A second extinct big cat from the Late Quaternary of Sri Lanka

Abstract

A second extinct big cat, tentatively considered to be a tiger (Panthera tigris), is recorded from Sri Lanka for the first time from a fossil left lower carnassial found in alluvium near Ratnapura in 1962 and a sub-fossil right middle phalanx 14 C dated to ~ 16,500 ybp, discovered in 1982 in a prehistoric midden at Batadomba Cave, near Kuruwita. The species is diagnosed from the only other big cats known from Sri Lanka, Panthera pardus and the extinct P. leo sinhaleyus Deraniyagala, 1938. This record significantly advances the timing of dispersal of tigers into the Indian peninsula. Tigers appear to have arrived in Sri Lanka during a pluvial period during which sea levels were depressed, evidently prior to the last glacial maximum ca. 20,000 years ago. The lion appears to have become extinct in Sri Lanka prior to the arrival of culturally modern humans, ca. 37,000 ybp.

Full text

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THE RAFFLES BULLETIN OF ZOOLOGY 2005
THE RAFFLES BULLETIN OF ZOOLOGY 2005 Supplement No. 12: 423–434
© National University of Singapore
A SECOND EXTINCT BIG CAT FROM THE LATE QUATERNARY OF SRI LANKA
Kelum Manamendra-Arachchi
Wildlife Heritage Trust of Sri Lanka, 95 Cotta Road, Colombo 8, Sri Lanka
Email: kelum@wht.org
Rohan Pethiyagoda
Wildlife Heritage Trust of Sri Lanka, 95 Cotta Road, Colombo 8, Sri Lanka (author for correspondence)
Email: rohan@wht.org
Rajith Dissanayake
23 Ranmoor Gardens, Harrow HA1 1UQ, United Kingdom
Email: rajd@nhm.ac.uk
Madhava Meegaskumbura
Wildlife Heritage Trust, 95 Cotta Road, Colombo 8, Sri Lanka
Department of Biology, Boston University, 5 Cummington Street, Boston, MA, 02215, USA
Email: madhava@bu.edu
ABSTRACT. – A second extinct big cat, tentatively considered to be a tiger (Panthera tigris), is recorded from
Sri Lanka for the first time from a fossil left lower carnassial found in alluvium near Ratnapura in 1962 and a
sub-fossil right middle phalanx 14C dated to ~ 16,500 ybp, discovered in 1982 in a prehistoric midden at
Batadomba Cave, near Kuruwita. The species is diagnosed from the only other big cats known from Sri Lanka,
Panthera pardus and the extinct P. leo sinhaleyus Deraniyagala, 1938. This record significantly advances the
timing of dispersal of tigers into the Indian peninsula. Tigers appear to have arrived in Sri Lanka during a pluvial
period during which sea levels were depressed, evidently prior to the last glacial maximum ca. 20,000 years ago.
The lion appears to have become extinct in Sri Lanka prior to the arrival of culturally modern humans, ca.
37,000 ybp.
KEY WORDS. – Sri Lanka, Pleistocene, tiger, lion, palaeontology, Panthera tigris.
INTRODUCTION
Apart from the leopard, which still persists in all natural
habitats across Sri Lanka, the only big cat recognised from
the island is an extinct lion, known only from two teeth found
in alluvial deposits at Kuruwita (06°47’N, 80°22’E) (Fig. 1).
Based on these, P. [E. P.] Deraniyagala (1939) erected a new
subspecies of lion, Panthera leo sinhaleyus, designating a
left lower carnassial (M1) as holotype (the other, a fragment
of a right lower canine, in too poor condition to facilitate
diagnosis, was lodged as a ‘metatype’ in NMSL: P. Deraniya-
gala, 1947).
The lion has been one of the most widespread of all non-
commensal mammals, having enjoyed a Pleistocene range that
included Africa, Eurasia, North America and tropical South
America (Nowak, 1999: 834). While the fossil record confirms
that the species’ range in the Indian subcontinent did extend
south to the 21st parallel and east to 87º E (Pilgrim, 1931;
Dutta, 1976)—approximately a line joining Gujurat to Bengal—
there is no evidence of the existence of the lion in Asia east of
Bengal or anywhere in peninsular India and Sri Lanka, except
for P. leo sinhaleyus.
The Holocene range of the tiger, however, extends to the
southernmost tip of peninsular India and to all of tropical
continental Asia (Hooijer, 1947; Aziz & de Vos, 1999). The
apparent absence of evidence of tigers in Sri Lanka and
Pleistocene peninsular India has led to the conclusion that
tigers arrived in south India “too late to get into Ceylon”

