Natural history of the leopard (Panthera pardus) in Peninsular Malaysia

Abstract

Modern leopards originated in Africa less than a million years ago and dispersed east through Central Asia, the Indian subcontinent up to the Far East and Southeast Asia, where it has a discontinuous distribution today. Scarce resources, intraguild competition and the eruption of Mt Toba may have led to extinction of leopards on Sumatra. In Peninsular Malaysia the range and population of leopards have decreased since the last century along with reduction in suitable forest habitat, to an estimated 500-700 individuals today. The fact that nearly all leopards on the peninsula are melanistic is an unusual phenomenon attributed to genetic drift and natural selection, the latter likely driven by advantage in concealment from tigers in a thickly forested environment. Melanism alone does not make leopards in Malaysia genetically distinct from other Indochinese leopards (Panthera pardus delacouri) to warrant classification as a different subspecies, but it does make them a globally unique subpopulation worthy of conservation and protection.

Full text

Natural history of the leopard (Panthera pardus)
in Peninsular Malaysia
S. Y. CHEW
Abstract : Modern leopards originated in Africa less than a million years ago and dispersed
east through Central Asia, the Indian subcontinent up to the Far East and Southeast Asia, where
it has a discontinuous distribution today. Scarce resources, intraguild competition and the
eruption of Mt Toba may have led to extinction of leopards on Sumatra. In Peninsular Malaysia
the range and population of leopards have decreased since the last century along with reduction
in suitable forest habitat, to an estimated 500-700 individuals today. The fact that nearly all
leopards on the peninsula are melanistic is an unusual phenomenon attributed to genetic drift
and natural selection, the latter likely driven by advantage in concealment from tigers in a
thickly forested environment. Melanism alone does not make leopards in Malaysia genetically
distinct from other Indochinese leopards (Panthera pardus delacouri) to warrant classification
as a different subspecies, but it does make them a globally unique subpopulation worthy of
conservation and protection.
Keywords: leopard, Panthera, pardus, delacouri, melanism, melanistic, Malaya, Malaysia,
panther, Felidae, Carnivora
“It exhibits in perfection that symmetry of form, that combination of grace, strength and agility
which distinguish its tribe. The panther is the embodiment of a beast of prey.”
– R.I. Pocock
INTRODUCTION
The leopard is the most widely distributed wild cat species, ranging from Africa, where it
used to occupy the entire continent but is nearly absent from North Africa today, through
Central Asia and the Indian subcontinent, including Sri Lanka, through most of East Asia up
to the Russian Far East. In Southeast Asia it is found from Myanmar to Thailand and Vietnam
on the mainland, down to Peninsular Malaysia* and finally on the island of Java. Leopards
in Malaysia belong to the subspecies P. pardus delacouri, which is distributed on mainland
Southeast Asia including Malaysia.
Though widely studied in Africa and India, there is less research on leopards in other
parts of Asia and hardly any carried out in Malaysia up till recent years. Leopards are
cryptic carnivores and are by nature scarce owing to their position as top predators. Adding
to this is the difficulty of conducting field studies in thick tropical rainforest compared to
other leopard habitats and lack of funding or interest compared to more charismatic species
like the tiger. Interestingly an overwhelming majority of individuals documented in the
peninsula are of the black or melanistic variety (Kawanishi et al. 2010). It is this
characteristic of Malaysian leopards that this paper shall focus on, discussing the various
theories explaining this unusual phenomenon and whether it is a sufficiently unique
attribute to make the black panthers here evolutionary distinct from other Southeast Asian
leopards. First, some background on the evolution of leopards and how they came to the
region, followed by a survey of literature on the natural history of leopards in Malaysia and
their ecology.
Malayan Nature Journal 2019, 71(2), 127-137
127
Lee Kong Chian Natural History Museum, Singapore (c/o)
chewsyian@gmail.com
*As there are no leopards on Borneo, ‘Malaysia’ would henceforth refer to West Malaysia excluding
Sarawak and Sabah.

