Journal of Mammalogy, 89(6):1435–1446, 2008
SPECIES DISTINCTION AND EVOLUTIONARY
DIFFERENCES IN THE CLOUDED LEOPARD
(NEOFELIS NEBULOSA) AND DIARD’S
CLOUDED LEOPARD (NEOFELIS DIARDI)
PER CHRISTIANSEN*
Zoological Museum, Universitetsparken 15, 2100 Copenhagen Ø, Denmark
Based on examination of molecular data and pelage patterns, it has recently been suggested that the island
populations of the clouded leopard, traditionally considered a subspecies, may, in fact constitute a separate
species. In this paper, I demonstrate that the island populations deviate strongly from the mainland populations in
a large number of cranial, mandibular, and dental characters. The differences far exceed those that have been
documented for subspecies within other pantherine felids, and are congruent with a separate species, to which the
name Sundaland clouded leopard, Neofelis diardi, has been given, although the name Diard’s cat has priority
based on historical precedence. I suggest that the vernacular name Diard’s clouded leopard be adopted for
Neofelis diardi. In contrast, mainland populations diverge less from each other, and are congruent with 1 species
(Neofelis nebulosa) and 2 subspecies, the western (N. n. macrosceloides) and eastern (N. n. nebulosa) clouded
leopard. Neofelis deviates from other large felids in many aspects of craniodental morphology, and most likely
also in several behavioral aspects. Diard’s clouded leopard appears more derived with respects to saber-toothed
craniodental features than the clouded leopard, indicating that the former may have gone farther than the latter in
convergently evolving craniomandibular features traditionally considered characteristic of primitive sabertoothed felids.
Key words: clouded leopard, craniodental morphology, Diard’s clouded leopard, evolution, saber-toothed cats, Sunda
Islands, species
In 1821, Edward Griffith described a new species of felid as
Felis nebulosa, based on a captive specimen housed in a private
menagerie in London. However, it is evident that he never saw
the specimen except from a drawing, probably made by Major
Hamilton Smith or Mr. Landseer, because he believed it was
the size of a Bengal tiger (Griffith 1821; Griffith et al. 1827).
This became the type description of the clouded leopard, later
referre to its own genus Neofelis by Gray (1867). It is not
known with certainty from where the specimen came, but it
may have originated from Canton in southern China (Griffith
et al. 1827; Osgood 1935; Pocock 1939a; Wozencraft 1993).
Milne-Edwards (1868–1874:208) commented on the animal
‘‘Thou-pao’’ from northern China as being the same as
Griffith’s felid. In 1823, Cuvier described the species Felis
diardi, reportedly from Java, which, however, has no population of clouded leopards today, although they were present in
* Correspondent: [email protected]
Ó 2008 American Society of Mammalogists
www.mammalogy.org
1435
the Pleistocene and into the Neolithic (Hemmer and von
Koenigswald 1964). The specimen was probably from Sumatra
(Kitchener et al. 2006). Hodgson in Gray (1853:plate XXXVIII),
depicted Felis macrosceloides (spelled macroselloides) based
on a specimen from Nepal, but with no description or diagnosis, and Swinhoe (1862) described Leopardus brachyurus
from Taiwan. Subsequently, these were recognized as subspecies of the clouded leopard (Neofelis nebulosa) by most
authorities (e.g., Ellerman and Morrison-Scott 1951; Hemmer
1968; Nowell and Jackson 1996; Pocock 1939a; Weigel 1961).
The 4 traditionally recognized subspecies have an allopatric
or parapatric distribution. N. n. brachyura and N. n. diardi are
supposedly insular forms, whereas N. n. nebulosa and N. n.
macrosceloides have adjacent distributional areas on the mainland. However, it is not generally agreed where these various
subspecies are found. According to a recent study (BuckleyBeason et al. 2006), N. n. macrosceloides is distributed in
Bhutan, India (including Assam and Sikkim), Nepal, and
western Burma, and N. n. nebulosa is present in eastern Burma,
Cambodia, southern China, Laos, Malaysia, Thailand, and
Vietnam. Another study in the same journal (Kitchener et al.
2006) depicted N. n. nebulosa as being the only Burmese
1436
Vol. 89, No. 6
JOURNAL OF MAMMALOGY
FIG. 1.—Variables included in the analyses. Cranium viewed from a) direct lateral, b) dorsal, and c) ventral perspectives; d) mandible in direct
lateral perspective; e) P3 and f) P4 in palatal perspective; and g) p4 and h) m1 in direct lateral perspective. A1–3 indicates angular variables.
Variables 5 and 6 denote upper canine crown height (5) and anteroposterior alveolar diameter (6), respectively, and variables 62 and 63 denote
lower canine crown height (62) and anteroposterior alveolar diameter (63), respectively. For description of variables, see Table 1. Average values
are listed in Table 2. Measurements of p3 crown length, and length and height of protoconid are not shown, but are similar to measurements
for p4.
subspecies. According to Ellerman and Morrison-Scott (1951),
N. n. macrosceloides is found in Burma, India, and Nepal,
whereas N. n. nebulosa is found in southern China and
‘‘Indochina’’ (¼ Cambodia, Laos, and Vietnam). According to
Buckley-Beason et al. (2006) and Kitchener et al. (2006), N. n.
diardi is an island form present on Sumatra, Borneo, and the
Batu Islands, but Pocock (1939a), Weigel (1961), and Nowell
and Jackson (1996) also state its presence on the Malacca
Peninsula, where, according to the former authors, only N. n.
nebulosa is present.
Buckley-Beason et al. (2006) found that molecular evidence
suggested that N. n. diardi was sufficiently different from the
other subspecies to warrant the rank of species, a suggestion
that has subsequently been widely reported in the popular
press. Kitchener et al. (2006) found that pelage patterns also
varied, although not as markedly, but that N. diardi differed
from the mainland populations in the size, number and morphology of the ‘‘clouds,’’ as well as in ground color of the coat,
as had been pointed out previously (Lekagul and McNeely
1977; Weigel 1961). Buckley-Beason et al. (2006:2375) and
Kitchener et al. (2006) both urged for caution in accepting
N. diardi as a separate species without a morphological study
and diagnosis, the latter stating that ‘‘Ideally, skull measure-
ments would have been desirable. However, few skulls were
available . . .’’ (Kitchner et al. 2006:2382). In this study, I provide
comparisons of cranial, dental, and mandibular morphology
among insular and mainland populations of clouded leopards
from throughout their range, and discuss the implications of the
findings, in particular whether the reported differences are in
accord with N. diardi constituting a separate species.