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Manamendra-Arachchi et al.: A second extinct big cat from Sri Lanka
(Pocock, 1930) as a result of the India-Sri Lanka land bridge
having been submerged since the Late Pleistocene. On the
basis of the few known Indian tiger fossils dating to the
Holocene (Lydekker, 1886a,b; Sankhala, 1978; Herrington,
1987; Turner & Antón, 1997; ) the recent literature too, dates
the arrival of tigers in the Indian peninsula only to the end of
the last glacial maximum, ca. 12,000 ybp (Hemmer, 1987;
Kitchener, 1999; Kitchener & Dugmore, 2000).
Recognizing that “Distinguishing apart the teeth of a tiger
from those of a lion is difficult”, P. Deraniyagala (1939)
distinguished the holotype M1 of Panthera leo sinhaleyus
from those of P. tigris entirely by its larger size, concluding
that “The Ceylon fossil, although narrower and more elongate,
agreed in general size with the lions’ teeth”.
In 1962, second a complete M1 was discovered in alluvium
in the course of excavating a gem pit near Ratnapura.
Tentatively identified as having come from a lion, it was
lodged in the National Museum’s Ratnapura branch. In
1982, a complete felid right-limb middle phalanx
(45.3×20.7×18.5 mm l×w×h) 14C dated to 13,500 ybp and
identified tentatively as belonging to a “large lion” (S. [U.]
Deraniyagala, 1992; subsequently revised to 16,500 ybp:
see S. Deraniyagala, 2001), together with undiagnosable
fragments of two other big-cat phalanges and an upper
premolar were found in a late Pleistocene midden in
Batadomba Cave (Fig. 2), at Kuruwita, a prehistoric human
habitation (11,500–37,000 ypb: S. Deraniyagala, 2004). This
phalanx clearly does not belong to a leopard.
Here we show that the 1962 Ratnapura M1 belongs to a tiger,
significantly advancing the timing of the dispersal of this
species into peninsular India. We also show that the
“Batadomba phalanx” belonged to a big cat more closely
related to the tiger than to the lion; and confirm from an
examination of its holotype M1 and a large series of recent
tiger and lion teeth that Panthera leo sinhaleyus was indeed
a lion, and that its demise appears to have preceded that of
the tiger in Sri Lanka.
MATERIALS AND METHODS
Materials referred to in this study are deposited in the National
Museum of Sri Lanka (Ratnapura) (NMSL); Department of
Archaeology, Colombo (DASL); the Field Museum of Natural
History, Chicago (FMNH); and The Natural History Museum,
London (BMNH).
Osteological terminology follows Turner & Antón (1997);
dental terminology follows de Muizon & Cifelli (2000). M1 =
lower carnassial (only those of the left dentary were used).
Measurements. – Metric measurements were made point-to-
point, using dial vernier callipers, to an accuracy of ±0.05 mm.
Angles were measured using a protractor, to the nearest 5°.
Carnassials—the following measurements were made: total
length (maximum anterior-posterior length of crown, Fig. 3a);
inter-apex length (distance between protoconid and hypoconid
Fig. 1. Map of Sri Lanka, showing location of Batadomba Cave,
Kuruwita (star), indicating also the climatic zones: DZ, dry zone;
IZ, intermediate zone; WZ, wet zone; the principal mountain ranges:
C, Central hills; K, Knuckles hills; and R, Rakwana hills; and the
500 m and 1,000 m contours. Scale bar: 50 km.

A
B
Fig. 2. (A), external and (B), internal views of Batadomba Cave,
Kuruwita, Sri Lanka, the source of the tiger phalanx DASL 1982.01.
DZ
IZ
WZ
K
C
R

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THE RAFFLES BULLETIN OF ZOOLOGY 2005
apices, Fig. 3b); protoconid length (length along superior
edge (blade) of protoconid, Fig. 3c); hypoconid length (length
along superior edge (blade) of hypoconid, Fig. 3d); protoconid
height (vertical distance from enamel margin to protoconid
apex, Fig. 3e); hypoconid height (vertical distance from enamel
margin to hypoconid apex, Fig. 3f); crown width (maximum
labial-lingual width of crown, Fig. 3g); crown depth (vertical
distance from enamel margin to posterior angle of hypoconid,
Fig. 3h); notch depth (vertical distance from base of median
notch to enamel base, Fig. 3i); carnassial angle (angle between
superior edges (blades) of protoconid and hypoconid).
Phalanges—the following measurements were made (see Fig.
3): distal width (maximum width of distal articulation, Fig.
3A); proximal width (maximum width of proximal articulation,
Fig. 3B); dorsal length (maximum dorsal length, Fig.
3C); ventral length (maximum ventral length, Fig.
3D); minimum width (minimum width of neck of phalanx, Fig.
3E); maximum height (maximum height of proximal articulation,
Fig. 3F); minimum height (minimum height of proximal
articulation, Fig. 3G).
SYSTAT for Windows XP, Version 11.00.01 was used for the
statistical analysis of dental and phalangeal measurements.
Principal components analysis (PCA) of the character
correlation matrix was used to reduce dimensionality of the
morphological variables and to identify those variables that
best discriminate between species. Various axis rotations were
tested and one selected for optimal interpretability of variation
among the characters. Discriminant function analysis (DFA)
was used to confirm the results that were obtained from the
PCA and to highlight the variables that best discriminate
between groups (lower carnassials and phalanges of recent
tigers, recent lions and the fossil examples). Both direct and
stepwise methods of discriminant analysis were employed.
TAXONOMY
Panthera leo
(Fig. 4; Table 1)
Panthera leo sinhaleyus P. Deraniyagala, 1939.
Material examined. – Holotype, lower left M1, BMNH Pal. Dept.
M 51883, 30.3 mm dorsal length; “Found immediately above gem
sand that was 19 ft below the surface in a Pit at Pan Vila, Edandé
Vala, Kuruwita, near milestone 51 on the Kuruwita to Ratnapura
Road, Sri Lanka” (06°43’ N, 80°23’ E, alt. ~ 30 m above sea level),
1936.
Identification. – (See Fig. 4A, B, D). The lower carnassial of
P. leo is distinguished from that of P. tigris most easily by
having only a single horizontal notch on the lower half of its
distal surface, see Fig. 4A, B, D (vs. two in P. tigris: see Fig.
4I). The carnassials of P. leo may also be distinguished from
those of P. tigris by having the talonid as a fairly well-
developed cusplet (vs. talonid elongate, swollen, in P. tigris);
the aboral protoconid slope convex in appearance (vs. the
smooth longitudinal ridge on aboral protoconid gives the
slope a flat or concave appearance in P. tigris); the saddle of
the talonid trough relatively short and shallow (relatively long
and deep in P. tigris); and the inferior enamel margin curved
Fig. 3. (Left), measurements of M1 (buccal aspect; see Materials and Methods): a, total length; b, inter-apex length; c, protoconid length; d,
hypoconid length; e, protoconid height; f, hypoconid height; g, crown width; h, crown depth; i, notch depth; and (right) middle phalanx (see
Materials and Methods): A, distal width; B, proximal width; C, dorsal length; D, ventral length; E, minimum width; F, maximum height; G,
minimum height.
b
cd
e
ih
f
a
g
A
B
CD
E
FG
DORSAL ASPECT LATERAL ASPECT VENTRAL ASPECT
PROXIMAL SAGITTAL ASPECT