Origin and evolution
Leopards originated in Africa, diverging from a common ancestor with lions in the Panthera
lineage 2 to 3 million years ago during the Pleistocene. Subsequently, these early forms
were found in Europe, India and Java (Turner and Anton 1997). However genetic analysis
posits a second radiation of the modern leopard out of Africa between 470,000 to 825,000
years ago. The route taken out of Africa into Europe and Asia largely parallels that of
Hominids, suggesting that the history of leopards is closely tied to our ancestors, like a
shadow on the fringes of campfires as they made the same journey out of Africa (Uphyrkina
et al. 2001). This long prehistorical association with humans perhaps explains why leopards
are able to co-exist so successfully with us compared to other large carnivores, as they
became behaviourally adapted to our ways, first as predator and competitor for common
prey animals but eventually as subordinates as our numbers grew.
Discontinuous distribution in Southeast Asia
By two to three hundred thousand years ago, leopards have spread throughout the Asian
continent up to the Pacific Ocean in the east and the island of Java to the south, with the
associated populations differentiating into the current array of eight subspecies (Kitchener
et al. 2017). During the Pleistocene ice ages the Sunda shelf was exposed when sea level
was over a hundred metres lower than present day, thereby connecting Malaysia, Sumatra,
Borneo and Java by land. Though leopards used to occur on Sumatra (Whitten 2000), there
is no evidence of them ever being on Borneo despite the seemingly similar habitat found on
all the Sunda islands. As sea level rose around 14,000-10,000 years ago after the most recent
glacial maximum, these land masses were cut off as connecting land bridges disappeared,
leaving the Javan leopard (P. pardus melas) an isolated subspecies today.
A leading hypothesis of why leopards could not persist on Sumatra and perhaps
Borneo, if they ever existed on the latter, is the scarcity of food resources combined with
competitive pressure from a rich guild of carnivores found in these parts. Comparative
studies of the biomass of potential prey species per unit area in different habitats found
unsurprisingly that tropical evergreen forests are significantly less productive for ungulates,
which comprise the majority of big cat prey. Where the more open woodland savannas and
deciduous forests of Africa and South Asia supported a few tons of prey per km2 on average,
rainforests in the Sunda region contained typically less than a ton, even as low as a few
hundred kilograms per km2 (Meijaard 2004). It is argued that smaller herbivores of the
rainforests, such as monkeys, squirrels and other rodents could not support a mid-sized
predator like the leopard in the long-term, while a host of smaller carnivores that are more
adapted to hunt smaller prey could subsist on them. Besides five smaller felids, ranging from
the clouded leopard to the flat-headed cat, that are found on Sumatra, there are also nine
Viverrid species like the Malay civet, banded palm civet and linsang that could be competitors
for smaller prey. Further, the presence of the Sumatran tiger (Panthera tigris sondaica), the
smallest tiger subspecies and one adapted to the rainforest environment likely posed a
significant threat, directly and indirectly through competitive displacement as they targeted
the same prey species like barking deer and wild boar (Volmer et al. 2017). However, the
same conditions exist more or less in Malaysia, which has the same habitat and a similar
complement of predator and prey species, including the tiger (Table 1). Notably the biomass
of primates was found to be much higher in Malaysia than Sumatra, 1,234kg/km2 vs.
727kg/km2 (Marsh and Wilson 1981),which could help support leopards given their greater
usage of this resource compared to tigers (see ‘Diet’ below).
Do leopards persist in Malaysia despite the suboptimal conditions due to the
peninsula being connected to mainland Southeast Asia and a replenishing source population
i.e. is Malaysia a population sink for leopards? Or have they become better adapted over
time to the environmental conditions here?
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Table 1. Number of predator and prey species in Sundaland (Cranbrook 1987)
Another compelling reason postulated for the extinction of leopards in Sumatra is
the eruption of Mount Toba over 70,000 years ago, which was the largest volcanic eruption
in the Quaternary (Volmer et al. 2017). This catastrophe would have had an impact not just
locally but globally due to the volume of ash and debris that would have altered the weather
worldwide for months. This could very well have been the final push that resulted in local
extinction of leopards in the vicinity. Rising seas would have prevented subsequent
recolonisation of what would become the island of Sumatra. Interestingly, early twentieth
century Malayan naturalists like F. N. Chasen and C. B. Kloss believed that leopards existed
on Sumatra at the time (Chasen 1940), albeit with no firm evidence. If so it is possible that
extinction was more recent on Sumatra.
