0.05 (14) 3.94 ± 0.04 (38) 3.83 ± 0.04 (55) 24.07 <0.0001
Derived variables
Cranial index 2.31 ± 0.022 (45) 2.31 ± 0.046 (60) 2.61 ± 0.015 (101) 2.55 ± 0.019 (141) 33.92 <0.0001
I.O.F./snout 1.19 ± 0.005 (61) 1.12 ± 0.004 (101) 1.09 ± 0.004 (101) 1.09 ± 0.004 (112) 73.33 <0.0001
Palatal/Pm2-M11.85 ± 0.008 (58) 1.79 ± 0.010 (76) 1.74 ± 0.009 (89) 1.64 ± 0.007 (109) 113.44 <0.0001
POC/IOB 1.80 ± 0.019 (59) 1.75 ± 0.016 (92) 1.70 ± 0.016 (101) 1.89 ± 0.015 (113) 27.82 <0.0001
CP/AP 0.97 ± 0.003 (55) 1.01 ± 0.003 (79) 1.00 ± 0.003 (99) 1.03 ± 0.002 (111) 50.25 <0.0001
Derived variables (¥ 10-2: ratio to the greatest length of the skull)
Pm2-M122.42 ± 0.12 (51) 22.69 ± 0.12 (76) 23.49 ± 0.11 (91) 24.25 ± 0.07 (111) 64.68 <0.0001
Pm2-Pm421.39 ± 0.12 (51) 21.65 ± 0.11 (81) 22.25 ± 0.11 (92) 23.18 ± 0.08 (111) 61.49 <0.0001
Auditory bulla 21.10 ± 0.14 (52) 20.99 ± 0.10 (85) 21.46 ± 0.11 (102) 22.76 ± 0.10 (115) 65.34 <0.0001
Pm3-M1 (mandible) 21.60 ± 0.13 (52) 21.82 ± 0.12 (76) 23.02 ± 0.10 (100) 22.80 ± 0.09 (110) 39.50 <0.0001
Pm4 breadth (mandible) 3.24 ± 0.02 (52) 3.21 ± 0.02 (76) 3.74 ± 0.02 (99) 3.46 ± 0.02 (111) 102.40 <0.0001
*M1 length (mandible) 8.08 ± 0.08 (33) 9.24 ± 0.07 (98) 8.97 ± 0.05 (110) 45.81 <0.0001
Variable
Mean ± Standard Error (Number examined)
FPUK Europe Central Asia Southern Africa
Statistically significant differences were detected by ANOVA (d.f. = 3 for all tests). Measured variables in mm unless stated.*Measurement was taken by N.Y. only
and the coefficient of variation was 0.36% and d.f. = 2. **Measurement was taken by N.Y. only and the coefficient of variation was 0.71% and d.f. = 2. Additional
derived variables were calculated by dividing the six measured variables with the highest F values by the greatest length of the skull.