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Manamendra-Arachchi et al.: A second extinct big cat from Sri Lanka
A
BF
E
10 mm
G
H
10 mm
Fig. 4. Left lower M1: A, lingual aspect, Panthera leo sinhaleyus, holotype, BMNH Pal. Dept. M 51883; B, buccal aspect, P. l. sinhaleyus,
holotype, BMNH Pal. Dept. M 51883; C, buccal aspect, P. pardus, NMSL uncat., Sri Lanka; D, buccal aspect, P. l. persica, female, BMNH
31.4.13.2, Gir Forest, India; E, lingual aspect, P. tigris, NMSL (Ratnapura), F559; F, buccal aspect, P. tigris, NMSL (Ratnapura), F559; G,
buccal aspect, P. tigris, NMSL (Ratnapura), F559; H, lingual aspect, P. tigris, NMSL (Ratnapura), F559; I, buccal aspect, P. t. tigris, BMNH
79.11.21.197, India. Arrows indicate notches on distal surface.
D
C
I
x
x
x
x
x
x
x
x
x
x
x

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THE RAFFLES BULLETIN OF ZOOLOGY 2005
downwards (vs. inferior enamel margin relatively straight in
P. tigris).
Measurements of holotype M1, BMNH M 51883 (in mm). Total
length, 30.3; inter-apex length, 19.9; protoconid length, 9.3;
hypoconid length, 14.2; protoconid height, 17.2; hypoconid
height, 15.0; crown width, 15.3; crown depth, 14.2; notch depth,
9.7; carnassial angle, 105º.
Note. – The Sri Lankan lion was allocated to a distinct subspecies
P. leo sinhaleyus by P. Deraniyagala (1939), but there is insufficient
information to determine how it might differ from other subspecies
of Panthera leo; for the purposes of the present paper therefore,
we consider this taxon as P. leo.
Panthera tigris
(Figs. 4, 5; Table 1)
Material examined. – Lower left M1, NMSL (Ratnapura) F559,
length 25.7 mm, Lindagava Kumbura, Muvagama, Ratnapura
(06°40’30” N, 80°24’12” E, alt. ~ 30 m above sea level), Sri Lanka,
20 May.1962. Complete sub-fossilised middle phalanx,
DASL1982.01, 45.3×20.7×18.5 mm (l×w×h) from Stratum 5; two
fragments of phalanges (DASL1982.02–03) from Stratum 4; and
one fragment of an upper premolar (DASL.1982.04) from Stratum
4, excavated from a midden in Batadomba Cave (see Fig. 2), Kuruwita
(06°47’ N, 80°23’E, alt. 460 m), Sri Lanka, 1982.
Identification. – (See Figs. 4, 5).The lower carnassial of P.
tigris is distinguished from that of P. leo most easily by having
two horizontal notches on the lower half of its distal surface
(vs. a single notch in P. leo). The carnassials of P. tigris may
also be distinguished from those of P. leo by having the talonid
elongate and swollen (vs. talonid a fairly well-developed
cusplet, in P. leo); the aboral protoconid flat or concave in
appearance as a result of the smooth, longitudinal ridge on
its buccal side (vs. aboral protoconid slope convex in
appearance in P. leo); the saddle of the talonid trough relatively
long and deep (vs. saddle of talonid trough relatively short
and shallow in P. leo); and inferior enamel margin relatively
straight (vs. inferior enamel margin curved downwards in P.
leo).
Principal components analysis (Fig. 6) with unrotated axes
on the correlation matrix of morphometric characters from
the phalanges of recent tigers, recent lions and the
Kuruwita right middle phalanx (DASL 1982.01) shows that
the fossilized example does not overlap with the recent
Fig. 5. A–C, dorsal, lateral and ventral aspects respectively, of right middle phalanx of Panthera pardus, NMSL uncat., Sri Lanka; D–F,
dorsal, lateral and ventral aspects respectively, of right middle phalanx of Panthera tigris, DASL 1982.01, Batadomba Cave, Kuruwita, Sri
Lanka. Scale bars = 10 mm.
A
B
D
E
F
C