Leopards on Java formed an isolated population for a sufficiently long period to
have become a subspecies distinct from those found on mainland Southeast Asia. Though
the Malayan peninsula was never entirely cut off from continental Asia by water, it is
connected to it via the Kra Isthmus, which is today a mere 44 km wide at its narrowest.
Could this tenuous land bridge be a significant bottleneck for leopards such that genetic
exchange would also have been compromised?
The Leopard in Malaya
“The most vicious killer in the jungle is the panther or black leopard.”
– Charles Shuttleworth
Distribution
“In Malaya uncommon but widespread throughout the forest of the mainland from lowlands
to hill tops.” – Lord Medway
Up till the middle of the twentieth century, when forests covered a larger extent of the
Malayan peninsula than it does today, leopards presumably roamed from the Thai border up
north to Singapore Island at the southern tip, including larger islands like Penang and
Langkawi (Harrison 1966). Now locally extinct on the islands, their range on mainland
129
Malaya
Sumatra
Borneo
Monkeys
5
5
8
Suids
2
2
1
Tragulids
2
2
2
Cervids
2
2
3
Bovids
2
1
1
Total
13
12
15
Canids
1
1
0
Mustelids – excl. otters
3
5
4
Viverrids
11
9
9
Herpestids
2
2
3
Felids
7
7
5
Total
24
24
21

Malaysia has been reduced, commensurate with the loss of forest cover from over 75% in
the 1950s to less than 50% by the 1990s (Aiken 1994). A recent survey of leopard
populations in the region depicted confirmed and potential leopard habitat in Malaysia,
which is mainly down the central mountain range and three major protected areas:
Belum-Temengor, Taman Negara and Endau Rompin (Figure 1). As a testament to the
leopard’s tenacity, they can still be found in small forest fragments surrounded by urban
development and plantations, like Krau Wildlife Reserve (600km2), Pasoh (24km2) and
Ayer Hitam Forest Reserves (12km2), located within or on the outskirts of the capital city
Kuala Lumpur (Rostro-Garcia et al. 2016).
Figure 1. Leopard habitat in Peninsular Malaysia (Rostro-Garcia et al. 2016)
Population
Using infrared illuminated black and white (IR) photographs taken by camera traps,
researchers were able for the first time to distinguish individual black leopards as their
unique pattern of rosettes showed up clearly under IR illumination. This then enabled a
population and density estimate using standard statistical software (Hedges et al. 2015).
Where previously only absence/presence and occupancy surveys could be carried out, this
ground breaking study conducted in a logged secondary forest on the East coast in
Terengganu resulted in a density estimate of 3 leopards per 100km2, a low figure compared
to studies done elsewhere but expected given the suboptimal habitat as already discussed.
Combining this number with the area of potential habitat, a crude estimate of total
population in Malaysia was given as 282-847 individuals (Rostro-Garcia et al. 2016). Given
the large range in estimates, more surveys in other parts of the country using the new
methodology are needed to gather baseline population data.
It is not known how many leopards there were in the past, but as a comparison, tigers
were said to number around 3,000 in the 1950s (Locke 1954) a far cry from the estimate of
300 today (IUCN 2015). A similar trend would put the number of leopards at 5,000 and
above seventy years ago. Of course, the two populations are not independent and tigers
affect leopard numbers as a competitor species, so it is possible that the latter could have
fared better as tiger population declined. This is seen in Ujung Kulon, west Java where there
are more leopards today than fifty years ago when Javan tigers still roamed there
(Boomgaard 2001).
The author would be inclined to lean towards the higher end of the current estimate
for leopard numbers in Malaysia, i.e. 500-700.