MATERIALS AND METHODS
For the purpose of this study, 49 clouded leopard (sensu lato)
skulls and mandibles were studied, comprising 25 specimens of
N. nebulosa diardi/N. diardi (13 #; 12 $), 7 specimens of N. n.
macrosceloides (3 #; 4 $), and 17 specimens of N. n. nebulosa
(10 #; 7 $). The sample of N. n. diardi/N. diardi consisted of
8 specimens from Borneo (3 #; 5 $), and 17 from Sumatra
(10 #; 7 $). The sample of N. n. macrosceloides consisted of
3 specimens from Sikkim (1 #; 2 $), 3 specimens from India
(2 #; 1 $), and 1 female from Bhutan. The sample of N. n.
nebulosa consisted of 5 specimens from Thailand (1 #; 4 $), 6
specimens from Vietnam (4 #; 2 $), 5 specimens from China
(4 #; 1 $), and 1 male from ‘‘Indochina.’’ For comparative
purposes, a sample of 21 jaguars (Panthera onca), 18 leopards
December 2008
CHRISTIANSEN—SPECIES DISTINCTION IN CLOUDED LEOPARDS
TABLE 1.—Descriptive list of characters used in the analyses on
Neofelis species, in comparison with the jaguar (Panthera onca),
leopard (P. pardus), and tiger (P. tigris). For key to variables, see
Fig. 1. Angular variables (A1–3) are: 1, angle between narial aperture
and dorsal nasal profile; 2, angle between dental gumline and
postorbital process (‘‘facial angulation’’); and 3, angle between
pterygoid–palatine and occipital crest.
Variable no.
Variable 1
Variable 2
Variable
Variable
Variable
Variable
Variable
3
4
5
6
7
Variable
Variable
Variable
Variable
8
9
10
11
Variable 12
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
Variable 44
Variable 45
Variable 46
Variable 47
Variable 48
Variable 49
Description
Condylobasal length of skull from occipital condyles to
premaxilla tip.
Facial length from premaxilla tip to posterior end of maxilla
at gumline.
Posterior skull length from jaw cotyle to occipital condyles.
Anteroposterior length of sagittal crest.
Dorsoventral height of upper canine crown in straight line.
Anteroposterior width of upper canine crown at alveolus.
Height of narial aperture from ventral part of I3 alveolus to
ventral part of nasal.
Vertical height of muzzle, taken just posterior to canine.
Vertical height of face taken at P3-P4 interval.
Vertical height of skull taken at pterygoid.
Height of anterior part of zygomatic arch at jugalmaxilla
junction
Height of midpart of zygomatic arch immediately posterior
to postorbital jugal process.
Length of nasals.
Width across nasals at narial aperture.
Width across nasals at maxillafrontal suture.
Width across muzzle.
Width between orbital apertures.
Width across frontal postorbital processes.
Width of postorbital constriction.
Greatest width across braincase.
Length of hard palate.
Basicranial length to center of foramen magnum.
Width across upper incisor arcade.
Width of hard palate between upper canines.
Width of hard palate across center of P3.
Width of hard palate across center of P4.
Width of pterygoid palate.
Width across zygomatic arches.
Width across mastoid processes.
Width across occipital condyles.
Anteroposterior length of P3 crown.
Anteroposterior length of P3 metacone.
Anteroposterior length of P3 paracone.
Width of P3 crown across paracone.
Width of P3 crown across metacone.
Anteroposterior length of P4 crown
Anteroposterior length of P4 metastyle.
Anteroposterior length of P4 paracone.
Anteroposterior length of P4 parastyle.
Anteroposterior length of P4 protocone.
Width of P4 crown across protocone.
Width of P4 crown across paracone.
Anteroposterior length of mandible from mandibular
condyle to symphysis.
Inlever moment arm of temporalis muscle.
Inlever moment arm of masseter muscle.
Dorsoventral depth of horizontal mandibular ramus
posterior to m1.
Dorsoventral depth of horizontal mandibular ramus
at m1-p4.
Dorsoventral depth of horizontal mandibular ramus at p4-p3.
Dorsoventral depth of horizontal mandibular ramus
anterior to p3.
1437
TABLE 1.—Continued.
Variable no.
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
50
51
52
53
54
55
56
57
58
59
60
61
62
63
64
Description
Anteroposterior length of m1 crown.
Anteroposterior length of m1 protoconid.
Dorsoventral height of m1 protoconid.
Anteroposterior length of m1 paraconid.
Dorsoventral height of m1 paraconid.
Dorsoventral height of carnassial notch.
Anteroposterior length of p4 crown.
Anteroposterior length of p4 protoconid.
Dorsoventral height of p4 protoconid.
Anteroposterior length of p4 paraconid.
Anteroposterior length of p3 crown.
Anteroposterior length of p3 protoconid.
Dorsoventral height of p3 protoconid.
Dorsoventral height of lower canine crown in straight line.
Anteroposterior width of lower canine crown at alveolus.
(P. pardus), and 24 tigers (P. tigris) were included. The
material is housed at the American Museum of Natural History;
Natural History Museum, London (BMNH); Zoological
Museum, Copenhagen; Museum National d’Histoire Naturelle,
Paris; Naturhistoriska Riksmusset, Stockholm (NRM); National Museum of Natural History (Naturalis), Leiden (RMNH);
Naturmuseum Senckenberg, Frankfurt; Staatliches Museum für
Naturkunde, Stuttgart; Shanghai Science and Technology
Museum; Museum für Naturkunde, Berlin (ZMB); and the
author’s personal collection.
A total of 64 craniomandibular and dental measurements
and 3 angular variables were analyzed (Fig. 1; Tables 1 and 2).