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Manamendra-Arachchi et al.: A second extinct big cat from Sri Lanka
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Fig. 6. Plot of principal components 1 and 2 of seven metric variables of the Kuruwita right middle phalanx, DASL1982.01 (black triangle),
and available middle phalanges of pes and manus of two recent specimens of P. tigris (open magenta circles) and four recent specimens of
African and Asian P. leo (solid cyan circles) (for specimen details see Comparative Material).
tiger
lion
Table 1. Proportional measurements of lower left carnassials of Panthera tigris (Ratnapura M1, NMSL-R F559); ten recent P. tigris;
holotype of M1 Panthera leo sinhaleyus (BMNH M 51883); and 27 recent P. leo (details of recent specimens listed in Comparative Material).
tiger lion
NMSL-R F559 min. max. s.d. BMNH M 51883 min. max. s.d.
M1holotype M1
inter-apex length (% of crown length) 75.5 60.2 75.5 4.4 65.7 64.1 84.0 4.2
protoconid length (% of crown length) 40.9 30.7 40.9 3.0 30.7 30.2 40.4 2.8
hypoconid length (% of crown length) 54.9 44.0 54.9 3.3 46.9 41.5 58.3 4.2
protoconid height (% of crown length) 54.1 50.7 61.4 3.7 56.8 48.6 61.7 3.4
hypoconid height (% of crown length) 51.8 43.2 55.0 3.6 49.5 47.2 59.4 3.0
crown width (% of crown length) 51.0 47.7 59.4 3.0 50.5 47.7 55.6 1.9
crown depth (% of crown length) 47.5 41.3 53.8 14.1 46.9 38.3 51.7 3.7
notch depth (% of crown length) 31.9 28.3 38.2 2.5 32.0 23.4 34.9 2.6
inter-cusp angle (degrees) 105 90 115 8 105 100 120 5
protoconid length : hyperconid length (%) 74.5 64.2 74.8 3.6 65.5 63.4 80.0 4.0
crown depth : protoconid height (%) 87.8 76.7 96.4 6.0 82.6 72.4 99.3 7.1
crown depth : hypoconid height (%) 91.7 84.7 112.5 8.7 94.7 72.1 108.8 9.7
notch depth : crown depth (%) 67.2 60.0 73.8 4.2 68.3 57.3 76.2 5.9

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THE RAFFLES BULLETIN OF ZOOLOGY 2005
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factor (1)
factor (2)
Fig. 7. Canonical variables plot of discriminant function analysis of
available middle phalanges of pes and manus of two recent specimens
of P. tigris (magenta open circles), four recent specimens of African
and Asian P. leo (cyan open circles) and right middle phalanx from
Batadomba Cave, Kuruwita (solid black circle); 95% confidence
ellipses are centered on the centroid of each group. For specimen
details see Comparative Material.
tiger
lion
lions on the PC 1 axis. The PC 1 axis, which explains about
65% of the total variation, represents maximum height
(factor score 0.955), proximal width (factor score 0.948) and
distal width (factor score 0.936). These dimensions were
larger in the Sri Lankan fossil big cat than in recent lions.
The PC 2 axis, which explains about 17% of the total
variation, represents minimum width (factor score 0.722)
and dorsal length (factor score -0.520). The fossil overlaps
with—and cannot be distinguished from—recent lions on
the PC 2 axis.
Stepwise backward DFA (Fig. 7) shows that the fossilized
phalanx is distinct from but closer to recent tigers than
recent lions. This analysis correctly classified 100% of the
fossil example, together with 93% of recent lions and 91%
of recent tigers (Wilks’ lambda 0.2009; p = 0.000). The first
canonical variable best discriminates the groups and
accounts for 88% of total dispersion of the groups
(eigenvalue 2.542). The second canonical variable accounts
for 10% of the dispersion (eigenvalue 0.35). In the canonical
variables plot, the centroid for the fossil phalanx is (-3.727,
4.345), and those for recent lions (1.193, 0.063) and recent
tigers (-1.913, -0.299). The first canonical variable represents