Description
“The black panther or melanistic leopard is very dark grey or brown, against which the
rosettes of black spots are just discernible. In Malaya, melanistic animals outnumber
conventionally coloured leopards.” – Lord Medway
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Source
Sex
TL (cm)
HB (cm)
Weight (kg)
Locke 1954
M
188
Tweedie and Harrison 1965
122
Harrison 1966
210
120
40
Medway 1983
185-215
ZRC#
F
192
116
ZRC
F
182
110
The Straits Times 29 Dec 1883*
187
Parr 2003^
180-220
100-120
#Zoological Reference Collection,
Singapore
*Singapore specimen shot
^for Thailand
For a species with such an extensive global range, leopards exhibit a wide range of size and
colouration depending on the habitat and subspecies (Stein and Hayssen 2013). There is also
a fairly large sexual dimorphism between males and females. With this in mind, a
‘typical’ leopard found in Malaysia seems to be small compared to those from other parts of
its range, with a Total Length (TL) of less than 2 metres and a Head and Body (HB)
measurement of 1.0 to 1.2 metres (Table 2). Admittedly the sample size is not large, but can
be compared to HB of over 1.2m and TL of over 2m for African and Indian leopards
(Sunquist and Sunquist 2002). This is in line with Bergmann’s rule, where body size is smaller
in warmer environments along the equator compared to cooler regions at higher latitudes.
Table 2. Size of Malayan leopards
Diet
Leopards are known for their adaptability, and this is reflected in their broad diet
encompassing over 100 species of mammals and birds. They eat animals ranging in size from
insects to large antelope like the Eland. The preferred or optimal prey size is in the 10-40kg
range, with an average of 25kg. This seems to provide the highest calorific return with the
least risk of injury, taking into account the general size of leopards (Hayward et al. 2006).
Though no systematic studies appear to have been done on diet of leopards in
Malaysia, naturalists listed known prey items as follows, some as evidenced by scat:
monkeys, mouse deer, wild pig, ground birds, binturong, goats and dogs. The last well
known as a favourite of leopards throughout their range (Sunquist and Sunquist 2002). This
is also the case in Malaysia, where a panther had accounted for 50 dogs before it was finally
shot by Arthur Locke.
“..by far its favourite food is dog and it has been known to kill and carry off dogs in the
estates in broad daylight.” – G.T.C. Metcalfe
Of note is the importance of primates as a food source for leopards in Malaysia in
the context of competition for scarce ungulate prey from tigers. This was seen in Taman
Negara where leopard numbers were higher in areas where macaques were more abundant
(Kawanishi 2002). Unlike in Africa, where prey is often taken up onto trees to keep them
away from larger predators and scavengers, this is less necessary in the more forested
terrain of Asia where thick foliage aids in reducing incidents of kleptoparasitism.
Interestingly, leopards are said to start feeding from the stomach area of their prey more
often than not, compared to the hind quarters for tigers (Locke 1954).
131

132
Activity pattern
The time of day when predators are most active are influenced by a number of factors,
typically coinciding with the time when their prey are active. For large cats that prey on
ungulates in the tropics, this usually means they are mainly crepuscular animals, as the latter
tend to feed more during the cooler times of day. Periods of activity as determined from
camera traps in two different forested areas in Malaysia indeed showed peaks during dawn
and dusk (Figure 2). However compared with the tiger, leopard activity was not as markedly
crepuscular, being more evenly spread, and generally more diurnal. This could be avoidance
behaviour and an adaptive response to a more dominant competitor, by reducing the
likelihood of confrontation while sharing the same space.
Taman Negara (Kawanishi and Sunquist 2004)
Jerangau, Terengganu (Azlan and Sharma 2006)
Figure 2. Activity pattern of leopard in Malaysia
Melanism in leopards
Melanism in brief is the occurrence of darker forms than what is typical in the same species.
In leopards it is a recessive trait manifested when there are two sets of the mutated ASIP
(Agouti Signalling Protein) gene present. It is thus possible even when both parents are of
the spotted variety when each has only one allele of the gene, but both pass on this recessive
allele to their offspring resulting in expression of the melanistic phenotype. There is also
some evidence that female black leopards have smaller litter sizes and therefore lower
fertility rates than spotted ones (Lekagul and McNeeley 1977). These factors point to a low
likelihood of the rate of melanism increasing and becoming widespread in any given
population unless it is beneficial and naturally selected for (Schneider et al. 2012). This
makes the concentration of black leopards in Malaysia a phenomenon that warrants further
investigation. This topic was extensively discussed and the subject of Kawanishi et al.