The measurements were used to construct 136 ratio variables,
and subsequent analyses of variance and post hoc Tukey’s
honestly significant difference tests were carried out on arcsinetransformed ratios, because this restored normality to the data
(Sokal and Rohlf 1995). Angular variables were not included in
ratios, and were analyzed without transformation. Principal
component analysis and stepwise discriminant analysis were
performed on the measured variables. Principal component
analysis does not require predefined groups, and was used to
examine clustering among individual specimens. If such were
present, discriminant analysis was used to compute axes of
dissimilarity among the groups, because this approach is effective in analyzing separation among groups, emphasizing variation among groups relative to within groups by identifying
canonical axes (kiXi,) that are linear functions of the included
variables, where ki represents coefficients and Xi represents
variables (Sokal and Rohlf 1995). The multiple regression
derived from discriminant analysis yields the best least-squares
predictor of group assignment, and thus facilitates post hoc
assignments of individual specimens to the groups in classification analyses.
RESULTS
The Sunda Island populations of clouded leopards were
significantly different from the mainland populations in a large
number of anatomical ratios, indicating a substantial morphological difference, as evidenced by 76 of the computed 136
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JOURNAL OF MAMMALOGY
TABLE 2.—Average measurement values for the included groups of Neofelis nebulosa and N. diardi, along with standard deviations. All values
are in millimeters, except angular variables 1–3, which are in degrees. For key to variables, see Fig. 1 and Table 1. Abbreviations: N. d.: Neofelis
diardi; N. n. m.: Neofelis nebulosa macrosceloides; N. n. n.: Neofelis nebulosa nebulosa.
N. d. (#)
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
54
55
56
57
58
59
60
61
62
156.69
63.31
48.46
73.55
37.90
13.01
23.87
38.44
53.95
55.70
21.24
14.35
48.15
22.82
15.70
40.63
31.51
54.13
33.20
57.42
71.10
46.47
18.82
20.67
43.44
57.18
18.42
110.63
69.74
34.76
12.97
2.74
5.20
5.35
5.93
18.64
7.68
7.54
3.40
5.88
10.27
6.81
112.72
28.48
23.96
20.84
21.05
20.23
21.35
14.34
7.47
9.16
5.17
9.37
4.53
13.59
5.95
9.91
2.46
7.75
3.94
5.22
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
9.85
3.31
3.14
12.31
2.43
0.77
1.67
3.46
3.79
2.98
1.22
1.40
3.33
1.62
1.13
1.32
2.08
6.08
1.66
2.40
4.21
2.74
1.13
1.29
2.71
2.78
1.92
8.40
5.77
2.50
0.52
0.35
0.35
0.30
0.34
0.96
0.36
0.54
0.34
0.63
0.50
0.41
6.16
2.35
2.23
1.41
1.53
1.41
1.38
0.97
0.44
0.72
0.51
0.55
0.38
0.64
0.48
0.78
0.32
0.62
0.26
0.40
N. d. ($)
129.04
53.69
41.96
35.46
30.06
9.88
18.53
32.44
45.74
49.03
17.17
11.34
38.53
18.70
13.15
33.59
25.95
45.71
30.47
52.39
58.73
40.12
16.59
17.99
37.54
50.42
16.66
90.02
56.72
30.15
11.53
2.44
5.02
4.92
5.28
17.44
6.92
7.08
3.16
5.12
9.30
6.34
90.95
22.88
18.42
16.94
16.36
16.23
17.07
12.76
6.84
8.28
4.66
8.42
4.09
12.15
5.63
9.2
2.23
6.84
3.58
4.60
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
4.04
3.72
2.49
11.84
2.28
0.99
1.17
2.67
2.75
1.63
1.88
1.37
2.58
1.12
1.31
2.28
1.04
3.92
1.41
1.27
2.31
1.42
1.09
0.93
1.51
1.34
0.93
3.98
2.17
1.65
0.64
0.22
0.39
0.35
0.36
0.79
0.57
0.37
0.28
0.62
0.54
0.31
3.49
1.10
1.92
1.03
1.00
0.93
1.14
0.76
0.40
0.59
0.38
0.50
0.47
0.66
0.32
0.65
0.22
0.72
0.44
0.37
N. n. m. (#)
150.09
63.06
43.70
75.11
32.30
12.04
22.03
34.81
48.18
52.02
21.17
14.55
46.10
21.86
14.25
39.77
27.83
42.98
27.05
51.14
68.58
44.77
18.80
22.25
42.19
54.19
16.29
98.22
64.09
32.13
12.61
2.47
5.86
5.27
5.71
19.41
7.35
8.06
3.78
5.98
10.09
6.58
106.96
26.75
25.09
21.38
20.24
19.10
19.58
14.2
7.58
8.83
5.42
10.00
4.78
12.40
6.23
8.88
2.10
8.77
4.26
5.40
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
5.99
2.80
2.09
10.47
2.50
0.94
1.26
1.83
1.83
1.03
1.13
0.82
2.25
0.88
1.64
0.33
1.87
6.44
1.44
1.86
1.90
1.79
0.65
1.27
1.39
0.98
1.52
7.11
2.28
1.20
0.35
0.22
0.41
0.20
0.36
0.29
0.17
0.35
0.25
0.54
0.47
0.08
3.65
1.94
1.84
0.21
0.40
0.72
0.64
0.46
0.23
0.28
0.21
0.50
0.22
0.79
0.40
0.13
0.39
0.33
0.27
0.30
N. n. m. ($)
126.41
55.59
39.92
40.49
28.41
10.27
19.58
32.36
44.17
48.61
16.84
12.13
38.50
19.23
13.57
34.53
24.50
39.90
27.57
50.28
58.58
38.68
16.14
18.86
36.66
49.11
14.62
86.46
54.96
30.15
11.65
2.31
5.62
4.70
5.25
17.52
6.99
7.26
3.24
5.45
8.92