mostly proximal width (standardized canonical discriminant
function (SCDF) -1.573) and minimum height (SCDF - 1.055).
The second canonical variable represents mostly maximum
height (SCDF -2.213) and minimum width (SCDF 2.164).
There is a slight overlap on the first canonical variable
between recent tigers and the fossil example, but no
overlap between this and recent tigers or lions on the
second canonical variable. Thus, according to the DFA,
some of the variables of the second canonical variable could
be used to distinguish the fossilized tiger from recent tigers
and recent lions: indeed, a review of the measurements
shows that the distal width of the fossil example does not
overlap with those of either recent lions or tigers. While
assigning this phalanx tentatively to a tiger, we consider it
possible that this may represent an as yet unknown species
of big cat.
Measurements of M1, NMSL (Ratnapura) F559 (in mm). Total
length, 25.7; inter-apex length, 19.4; protoconid length, 10.5;
hypoconid length, 14.1; protoconid height, 13.9; hypoconid
height, 13.3; crown width, 13.1; crown depth, 12.2; notch depth,
8.2; carnassial angle, 105º.
Measurements of right middle phalanx, DASL 1982.01 (in mm),
width of distal articulation, 20.1; width of proximal articulation,
20.7; dorsal length, 45.3; ventral length, 42.4; minimum
horizontal width, 11.3; outer height of proximal articulation,
18.5; inner height of proximal articulation, 12.5.
COMPARATIVE MATERIAL
Panthera tigris, phalanges: BMNH 1884.1.22.6, female, “Deccan”,
N=16; BMNH 114.K.K., male, “Mizapore, Decca[n]”, N=7; FMNH
31153, sex undetermined, “Allapalli, India”, N=15. Left lower
carnassials: BMNH 82.12.10.1, male, “Bengal”; BMNH 10.7.21.1,
female, “south-west India”; BMNH 29.11.2.1, sex undetermined,
“North Canara, Bombay” [sic]; BMNH 8.8.11.19, sex
undetermined, “North China”; BMNH 88.8.7.1, male,
“Afghanistan”; BMNH 13.5.11, sex undetermined, “West China”;
BMNH 1849.7.27.4, sex undetermined, “India”; BMNH
79.11.21.197, sex undetermined, “India”; BMNH 1938.8.12.3, male,
“India”; BMNH 1884.10.30.3, sex undetermined, “India”.
Panthera leo, phalanges: BMNH 68.657, female, Zambia, N=9;
BMNH 1932.6.6.4, male, “Tanganyeka”, N=8; BMNH
1952.11.13.1, female, “India”, N= 8; BMNH 1952.11.13.1, male,
“India”, N= 9; BMNH 1932.6.6.4, male, “Tanganyeka”, N=7;
FNMH 15530, sex undetermined, “Africa”, N=4; FMNH 73.175,
female, “Africa”, N=8. Left lower carnassials: BMNH 31.2.1.5,
female, “Bechuanaland”; BMNH 19.7.7.942, female, “Natal”;
BMNH 35.3.16.1, male, “Northwest Rhodesia”; BMNH 31.2.1.4,
male, “Bechuanaland”; BMNH 25.6.17.12, sex undetermined, “NE
Transvaal”; BMNH 30.12.3.1, male, “E. Transvaal”; BMNH
45.136, female, “India”; BMNH 34.11.1.4, male, “E. Transvaal”;
BMNH 25.6.17.5, sex undetermined, “NE Transvaal”; BMNH
25.6.17.7, sex undetermined, “NE Transvaal”; BMNH 25.6.17.9,
sex undetermined, “NE Transvaal”; BMNH 57.2.24.1, female,
“India”; BMNH 31.4.13.1, male, “India”; BMNH 31.1.5.1, male,
“India”; BMNH 31.1.5.2, male, “India”; BMNH 31.4.13.2, female,
“India”; BMNH 1642 a, sex undetermined, “Moshonaland”; BMNH
1893.5.20.1, male, Zimbabwe; BMNH 35.3.16.2, male, “Northwest
Rhodesia”; BMNH 34.11.1.3, female, “E. Transvaal”; BMNH
31.2.1.6, female, “Bechuanaland”; BMNH 93.5.21.1, sex
undetermined, “Bechuanaland”; BMNH 1893.5.201, sex
undetermined, “Zimbabwe”; BMNH 19.7.15.32, sex undetermined,
“S. Rhodesia”; BMNH 87.5.16.1, male, “south of Victoria Falls”;
BMNH 25.6.17.10, sex undetermined, “NE Transvaal”; BMNH
1992.167, male, Transvaal.
Panthera pardus, NMSL uncat., manus right 4th-digit middle
phalanx and left lower M1, sex undetermined, Sri Lanka.