(2010), which collated the results of past surveys in the country evidencing the overwhelming
prevalence of black leopards in the peninsula. Here I summarize the theories and elaborate
on those I believe to be most likely explanations.

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Where and why leopards are black
The unique history and situation of Peninsular Malaysia in Sundaland has already been
discussed, and the existence of a bottleneck at the Kra Isthmus mentioned as a potential
inhibitor of genetic exchange for leopards with the rest of mainland Southeast Asia.
Kawanishi et al. (2010) conjectured that an isolated small population would have made it
possible for genetic drift alone to account for melanism becoming fixed over a period of
20,000-40,000 years.
Being darker also has selective advantages for leopards under particular
circumstances, and is associated with certain environmental conditions. A study mapping
the global distribution of melanistic leopards confirmed environmental variables like
precipitation and vegetative cover being main factors correlated with melanism in the
species (da Silva et al. 2017). Black leopards were found to occur almost exclusively in
moist evergreen forests with low variance in rainfall throughout the year. In South Asia this
is found in the Western Ghats, northeast India and the Lower Himalayan range in Bhutan.
In Southeast Asia it extends from western Thailand down the Tenasserim range into
Peninsular Malaysia (Figure 3). Such favourable habitat is also found in west Java, where
melanistic individuals of the Javan subspecies are more commonly found than on the rest of
the island (Boomgaard 2001).
Figure 3. Distribution of melanism in Asian leopards (in dark circles) (da Silva et al. 2017)
Darker pigmentation as a characteristic of birds and mammals living in humid
environments near the equator gave rise to Gloger’s rule. One reason was that darker
feathers and hair were found to be more resistant to bacteria, conferring greater resistance
to disease. Logically, being dark in an environment with low light levels such as under
closed canopy forests would also be advantageous for camouflage from predators and prey
alike. These reasons could have selected for melanistic leopards in Malaysia and other parts
of Asia where conditions are similar. As plausible as they may sound, it is curious to observe
that melanism is not more prevalent in other species living in humid rainforests given the
advantages it brings. Comparing just within the Felidae, there are more spotted and
patterned pelages among the various smaller cats that live in the humid tropics than pure
melanistic individuals. For jaguars, the only other large cat species where melanism also
occurs, black individuals are also found more often in moist forest but nowhere are they as

134
concentrated in number, even in the Amazon rainforest (da Silva 2017). Most glaringly,
black leopards are not recorded from the equatorial rainforests of Central and West Africa,
ideal habitat where such individuals would presumably be better concealed from prey
(da Silva et al. 2017). Better camouflage would be effective if vision is the primary sense
used by prey. Most ungulates rely more on an acute sense of smell and hearing, especially
so where sight is limited in a thick and gloomy forest. Being black, which is the most
extreme form of dark colouration, may thus not provide a significant benefit for stalking
prey in such an environment.
The most likely reason then, for the occurrence of melanism in leopards would be
the advantage in concealment from predators or socially dominant competitors. In Asia
concealment from tigers was a reason to explain melanistic leopards postulated in
Kawanishi (2002) and I believe one that best explains the phenomenon. This competitive
pressure is absent in African forests, where leopards are the top predator, not counting
humans, since lions inhabit only the plains and more open woodland savannas. The same
could be said of jaguars as well, being the top predator in the forests of South and Central
America.
Co-existence of tigers and leopards
Today, with the exception of Sri Lanka, Sumatra and Java after the 1980s when the larger
cat became extinct there, tigers and leopards are sympatric throughout Asia. Temporal
separation by avoiding activity in the same time period is one strategy already mentioned
adopted by leopards as a form of ecological niche partitioning. The other two key ways are
resource and habitat differentiation. In places where ungulate prey species are abundant in
terms of size range such as south India, where spotted deer, sambar deer and gaur are
present in large numbers, the two cat species favour prey of different sizes (Karanth and
Sunquist 1995). In the absence of different size prey, tigers would displace leopards by
targeting the same prey species, as happened in Nepal (Odden et al. 2010). Lastly habitat or
spatial partitioning is when two species utilize different habitat type in the same landscape
to reduce competition. This was also observed in Nepal (Seidensticker 1976) where in
Chitwan leopards favoured more open terrain such as grassland and forest edge compared
to tigers.