6.23
92.14
22.31
20.91
17.91
15.84
15.06
15.84
12.49
6.83
7.86
4.72
8.72
3.72
11.54
5.77
7.91
2.06
8.28
4.53
5.03
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
5.71
1.29
2.34
18.3
1.75
0.22
1.95
3.52
3.79
2.92
1.73
2.15
0.85
0.97
1.35
1.79
2.93
4.90
1.07
2.43
2.32
2.73
0.27
0.77
0.95
2.19
1.38
5.63
3.32
1.66
0.37
0.26
0.47
0.11
0.40
0.87
0.30
0.11
0.29
0.43
0.55
0.27
3.90
2.33
1.92
1.25
1.61
1.45
0.98
0.54
0.29
0.37
0.42
0.42
0.29
0.79
0.70
0.23
0.27
0.37
0.22
0.15
N. n. n. (#)
158.25
64.24
47.44
81.61
35.25
12.37
23.18
35.47
49.93
54.00
19.72
14.22
46.66
23.11
15.20
40.64
29.33
45.70
28.04
54.64
71.41
46.52
18.57
22.46
43.77
56.27
17.26
105.38
66.68
33.52
13.27
2.65
6.01
5.24
6.19
19.47
7.59
7.91
3.50
5.54
9.89
6.35
113.33
27.43
25.18
21.15
20.28
19.31
20.06
13.91
7.40
8.41
5.66
9.16
4.15
12.49
5.96
8.16
2.31
9.27
4.71
5.49
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
9.78
4.61
3.97
12.38
2.99
1.19
1.87
2.28
2.87
3.62
1.34
0.83
3.65
2.43
1.15
2.58
1.67
3.32
1.65
1.98
5.71
1.89
1.30
1.49
3.21
3.36
1.03
5.56
4.07
1.44
0.96
0.30
0.37
0.51
0.42
1.36
0.64
0.59
0.39
0.51
0.62
0.36
7.28
1.72
1.61
1.44
1.79
1.95
1.92
0.79
0.51
0.41
0.60
0.43
0.35
0.67
0.32
0.34
0.33
0.53
0.32
0.25
N. n. n. ($)
138.85
56.51
41.96
50.71
28.94
10.00
20.53
30.67
43.72
49.28
16.75
11.66
39.66
20.26
13.40
35.96
26.40
42.75
28.07
52.11
63.25
41.33
17.12
20.32
40.11
51.78
15.42
94.97
59.54
31.07
11.65
2.50
5.23
4.74
5.43
17.92
6.99
7.57
3.25
5.23
8.99
5.77
98.95
23.63
21.46
19.24
17.93
17.24
17.81
12.59
6.73
7.60
5.02
8.66
4.01
11.87
5.79
7.80
2.05
8.59
4.40
5.11
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
6
3.66
1.62
3.00
14.50
1.36
0.49
2.02
1.99
1.93
1.16
2.01
0.94
2.19
0.98
0.93
1.01
1.81
5.00
0.96
1.44
2.33
0.91
0.52
0.61
1.45
1.35
1.25
4.50
1.52
0.84
0.63
0.37
0.37
0.19
0.38
0.44
0.37
0.29
0.24
0.46
0.18
0.21
2.02
1.17
1.70
0.68
0.66
1.37
1.31
0.55
0.19
0.42
0.54
0.62
0.41
0.46
0.29
0.40
0.26
0.43
0.33
0.32
December 2008
CHRISTIANSEN—SPECIES DISTINCTION IN CLOUDED LEOPARDS
1439
TABLE 2.—Continued.
N. d. (#)
Variable 63
Variable 64
Angle 1
Angle 2
Angle 3
28.48
11.25
147.68
85.93
129.82
6
6
6
6
6
1.53
0.44
3.13
3.00
3.51
N. d. ($)
22.36
8.68
146.18
84.53
126.63
6
6
6
6
6
1.91
1.05
2.59
2.21
4.18
N. n. m. (#)
25.92
10.59
148.3
82.57
129.7
ratio variables being significantly (P , 0.05) different in the
Sunda Island populations when compared to N. n. nebulosa,
and 61 being significantly different when compared to N. n.
macrosceloides. A principal component analysis demonstrated
a large divergence of the Sunda Island specimens from the
mainland specimens (Fig. 2). The 1st principal component
explained ;58% of sample variation (Table 3), and was
principally correlated with condylobasal and facial skull length,
palate length and basicranial length, width of the skull across
the palate (but not pterygoid palate) and mastoid processes, and
the length and dorsoventral height of the mandible and lower
canine. The 1st principal component primarily reflected
specimen size, as indicated by the divergence of specimens
of males and females. The 2nd principal component explained
;9% of sample variation, and indicated a genuine morphological difference between the insular and mainland populations, but not between the 2 mainland subspecies. The 2nd
principal component was positively associated with width
between the upper canines, length of P3 and P4 paracone,
angular variable 2, and the size and proportions of p3, and to
a lesser extent the size of P4 and C1, and mandibular moment
arm of the masseter, and the length of m1 paraconid. The 2nd
principal component was negatively associated with postorbital, braincase, pterygoid palate, and in particular postorbital
constriction width, P4 width across the paracone, m1
protoconid height, p4 length and in particular protoconid
height, and to a lesser extent the height of the skull posterior to
C1 and at P3-P4. Wilting et al. (2007) suggested that populations from Borneo and Sumatra represented different subspecies. In my study, 14 of 136 computed ratio variables were
significantly (P , 0.05) different between the 2 populations,
lending some support to the notion of geographic isolation. The
traditionally recognized mainland subspecies N. n. nebulosa
and N. n. macrosceloides were significantly different in 13 of
the computed ratio variables. However, neither of the subspecies differences was evident in the principal component
analysis (Fig. 2).