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Manamendra-Arachchi et al.: A second extinct big cat from Sri Lanka
DISCUSSION
P. Deraniyagala (1939) did not explain explicitly how he
diagnosed the holotype M1 of Panthera leo sinhaleyus as
belonging to a lion, though he justified its allocation to a
distinct subspecies of lion by its being “narrower and more
elongate” than those of recent lions in the BMNH collection.
It appears that he based his species-identification essentially
on Brongersma (1935), confirming this through the
examination also of two recent tiger carnassials in the BMNH
collection. His conclusion that the BMNH M 51883 M1 belongs
to a lion and not a tiger was supported by Hemmer (1966b)
and also by our own examination.
We exclude the possibility of the Kuruwita-Ratnapura M1
and phalanx belonging to the only other Sri Lankan big cat,
the leopard (P. pardus), as the carnassials and phalanges of
leopards are immediately distinguishable from those of lions
and tigers in both morphology and size (Figs. 4, 5). The length
of the lower carnassial of the largest leopard we examined
was 18.4 mm, significantly short of even the smallest lion
(23.5 mm) and tiger (25.0 mm) measured. The largest middle
phalanx in the leopard, in the 3rd or 4th-digit of the pes, at
22.7 mm dorsal length, is significantly smaller than even the
smallest tiger (31.2 mm) and lion (22.9 mm) measured. Fossil
leopards are known from an archaeological context (see
below), and are not different in size from recent ones. We
note, however, that old museum collections of big cats may
be biased towards larger animals, which may have been
preferred by the sport hunters who acquired these specimens.
The phalanges of tigers and lions lack non-overlapping
proportional measurements, although they separate well in
multivariate space (see Fig. 6). At 45.3 mm dorsal length, the
Batadomba phalanx is significantly (9.3%) longer than the
longest recent P. leo phalanx measured in the BMNH
collection; these average 32.8 mm (s.d. = 4.4, range 22.9–41.1
mm, N=40) in dorsal length. The dorsal lengths of the
measured tiger phalanges averaged 37.4 mm (s.d. = 6.5, range
14.3–46.1, N=23). Unfortunately, we could find no way of
discriminating between the middle phalanges of the digits of
pes and manus, or determining the digit to which the
Batadomba phalanx belongs, though based on its large size,
we suspect it belongs to digit 3 or 4.
Given their close relationship (O’Brien et al., 1987), the
diagnosis of lions from tigers based on bone fragments alone
is challenging (see also Turner & Antón, 1997). While lion
and tiger carnassials do not separate clearly in mensural
statistics, Herrington (1987: fig. 4a) showed them to be distinct
in shape in occlusal view. We were, however, unable
unambiguously to distinguish lion and tiger carnassials using
principal components analysis and discriminant function
analysis based on the seven measurements made (see Fig.
3a–g, and Comparative Material for details of measured
examples). The left lower carnassials of recent lions, tigers
and the fossil examples used in this analysis cannot be
distinguished from each other with confidence by either PCA
or DFA. Principal components analysis served to demonstrate
that M1 F599, the holotype carnassial of P. l. sinhaleyus, and
those of recent lions and tigers show considerable overlap
on both PC axes. Recent lion and tiger carnassials could not
be unambiguously discriminated from each other in DFA,
which correctly classified only 81% of the recent lions and
80% of the recent tigers (Wilks’ lambda = 0.5093; p = 0.0293).
Inclusion of the fossil teeth together with the recent lion and
tiger carnassials gave a similar result. DFA correctly classified
100% of the holotype of P. l. sinhalayus and 100% of F599,
70% for recent tigers and 41% for the recent lions
(Wilks’lambda = 0.3046; p = 0.20). Here the standardized
canonical discriminant functions were greatest for inter-apex
length (0.953) and crown depth (-0.719). Owing to their
inconclusive outcomes, the dental PC and DFA plots are not
shown.
Hemmer (1966a: 23: trans. Colin Groves, in litt.) showed that
lion and tiger molars could be differentiated on the basis of
several subtle morphological character states, as follows.
Lion—“The talonid is a fairly well-developed cusplet, and
the aboral protoconid-slope usually appears convex; the
trough is narrow, the saddle of the notch is usually low and
the inferior enamel margin on the buccal side is curved
downwards.” Tiger—“The talonid is an elongated swelling
and the diminutive hypoconid is bordered by a notch, so that
the aboral edge of the protoconid appears mostly concave;
the protoconid is narrow, the trough is wide and the saddle of
the notch is usually high; the inferior enamel margin on the
buccal side is fairly straight.” We found these character states
to be consistent across all 27 lion and 10 tiger left lower
carnassials examined, but note that they are difficult to translate
into mensural data of statistical value using the measurement
techniques employed here.
The most reliable binary character state for distinguishing the
lower carnassials of tigers from lions is the horizontal notch on
the lower half of the posterior face of the tooth. Lions have one
such notch (see Fig. 4C), whereas tigers have two (see Fig. 4G):
see also Hemmer (1966a: pl. 7). The Ratnapura M1, however, has
the lower of these notches more prominent (Fig. 4H) than those
on the recent tiger teeth examined (see Fig. 4G), a character that
may have taxonomic significance. It is possible also that dental
characters have undergone change in these cats during the past
16 millennia (see Szuma, 2003, for evidence of dental variation in
the Red fox in the decadal time frame).
We cannot be certain that the Batadomba phalanx (DASL
1982.01) and the Ratnapura M1 (NMSL F559) belong to the
same species. While we are confident that the Ratnapura M1
belongs to a tiger, the Batadomba phalanx, while separating
distinctly from the lion, is clearly distinct also from the tiger.
At 45.3 mm dorsal length, this phalanx is less than 2% shorter
than the longest of the 23 BMNH tiger phalanges (46.1 mm,
from a female, BMNH 1884.1.22.6, from the “Deccan”
[peninsular India]), which suggests that this cat was
comparable in size to the tiger. Pending the availability of
further evidence, we choose to assign both the M1 and the
phalanx to a single species, tentatively the tiger. While
conceding that the large size of the Batadomba phalanx could
be the result of taphonomic bias, the size of the Sri Lankan cat
appears to have been remarkable given that insular P. tigris