In Malaysia and other evergreen rainforests in Asia inhabited by both tigers and
leopards, prey and spatial partitioning are less effective due to low prey densities and
uniform habitat respectively, though for the former case leopards can subsist on more
unconventional prey like monkeys that are not feasible for tigers as seen in Taman Negara.
Having also seen the similar activity patterns for both cats in Malaysia, albeit with some
minor differences, temporal partitioning as a strategy also seems limited. A characteristic of
the evergreen rainforest habitat is the opportunity for better visual concealment amidst the
more three dimensional environment enabling sharing of the same area without confronta-
tion (Kawanishi 2002). To this end, melanistic leopards would have the edge in being better
hidden from tigers, which are primarily visual and aural hunters and possibly unable to
scent other animals even at close range.
The author contends that the leopard population on the peninsula have evolved to
become melanistic as this trait confers a significant selective advantage in enabling the
ecological separation with tigers in a moist evergreen rainforest environment. For this
reason when leopards were extant in Sumatra they would have been in all likelihood
melanistic given the very similar environmental factors favouring this trait. Although no
skins from the island have ever been preserved, intriguingly Sir Stamford Raffles
referenced black tigers or ‘Rimau Kumbang’ being numerous there (Pocock 1930).

135
Is the Malayan black panther evolutionarily distinct?
The Malayan peninsula, with its multiple episodes of faunal compression due to repeated
sea level rises constricting the already narrow Kra Isthmus would have created conditions
for differentiation both between and within species. Indeed, this is an important
biogeographical boundary for flora and fauna, dividing mainland Indochinese from Sundaic
mammal communities (Woodruff and Turner 2009). Did the relatively isolated leopard popu-
lation here give rise to significant intraspecific genetic variance? Analyses of gene samples
across a range of Southeast Asian wild felids found surprisingly that, unlike other cat species,
leopard DNA did not show significant variance between individuals from Malaysia and those
north of the isthmus (Luo et al. 2014). In contrast, there is evidence that tigers in Malaysia
are sufficiently divergent genetically from other Southeast Asian tigers (P. tigris corbetti) to
justify them being a different subspecies (P. tigris jacksoni), though superficially they are
indistinguishable.
It seems that black leopards in Malaysia are genetically similar to their counterparts
from the Southeast Asian mainland and the current subspecies classification as P. pardus
delacouri remains valid, despite the fixation of melanism in the population. The dramatic
appearance of the phenotype is merely superficial and was likely a rapid adaptive response
to local conditions as already elaborated. This also implies that there was never an impedance
of leopard gene flow across the isthmus, in contrast to the tiger that had resulted in a
different subspecies on the peninsula for the latter. In comparison, leopards seem to have
continued to move freely across the Indochinese-Sundaland boundary despite the historically
challenging conditions, demonstrating greater resilience. After spreading south into
Sundaland, their adaptability was evident in the plasticity of phenotype allowing for
melanism to be selected for as a competitive response to the presence of tigers in a thickly
forested habitat. The seeming lack of any bottleneck effect that would have resulted in an
isolated population favours natural selection over genetic drift as the primary driver of
leopard melanism in Malaysia, bolstered by the improbability of this recessively inherited
trait becoming fixed absent any adaptive benefit.
Implications for conservation
Unlike the tiger, it appears that black panthers in Malaysia are not genetically distinctive to
warrant classification as a different subspecies of leopard. Indeed, one could argue that
losses in their population could conceivably be replaced with Indochinese leopards from
elsewhere, which belong to the same subspecies. However it remains a fact that nowhere
else in the world is there such a concentration of black leopards, a phenomenon that arose
due to very specific conditions of habitat and community unique to this locality. This should
make the leopards of Malaysia worthy of conservation efforts because if ever they are lost,
it would be unlikely that the same circumstances can be replicated or recreated for
widespread melanism in a charismatic wild cat to recur.
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