The Sunda Island specimens were distinguished from the
mainland populations by a large number of proportional
differences pertaining to the skull, mandible, and dentition,
and despite clouded leopards (sensu lato) being distinctly
sexually dimorphic, as other pantherines (Gittleman and Van
Valkenburgh 1997; Mazak 2004; Mazak and Groves 2006;
Palmquist et al. 2007), these differences were most often independent of sex, and reflected genuine differences among the
populations (Fig. 3). Relative to mainland clouded leopards, the
Sunda Island populations have longer upper canines, narrower
palate between the upper canines but wider palate across the
6
6
6
6
6
1.75
0.54
1.85
1.17
6.06
N. n. m. ($)
22.96
9.61
145.7
81.98
125.93
6
6
6
6
6
0.67
0.33
3.51
2.39
3.32
N. n. n. (#)
28.37
10.94
147.05
86.29
134.91
6
6
6
6
6
2.89
0.65
2.56
2.85
4.57
N. n. n. ($)
24.18
9.65
148.54
86.56
134.13
6
6
6
6
6
0.85
0.64
2.33
3.18
4.08
pterygoid, greater postorbital and postorbital constriction
widths, thicker upper carnassials (P4) across the paracone,
different muscle invectors in the mandible, as indicated by
higher temporalis relative to masseter moment arms, differently
proportioned mandibular horizontal ramus, larger p4 and
smaller p3, and a large number of distinct proportional
differences in interdental ratios. The latter is unexpected for
subspecific population differences, because dental differences
in mammals are characters traditionally used to distinguish
species.
A discriminant analysis on the included variables in comparison with other pantherine species (Wilks’ lambda ¼
0.00001, F ¼ 423.817, P , 0.00001) indicated that the mainland populations group together, but that they group distinctly
distantly from the insular populations, which also cluster
closely together. F-matrix values (Table 4) indicated that the
differences between the insular and mainland populations (10–
20) were equal to differences among other pantherine species,
such as the clouded leopard (sensu lato) and the leopard, or the
leopard and jaguar. In contrast, the differences between the
mainland subspecies N. n. macrosceloides and N. n. nebulosa
were approximately 20–40 times less. This was corroborated
by a subsequent jackknifed classification analysis, where the
FIG. 2.—The first 2 principal components from an analysis on 64
measurement and 3 angular cranial, mandibular, and dental variables
in Neofelis. Symbols: u, Neofelis diardi (#); , N. diardi ($); n,
Neofelis nebulosa macrosceloides (#); , N. n. macrosceloides ($);
r, N. n. nebulosa (#); m, N. n. nebulosa ($). The letters inside the
symbols for N. diardi indicate specimens from Borneo (B) and
Sumatra (S).
1440
Vol. 89, No. 6
JOURNAL OF MAMMALOGY
TABLE 3.—Principal components from principal component
analysis on all included variables in Neofelis. For key to variables
see Fig. 1 and Table 1.
TABLE 3.—Continued.
Component loadings
1
2
3
4
5
0.345
0.653
0.935
0.872
0.265
0.432
0.354
0.805
0.458
0.061
0.079
0.054
0.040
0.458
0.090
0.146
0.046
0.014
0.048
0.007
0.391
0.127
0.041
0.003
0.227
0.407
0.502
0.294
0.023
0.228
0.027
0.132
0.136
0.321
0.008
Variance explained by components
1
2
38.664
5.979
3
3.262
4
2.071
5
2.016
3
4.869
4
3.091
5
3.009
Component loadings
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
Variable
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
25
26
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
53
54
55
56
57
58
59
60
1
2
3
4
5
0.961
0.923
0.811
0.865
0.901
0.888
0.879
0.739
0.787
0.889
0.771
0.774
0.876
0.899
0.755
0.926
0.859
0.440
0.341
0.692
0.962
0.932
0.865
0.787
0.916
0.930
0.688
0.903
0.930
0.842
0.885
0.503
0.427
0.763
0.774
0.759
0.775
0.610
0.504
0.579
0.813
0.620
0.957
0.848
0.831
0.909
0.890
0.934
0.939
0.800
0.741
0.620
0.525
0.620
0.516
0.727
0.403
0.379
0.529
0.494
0.118
0.162
0.098
0.217
0.276
0.094
0.020
0.337
0.385
0.174
0.147
0.014
0.013
0.144
0.088
0.149
0.243
0.423
0.757
0.371
0.122
0.035
0.054
0.492
0.134
0.043
0.384
0.123
0.071
0.117
0.150
0.023
0.587
0.131
0.168
0.334
0.059
0.452
0.184
0.003
0.209
0.439
0.124
0.118
0.299
0.247
0.027
0.064
0.116
0.096
0.093
0.408
0.375
0.041
0.223
0.392
0.154
0.718
0.239
0.782
0.170
0.068
0.076
0.153
0.108
0.017
0.248
0.164
0.191
0.182
0.014
0.190
0.095
0.022
0.285
0.004
0.251
0.376
0.089
0.139
0.066
0.125
0.009
0.141
0.177
0.124
0.055
0.310
0.170
0.044
0.126
0.043
0.271
0.225
0.055
0.280
0.135
0.284
0.324
0.307
0.287
0.437
0.203
0.187
0.219
0.033
0.058
0.042
0.004
0.458
0.368
0.460
0.304
0.424
0.381
0.256
0.426
0.339
0.061
0.039
0.081
0.112
0.183
0.122
0.059
0.215
0.043
0.305
0.262
0.090
0.196
0.323
0.007
0.134
0.194
0.045
0.068
0.124
0.227
0.246
0.077
0.041
0.048
0.033
0.147
0.139
0.236
0.076
0.144
0.160
0.039
0.059
0.349
0.163
0.010
0.087
0.079
0.003
0.352
0.204
0.020
0.143
0.036
0.066
0.127
0.040
0.017
0.079
0.095
0.023
0.194
0.089
0.330
0.062
0.188
0.223
0.212
0.088
0.161
0.039
0.003
0.025
0.090
0.167
0.044
0.100
0.084
0.051
0.018
0.146
0.300
0.134
0.018
0.111
0.062
0.013
0.019
0.049
0.090
0.022
0.020
0.015
0.076
0.019
0.108
0.054
0.005
0.012
0.025
0.019
0.177
0.408
0.168
0.214
0.019
0.327
0.235
0.115
0.292
0.099
0.204
0.142
0.066
0.170
0.132
0.137
0.035
0.037
0.033
0.111
0.091
0.225
0.100
0.508
0.487
0.024
0.034
0.050
0.567
0.084
Variable
Variable
Variable
Variable
Angle 1
Angle 2
Angle 3
61
62
63
64
Percent of total variance explained
1
57.708
2
8.924
insular population was identified with 100% certainty, as were
the jaguar, leopard, and tiger, but some overlap was present
between the mainland subspecies, corroborating the above.