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THE RAFFLES BULLETIN OF ZOOLOGY 2005
populations (e.g. in the Sunda Islands) have generally been
noted to be smaller than their mainland counterparts (Luo et
al., 2004).
The assignment of the Batadomba phalanx to P. tigris adds
confidence to our diagnosis of this species from Sri Lanka,
especially since trade in tiger teeth has been implicated in
prehistoric records of the tiger in islands such as Borneo,
where there is no other evidence of their presence and no
apparent reason for their extirpation (Wilkinson & O’Reagan,
2003). Indeed, the presence of sharks’ teeth in the Batadomba
Cave middens (pers. obs.), more than 50 km from the sea,
indicates that teeth may have played a role in commerce even
in prehistoric Sri Lanka. However, the discovery of the
Ratnapura tiger M1 in alluvium, together with hippopotamus
and rhinoceros fossils, demonstrates that tigers did indeed
occur in the island.
Based on present-day submarine topography, a functional
land bridge between Sri Lanka and India requires a sea-level
lowering of only ~ 10 m. Sea levels were ~ 120 m below
present-day levels at the last glacial maximum ca. 20,000 ybp
(Siddall et al., 2003), thus facilitating a more than 80 km-wide
terrestrial connection. It appears likely that sea levels were
sufficiently depressed during the final ~ 200,000 years of the
Pleistocene to have supported a land connection between Sri
Lanka and India for all or most of that time (Bossuyt et al.,
2004), and probably until 5,000–10,000 ybp (S. Deraniyagala,
1992; Anderson, 1998; Yokoyama et al., 2000).
While the present data push back the date of arrival of tigers
in peninsular India, they do not facilitate a conclusion as to
when these big cats first arrived in the peninsula. Tigers are
known from Java around two million ybp (Hemmer, 1987), and
may have reached India and Sri Lanka at any time during the
Pleistocene or Late Pliocene, though the lack of fossil
evidence does not permit a conclusive resolution of this
question. Indeed, there is no known barrier to the dispersion
of tigers into the Indian peninsula during the Pleistocene,
though Kitchener & Dugmore (2000) speculated that the
widespread presence of short grasslands may have resulted
in the tiger being altogether absent, or present only in very
small numbers, during this period, surviving successive glacial
maxima in refugia such as the moist forests of the south-
western Western Ghats mountains. The wet zone of Sri Lanka
may have provided another such refugium.
It appears however, that despite the existence of a land bridge,
an ecological impediment to the dispersion of moist-forest
faunas between the mainland and Sri Lanka did exist for much
of the past 500,000 years (Bossuyt et al., 2004), though the
nature of this barrier is not known. Although the climatic
history of South Asia is not well documented, there is evidence
that the climates of peninsular India and Sri Lanka experienced
protracted desiccation during Pleistocene glacial maxima (S.
Deraniyagala, 1992; Pant & Rupa Kumar, 1997), possibly
resulting in desertification of the land bridge between India
and Sri Lanka for much of that time. Even during the present
relatively pluvial period, southern India and northern Sri Lanka
are remarkably dry, precipitation being seasonal and rarely
exceeding 1,500 mm yr-1, with a vegetation of tropical dry
shrub-land, a habitat not associated with tigers. In view of
tigers having appeared in Sri Lanka, established a population
sufficient to have justified hunting, and then become extinct
at the end of the last glacial maximum, we suspect that their
entry to Sri Lanka (and therefore peninsular India) may have
coincided with a pluvial phase during or prior to the previous
interglacial, ca. 70,000–2000,000 ybp, their apparent absence
from Pleistocene India during this period being a sampling
artefact.
There is no fossil evidence in Sri Lanka that facilitates dating
of the appearance on the island of leopards, which arrived in
Asia 170,000–300,000 ybp (Uphyrkina et al., 2001; Meijaard,
2004). The leopard is known, however, from cave middens,
including those at Batadomba Cave, 14C dated to 31,000 ybp
(S. Deraniyagala, 1992), suggesting that it co-existed with the
tiger in Sri Lanka for several thousand years before the latter
disappeared. Leopards are also known from the Pleistocene
of India, from Billa Surgam cave in Karnul (Andhra Pradesh
State) and alluvial deposits at Susunia (West Bengal State)
(Saha et al., 1984). The case in Sri Lanka appears to have
been the reverse of the model proposed by Wilkinson &
O’Reagan (2003), which suggests that tigers were respons-
ible for the extirpation of leopards on Bali, if indeed leopards
did reach that island.
Despite the existence of a Pleistocene land bridge, there are no
records from Sri Lanka, either fossil or recent, of several present-
day south Indian large mammals, such as the wolf (Canis lupus),
nilgai (Boselaphus tragocamelus), four-horned antelope
(Tetracerus quadricornis) and blackbuck (Antilope cervicapra).
There is fossil evidence however, of the late Pleistocene presence
in the island of the dhole (Cuon javanicus) (P. Deraniyagala,
1958), which has since been extirpated. The gaur (Bos gaurus),
also known from middens at Batadomba Cave (S. Deraniyagala,
1992), appears to have persisted longer, becoming extinct only
in historical times (Knox, 1681: 78). In India and Southeast Asia,
the range of the tiger completely overlaps that of the gaur (Corbet
& Hill, 1992), the latter serving as a prey species for the former
(Lekagul & McNeely, 1988).
While tigers occur in a diversity of ‘closed’ habitats ranging
from tropical rainforests through mangrove swamps to tall
grasslands, lions are associated mainly with ‘open’ habitats
such as savannah, grassy plains and scrub (Nowak, 1999:
825, 832). The other fauna recorded from the same midden as
the Batadomba phalanx include the land snails Acavus and
Paludomus, the carp Tor khudree, the jungle fowl Gallus
lafayettii, gaur, and a variety of smaller mammals such as
monkeys and porcupines (S. Deraniyagala, 1992: 314; pers.
obs.). While many of these species occur in all Sri Lankan
forest types, members of the endemic Sri Lankan mollusc
genus Acavus are restricted to closed-canopy monsoon or
‘rain’ forest (Hausdorf & Perera, 2000). The late Pleistocene
fauna of the Ratnapura area also included a now-extinct
hippopotamus, Hexaprotodon sinhaleyus and rhinoceroses,
Rhinoceros sinhaleyus and R. kagavena (see P. Deraniyagala,
1963; S. Deraniyagala, 1992). Teeth of R. sinhaleyus (= R.
sondaicus: see Laurie et al., 1983), from Adavatta, Lunugala