Discriminant function 1 was primarily a size component,
clustering groups of similar overall size, but subsequent
functions were near size-independent, emphasizing proportional
differences among groups. A plot of discriminant analysis scores
2, 3, and 4 (Fig. 4) indicated a large distance between the insular
and mainland clouded leopards, equal to species differences
among pantherines in general, and that the mainland subspecies
group closely together.
DISCUSSION
In recent years, concerns have been raised about the dramatic
increase in the number of purported mammalian species, often
by elevation of allopatric populations, traditionally regarded as
subspecies, to the rank of full species, a term widely used as
a basal entity in other fields of science such as macroecology
and conservation (Agapow et al. 2004; Chaitra et al. 2004;
Helbig et al. 2002; Isaac et al. 2004; Meiri and Mace 2007).
Labeled taxonomic inflation, this has resulted in numerous
claims of multiple species from allopatric populations based on
vague criteria, often by using the phylogenetic concept stating
that species are designated by having at least 1 apomorphic
character. This is relevant for the current discussion of Neofelis,
and Meiri and Mace (2007) argued that more-robust evidence
should be advanced than simply differences, which are often
found in allopatric populations of the same species (e.g., see
Weigel [1961] for felid fur patterns). They argued that populations may be deemed as separate species if a genuine ecological
or evolutionary distinctiveness could be formulated, and
advocated using a comparative analysis of differences between
undisputed, sympatric species, in this case pantherine felids.
The latter has been done for Neofelis by Buckley-Beason et al.
(2006) and in this study. N. nebulosa and N. diardi also appear
to diverge in a number of unusual craniomandibular characters,
as noted below, indicating different evolutionary adaptations.
December 2008
CHRISTIANSEN—SPECIES DISTINCTION IN CLOUDED LEOPARDS
1441
FIG. 3.—Box plots of some population differences among clouded leopards; top) Neofelis diardi, center) Neofelis nebulosa macrosceloides, and
bottom) N. n. nebulosa. Box length indicates the length of the central 50% of values, with box hinges at the 1st and 3rd quartiles. The whiskers
indicate values falling within the inner (1.5 times hinge median) fences, and outliers between inner and outer (3 times hinge median) fences are
indicated with asterisks. a) Width across postorbital processes/condylobasal skull length; b) width between upper canines/condylobasal skull
length; c) width across pterygoid palate/condylobasal skull length; d) width of braincase/width of postorbital constriction; e) lateromedial width of
upper carnassial (P4) across paracone/anteroposterior length of P4 crown; f) height of upper canine (C1) crown/condylobasal skull length; g)
height of lower canine (c1) crown/height of upper canine (C1) crown; h) anteroposterior length of p3 crown/mandibular length; i) mandibular
moment arm of masseter muscle (MAM)/mandibular moment arm of temporalis muscle (MAT); j) dorsoventral height of lower carnassial (m1)
protoconid/dorsoventral height of m1 paraconid; k) anteroposterior length of p4 protoconid/dorsoventral height of p4 protoconid; and l)
anteroposterior length of p3 protoconid/dorsoventral height of p3 protoconid.
Cranial, mandibular, and dental morphology indicate that the
Sunda Island populations of clouded leopards are significantly
different from the mainland populations, but the taxonomic
status of the insular form is not readily solvable by
unambiguous methods, because the 2 forms evidently constitute sister taxa (see Hennig 1966). In relation to the above,
a possible solution may be found by comparison with other
pantherines that are divided into subspecies. The discriminant
analysis scores and F-matrix values reported in this study
indicate that the insular populations are as distinct from the
mainland populations as are other species of pantherine felids
from each other. Clearly, the morphological differences
between the insular and mainland populations of clouded
leopards far exceed those traditionally reported for subspecies
identification in other pantherines, for example, the tiger
(Hemmer 1967; Hooijer 1947; Mazák 1981, 1983; Pocock
1929; but see Kitchener 1999; Kitchener and Dugmore 2000),
leopard (Pocock 1930a, 1930b, 1932; Schmid 1940), jaguar
(Hemmer et al. 1971; Nelson and Goldman 1933; Pocock
1939b; but see Larson 1997), or lion (Hemmer 1974; Mazák
1970, 1975; Pocock 1930c; Todd 1965). Recent, more-detailed
studies of subspecies status in several pantherines, where
a comparison of molecular and morphological data can be
made, strongly corroborate the notion that the insular
populations of clouded leopards warrant the status of species.
Miththapala et al. (1996) and Uphyrkina et al. (2001) found
that molecular data supported the Javan leopard as a clearly
defined subspecies, thus constituting a separate evolutionarily
significant unit, which had split off from other leopards as long
ago as the mid- to late Pleistocene (800,000–300,000 years
ago). The craniometric analysis by Meijaard (2004) confirmed
this, implying a distinct taxonomic clade, as indicated by
distinction of groups in discriminant analysis. Several traditional subspecies of lions have been confirmed to constitute
separate evolutionarily significant units in molecular studies
(Burger and Hemmer 2006; Burger et al. 2004; Dubach et al.
2005), which is corroborated by cranial morphology (Christiansen 2008b). Several tiger subspecies have been found to be
clearly distinguishable and separate evolutionarily significant
units in molecular studies (Cracraft et al. 1998; Hendrickson et
1442
Vol. 89, No. 6
JOURNAL OF MAMMALOGY
TABLE 4.—Discriminant analysis significance matrix and post hoc jackknifed classification matrix on 67 variables in Neofelis, and the jaguar
(Panthera onca), leopard (P. pardus), and tiger (P. tigris).
N. diardi
N. nebulosa macrosceloides
N. n. nebulosa
P. onca
P. pardus
P. tigris
0.0
10.553
20.119
50.371
27.516
159.418
0.0
0.575
31.078
15.293
76.294
0.0
44.646
20.478
128.220
0.0
12.738
46.882
0.0
38.297
0.0
0
0
0
21
0
0
0
0
0
0
18
0
% correct
Between-groups F-matrix
N. diardi
N. n. macrosceloides
N. n. nebulosa
P. onca
P. pardus
P. tigris
Jackknifed classification matrix
N. diardi
N. n. macrosceloides
N. n. nebulosa
P. onca
P. pardus
P. tigris
25
0
0
0
0
0
0
6
3
0
0
0
al. 2000; Luo et al. 2004, 2006), which has been corroborated
by craniometric studies (Mazak and Groves 2006). Yet, in all
these cases, the reported distinctions in multivariate analyses on
craniometric data, even among groups that were identified with
100% certainty in classification analyses, and thus would
constitute phylogenetic species according to the phylogenetic
species concept (Cracraft 1997; Cracraft et al. 1998; Groves
2001), were far less than reported in the present study for
insular and mainland populations of clouded leopards.
Accordingly, the results of this study corroborate the recent
studies on molecular and pelage data, and are suggestive of full
FIG. 4.—Three-dimensional plot of discriminant function scores 2,
3, and 4 from a discriminant analysis on 64 measurement and 3
angular cranial, mandibular, and dental variables in Neofelis, and
jaguar (Panthera onca; n ¼ 21), leopard (P. pardus; n ¼ 18), and tiger
(P. tigris; n ¼ 24). Symbols: u, Neofelis diardi; n, Neofelis nebulosa
macrosceloides; , N. n. nebulosa; n, Panthera onca; r, Panthera
pardus; n, Panthera tigris.
0
1
14
0
0
0
0
0
0
0
0
24
100
86
82
100
100
100
species status for the insular populations of clouded leopards,
which have recently been given the name Sundaland clouded
leopard (Wilting et al. 2007:5). However, N. diardi already has
a vernacular name. Georges Cuvier (1823:437) originally
named ‘‘Felis diardi’’ in honor of French naturalist and
explorer Pierre Médard Diard. This scientific name resulted in
the vernacular name of Diard’s cat immediately being applied
to it (‘‘Chat Diard’’—Audouin 1823:495), and throughout the
19th century it was commonly known by this name (e.g.,
Ripley 1858:543), prior to it erroneously being regarded as
a junior synonym of the clouded leopard. The 1st description of
this animal by Griffith et al. (1827), disregarding Horsfield’s
(1825) description of the ‘‘Rimau-Dahan’’ of Sumatra, which
was not yet demonstrated to be the same animal, also confirms
this, because the authors write: ‘‘There appears, however, to be
in Java another wild species of the cat, much larger, and very
remarkable for the beautiful regularity of its spots, which our
author [Cuvier] names from M. Diard, its describer, Felis
diardi’’ (Griffith et al. 1827:484). Although no fast rules exist
for vernacular names (International Commission on Zoological
Nomenclature 2000), the original name should be given
priority, in particular because it correctly honors Mr. Diard,
who sent Cuvier the 1st skin and a drawing, as was originally
intended. In keeping with its close affinity to the clouded
leopard, I propose the vernacular name of Diard’s clouded
leopard for N. diardi.
The mainland population collectively represents another
species, the clouded leopard (N. nebulosa), and may conveniently be subdivided into 2 subspecies, as traditionally held,
N. n. brachyura in the eastern part of its range, and N. n.
macrosceloides in the western part, to which the names eastern
and western clouded leopard, respectively, may be applied.
Although morphologically much closer to each other than
either is to N. diardi, there are nonetheless distinct morphological differences between them, as noted above. The status of
N. n. brachyura is not addressed in this study. This subspecies
is considered very rare today (Rabinowitz 1988), or possibly
already extinct (Pei and Chiang 2004). Recent studies on
molecular (Buckley-Beason et al. 2006; Wilting et al. 2007)
December 2008
CHRISTIANSEN—SPECIES DISTINCTION IN CLOUDED LEOPARDS
and pelage (Kitchener et al. 2006) data found no support for
a distinct N. n. brachyura subspecies, however.
Examination of molecular data suggests reproductive isolation of N. diardi from mainland clouded leopards by
approximately 1.4–2.9 million years ago, comparable to
species isolation in other pantherines (Buckley-Beason et al.
2006; Wilting et al. 2007). The distinct morphological
differences reported in this study are congruent with a long
reproductive isolation. John Edward Gray was probably the 1st
to notice the unusual craniomandibular morphology of Neofelis
and its resemblance to that of extinct saber-toothed felids (‘‘The
Felis macrocelis has very long, rather compressed canine
teeth . . . . Its skull presents the nearest approach to . . . Felis
cultridens . . . and . . . F. megantherion and F. smilodon . . .’’
[Gray 1867:260]; and ‘‘Lower jaw truncated and high in
front . . . . Canine teeth . . . very long . . . . This skull most nearly
resembles that of the celebrated fossil Felis smilodon . . .’’
[Gray 1867:265]). This has subsequently been repeated by
numerous authors, for example, Prater (1965:70— ‘‘very
striking . . . is the enormous relative development of the upper
canine teeth, which present the nearest approach among living
cats to the great tusks of the extinct sabretoothed tiger’’),
Rabinowitz et al. (1987), and Werdelin (1983), who noted that
Neofelis also has large lower canines, indicating its feline
affinity.
However, craniomandibular morphology of the sabertoothed felids (Felidae: Machairodontinae) differed from that
of extant felids in many more characters than the development
of hypertrophied upper canines (Emerson and Radinsky 1980),
although recent studies have suggested that it may have been
enlargement of the upper canines that largely dictated the
evolution of skull morphology in those felids, whereas skull
morphology of extant felids is more size-dependent (Van
Valkenburgh and Slater 2007). In several studies, I have drawn
attention to the unusual craniomandibular traits of the clouded
leopard (sensu lato), which collectively imply convergent
resemblances with primitive saber-toothed cats, such as greater
posterior inclination of the facial relative to the basicranial part
of the skull, lowered jaw joint, smaller and posteriorly inclined
paroccipital process, large and somewhat bladelike upper
canines, proportionally shorter lower canines, and a so-called
‘‘verticalized’’ mandibular symphysis (Christiansen 2006, 2007,
2008a). Mandibular bending resistance patterns in clouded
leopards also diverge from those of other extant felids, and bear
some resemblance to those of the extinct saber-toothed cats
(Therrien 2005). Interestingly, N. diardi appears more derived
than N. nebulosa in several of those respects.