432
Manamendra-Arachchi et al.: A second extinct big cat from Sri Lanka
(Sri Lanka) have been thermoluminescence dated to 80,000 ±
20,000 ybp (S. Deraniyagala, 2004). Indeed, R. sondaicus, the
Javan rhinoceros, is a rainforest species (whereas the Indian
rhinoceros, R. unicornis, is typical of the floodplains
ecosystem of the terai). These data suggest that the late
Pleistocene habitat of Kuruwita comprised swampland and
moist, closed-canopy rain forest that seems to have persisted
until large-scale clearing commenced ca. 150 ybp: elsewhere
in the range of these species, such habitats are associated
closely with tigers, but not with lions.
Further, rhinoceros, hippopotamus and lion remains are not
represented in Sri Lankan cave middens: they seem to have
disappeared before the occupation of these caves by early
modern humans. (The records of lion remains in the Batadomba
Cave middens—see S. Deraniyagala, 1992—are erroneous:
the Batadomba phalanx, here attributed to a tiger, is the only
specimen in sufficiently intact condition as to facilitate
definitive identification). The lions therefore appear to have
been victims of the advancing rainforests and dense monsoon
forests that accompanied the pluvial phase that saw the
advent of the tiger in Sri Lanka. While the Kuruwita and
Ratnapura fossils show that lions and tigers were sympatric
in this area, however, there is no evidence to suggest they
were syntopic.
The Late Pleistocene is also significant because it was during
this time that the initial dispersion of modern humans occurred.
Although stone tools probably dating back to the Mid-
Pleistocene have been found (S. Deraniyagala, 1992), the
earliest direct evidence of modern humans in Sri Lanka dates
to ca. 37,000 ybp (S. Deraniyagala, 2004). Whether hunting
pressure was sufficient to extirpate tigers from the island,
however, is open to question: there is no direct evidence to
support or refute the idea that modern humans impacted
negatively on the fauna, resulting in ‘prehistoric overkill’ sensu
Martin (1984).
Extinctions on islands have generally been associated more
with predation and prolonged attrition (“sitzkrieg”) than with
environmental change (Barnosky et al., 2004; Guthrie, 2004).
While hunting may have impacted on the population of tigers
in Sri Lanka, habitat loss too, might have been an important
determinant. Premathilake & Risberg (2003) show from a study
of pollen that at Horton Plains, a present-day tropical montane
rainforest (2,100 m a.s.l., ~ 40 km distant from Kuruwita), that
a significantly cooler climate dominated 24,000 ybp, giving
way to grasslands 18,000 ybp, semi-deciduous seasonal forest
establishing itself about 14,000 ybp, with the final transform-
ation into rainforest taking place only about 9,000 years ago.
Ungulate prey of 5.3–63.8 animals km-2 are required to support
typical tiger densities of 3.2–16.8 100 km-2 (Karanth et al.,
2004; Karanth & Stith, 1999). Both reduced prey density and
shrinkage of dense forest (resulting from desiccation during
the last glacial maximum) may significantly have reduced the
range and population of the tiger in Sri Lanka, with human
predation accelerating its demise. Opposing phases of climate-
driven habitat flux appear to explain the disappearance of
both the lion and the tiger from Sri Lanka, leaving this territory
to the only truly generalist Asian big cat, the leopard. Even
today, leopards are ubiquitous in Sri Lanka, persisting in all
natural habitats and many anthropogenic ones, from sea level
to montane cloud forest at up to 2,400 m a.s.l.
P. Deraniyagala (1939) supported his assignment of the
Kuruwita M1 to a lion rather than a tiger in part because of the
“almost complete lack of a reference to the tiger in Ceylon’s
art, legend and folk lore when compared with the frequent
appearance of the lion in these fields…” (P. Deraniyagala,
1958: 87). The appearance of the lion in Sri Lanka’s art, legend
and folklore, however, cannot serve as evidence of the
presence of lions in the island during historical times. The
majority ethnic group of Sri Lanka, the Sinhalese, derive their
name from ‘sinha’, the Sanskrit for lion; the name of the island
being a corruption of “si[n]hala-dweepa”, which translates
as “island of the lion race”. This association with lions appears
to have originated from Indo-Aryan colonisers of the island
in the sixth century BCE. “The original home of the first Indo-
Aryan immigrants to Sri Lanka was probably north-west India
and the Indus region” (de Silva, 1981: 3). As it happens, north-
west India remains the only part of the subcontinent in which
lions have been reported during historical times. As P.
Deraniyagala (1958) himself points out, the lions in Sri Lankan
art are heavily stylised and were evidently the work of
sculptors who had not seen the living animal (see Fig. 8).
The frequency with which the ‘sinha’ root appears in Sri
Lankan place names cannot be used as evidence for the former
occurrence of lions there. Singapore, which derives from the
Sanskrit ‘sinha’ (= lion) and ‘pura’ (= city), and Singaraja (=
Fig. 8. The lion in Sri Lankan art. Stylized lions such as this one at
Anuradhapura (6th century, AD) are common in Sri Lanka—evidently
the work of sculptors who had not seen the living animal.

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THE RAFFLES BULLETIN OF ZOOLOGY 2005
lion king) in Bali are two prominent instances where the lion
has infiltrated the etymology of places in which its has never
existed, thanks to the spread of Indo-Aryan culture. That such
place names should be commonplace in Sri Lanka while being
almost entirely absent in peninsular India (where, too, they ought
then to have been present) is also evidence of an origin based
on culture and not the physical presence of lions.
ACKNOWLEDGEMENTS
We thank Colin Groves (Australian National University,
Canberra) and David Polly (Queen Mary, University of
London) for critical review that served significantly to improve
the manuscript; and are further grateful to the former for
helpful discussion and for translating the portion of Hemmer
(1966a) quoted in the text. Richard Sabin, Norman McLeod,
David Gower, J. J. Hooker, Daphne Hills and Andy Currant (all
of BMNH); and Bill Stanley and Robert F. Inger (FMNH)
provided access to material, hospitality and guidance during
visits to their institutions. We are grateful also to Siran U.
Deraniyagala, former Director General of Archaeology, Sri
Lanka, for inviting K. M.-A. to work on the identification of
fossil animals in the Department’s collection, for the loan of
specimens, and for helpful discussion.
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