The ratio of C1 to condylobasal skull length (Fig. 3f) in N.
diardi (0.239 6 0.012 SD) is significantly greater (P , 0.001)
than in N. nebulosa (0.218 6 0.013), whereas c1 to C1 is much
lower (Fig. 3g; P , 0.001; 0.745 6 0.038, and 0.815 6 0.031,
respectively). Some specimens of N. diardi have enormous C1
relative to condylobasal skull length, such as BM1939.1656
(0.271), and several have C1 to condylobasal skull length ratios
of ;0.25 (e.g., BM1938.11.30.24, RMNH a, and ZMB22153);
in contrast, only a single specimen of N. nebulosa (NRM585711)
had a C1 to condylobasal skull length ratio of 0.241, and most
1443
specimens range between 0.205 and 0.220. Thus, the true,
tusklike canines often alluded to in the literature are present in
Diard’s clouded leopard. Additionally, this species usually has
more laterally compressed (‘‘bladelike’’) upper canines than the
clouded leopard (P. Christiansen, pers. obs.).
The ratio of the mandibular masseter inlever moment arm to
mandibular length in N. diardi (0.207 6 0.014 SD) is
significantly lower (P , 0.001) than in N. nebulosa (0.223
6 0.014 SD), whereas the mandibular temporalis inlever
moment arm is significantly higher (P ¼ 0.018; 0.252 6 0.012,
and 0.242 6 0.016, respectively), causing the mandibular
masseter inlever moment arm to mandibular temporalis inlever
moment arm ratio (Fig. 3i) in N. diardi (0.825 6 0.071) to be
significantly lower (P , 0.001) than in N. nebulosa (0.922 6
0.063). This indicates that N. diardi relies more on temporal
than masseter muscles for jaw adduction than does N.
nebulosa, which was probably also the case among sabertoothed cats, where the posterior part of the skull was typically
tall, and along with the reduced coronoid process provided
a longer inlever moment arm for the temporalis, but the
zygomatic space for the masseter was reduced (Christiansen
2006; Emerson and Radinsky 1980).
Neofelis diardi has a significantly (P ¼ 0.009) taller horizontal mandibular ramus anterior to p3 relative to mandibular
length (0.189 6 0.009) than N. nebulosa (0.177 6 0.010), and
a taller skull relative to condylobasal skull length than N. n.
nebulosa, but not N. n. macrosceloides (Appendix I). Many
saber-toothed cats also had tall skulls relative to skull length,
and all had a taller mandibular symphysis relative to the
dorsoventral depth of the horizontal ramus, in particular
derived forms, such as Megantereon or Smilodon, and
especially Homotherium. N. diardi has a larger p4 and a smaller
p3 than N. nebulosa, and in saber-toothed cats, the anterior
postcanine dentition was often markedly reduced, whereas the
posterior dentition was enlarged. Accordingly, N. diardi appears to have evolved farther toward convergent craniomandibular morphology with primitive saber-toothed cats than
mainland N. nebulosa, possibly because of differences in prey
profile or prey density (see Meijaard 2004; Seidensticker 1986).
Numbers of remaining populations of Diard’s clouded
leopard are currently unknown. A survey of Sabah, Borneo
(;76,000 km2), indicated that Diard’s clouded leopard was
probably only present in one-fourth of the province’s reserves
above 350 km2, which was set as the minimal size for a
sustainable (;50 individuals), long-term population, and that
only 4 of such reserves, collectively representing ;5% of
Sabah’s area, were adequately protected (Wilting et al. 2006).
Wilting et al. (2006) estimated a density of 8–17 individuals/
km2, and a tentative total population in Sabah of 1,500–3,200
individuals, of which only 275–585 were present in sufficiently
large reserves with adequate protection. However, these
numbers are fraught with substantial uncertainty (Gordon
et al. 2007).
Rabinowitz et al. (1987) reported that Diard’s clouded
leopard was still relatively common in Sarawak and Sabah
provinces, and that was only little affected by human activities.
They also found that Diard’s clouded leopard was no longer
1444
JOURNAL OF MAMMALOGY
hunted to any degree, and that there appeared to be little market
for skins or body parts. In contrast, Santiapillai and Ashby
(1988) reported that in Sumatra, 65–80% of the lowland forests
were degraded or destroyed by the beginning of the 1980s, and
highland forests also were severely affected, but it was
estimated that only around 15% of forest cover was destroyed.
They estimated that before 1900, when most of Sumatra was
covered by primary forest, Diard’s clouded leopard was present
on most of the island, but by 1988, there was direct evidence of
it from only about 3% of the island’s area. The unusual nature
of Neofelis, and the possibility of Diard’s clouded leopard
having evolved along slightly different lines than the clouded
leopard, corroborate previous studies (Buckley-Beason et al.
2006; Kitchener et al. 2006; Wilting et al. 2007) in underscoring an urgent need for enhanced protection of Diard’s
clouded leopard and the clouded leopard separately, because
continuing habitat degradation is regarded as the most serious
threat for future survival (Cat Specialist Group 2002). This
work was supported by the Carlsberg Foundations
ACKNOWLEDGMENTS
I am indebted to D. Hills, P. Jenkins, H. Turni, I. Mann, J. LesurGebremariam, F. Renault, J. Barreiro, K. Krohmann, D. Möricke,
O. Grönwall, and H. van Grouw for much valuable assistance and
hospitability during my visits to their respective institutions. Three
anonymous reviewers provided much constructive criticism of an
earlier draft of this manuscript. I am especially indebted to J. Mazak
for material and information on clouded leopards from China. This
work was supported by the Carlsberg foundation.
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Submitted 11 January 2008. Accepted 27 June 2008.
Associate Editor was Jesús E. Maldonado.