(THE SUBSPECIES AND EVOLUTIONARY DEBATE)
nterestingly, many wild cats frequent regions where there is water, as an example, those belonging to the panthera
group, although most avoid direct contact and swim only if necessary (such as crossing a river to reach a viable food source).-
On the other hand, ALC
subspecies, that live in environments where there are high concentrations of water,-
appear to exhibit a degree of adeptness
and can swim easily
(see accounts of Pocock, 1917; Cai, et al, 1989; Lekagal and McNeely, 1977; Gao, et al, 1987).-
This association points to a certain affinity and natural attraction to water,
in a sense, "aquatic ability."-
In our view, this is a form of specialization
, and depending on habitat, is probably important for survival.-
Still, after a review of the literature, we believe, since there are reports of ALC
on off-shore islands within their habitable range (Lu and Sheng, 1986; Santiapillas and Suprahman, 1985), such behavior suggests
, that small groups of ALC may have swam short distances to other islands and established viable populations. -
This possibility has been alluded to by Nowell and Jackson (1996), who also speculate, that in such environments, over an evolutionary time frame, they evolved
to the point of -
This particular change or transformation is referred to as SPECIATION
and is common to biological organisms. Nonetheless, such occurrence, regardless of its extent, may explain the presence of so many ALC subspecies. However, this premise is theoretical, and although plausible, is not conclusive; an introduction to this process can be found at Wikipedia, see the section on allopatric speciation
As an addendum, explanations of evolutionary events and genetic change, as they relate to habitation and species development in specific environments
, such as islands and enclosed inland "pocket" habitats
, thereof, is conjecture, and has not been clearly defined in the scientifiic literature. However, based on current phylogenetic data, it is reasonable to assume that such development is highly possible.
s one would expect, applying the principles of "speciation" -
to explain evolutionary events, developmental tendencies, and physical changes in "animalia" (such as the inherent genetic expression
for specific characters and traits), -
is both an interesting and a complex subject. After all, trying to extract and draw logical conclusions from information, that, in many instances, is based on theory and limited genetic data, is often perplexing (Baker, J. M., 2005; Rice, W. R. and Hostert, E. E., 1993). Nonetheless, aften an extensive review of the literature, and as it may apply to the ALC and its subspecies, the occurrence of such events are possible for the following reasons:
- ALC subspecies and other Asian SWC have been observed on
islands throughout areas of: Indonesia, the Philippines, and Japan.
- The Asian basin and surrounding regions comprise an immense area,
much of which, has not been fully explored.
- The geographic regions of southeastern Asia consist of unique habitats,
many of which are suitable for cats (i. e., as long as they have
an adequate fresh water supply and a sustainable food source).
lthough the Asian Leopard Cat has evolved into a number of subspecies
, very little is known about the -
"chain of events"-
or causative factors
that produced these cats. From a theoretical perspective, it is possible, that this particular species developed a genetic predisposition
, in a sense, a propensity and natural inclination for change. It is here, perhaps, that a readiness for
, as well as a heightened state
of transformation was quickly
reached, and, at some critical developmental point
, accelerated changes in outward physical appearance were expressed
(ergo, the beginning of subspecies development).-
Further, whether through speciation
),- chance occurrence
, or, as yet, some unexplained
change in its DNA, physical variability
began (but, at this particular juncture, the affects of environment
and other external forces probably influenced the outcome, as well). However, the aforementioned premise is purely conjecture, but when viewed within the broader context of phylogenetics, such developmental theories, have, nonetheless, contributed to the ongoing ALC subspecies debate
. An in depth discussion of this subject, is, of course, beyond the scope of the present writing, for this, the reader is referred to the literature. We do hope, however, that this short narrative, has, at least, increased your interest and awareness, and perhaps, in a broader philosophical sense, deepened your appreciation for the complexity and beauty observed in the "natural"-
ot surprisingly, as one investigates the greater body of research regarding ALC subspecies, it is evident that current data creates even more questions. Here, finding answers, and constructing models that clearly reflect evolutionary change
and physical differences
among these cats, continues to be elusive. Still, what variables
are known, and their preponderance "to effect" change, have only added fuel to the debate:- Why are there so many subspecies of the ALC (listed as ten by some sources), when there is only a limited genetic database to support the current classification? -
In this regard, it has not been shown
there is a similar
or uniform genetic footprint
across subspecies populations, nor, is there proof, there are specific genetic markers
that would, conclusively
, differentiate these cats. However, such information, that is, an accurate and descriptive genetic account, does not currently exist
Therefore, until such information is found, in a sense, until more pieces of the genetic puzzel can be put together, the subspecies controversy will continue.-
As reported by the IUCN Cat Specialist Group (1996-2007):
" very little progress has been made in re-defining species using modern
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--molecular analysis, including genetic analysis, and most classically described
--cat subspecies are not valid based on current genetic information."
you examine pictures of the ALC and its subspecies throughout this writing, you will notice that no two cats look alike
In our opinion and discounting genetic influences, such development is due, in part, to climate differences
, geographic separation
, and isolation
, especially in the southern basin
of southeastern Asia,
where unique habitats and island land masses frequent the region (as seen in areas of Borneo, Malaysia, Sumatra, and Jakarta, as examples).-
It is in such environments, and over an evolutionary time period, that variability
and physical fragmentation
has occurred across populations (with attributes, that, for the most part, are easily recognized. As examples, differences
in color, markings, and in some cases, overall size.-
See Heptner, V. H., 1971; Johns, A. D., 1989; Yu, J., and Wozencraft, 1993).
rom a phylogenetic perspective, the primary
mechanisms responsible for physical variability
among ALC subspecies remain unexplained.-
However, speciation events
,- random occurrences
, incremental genetic drift
, and geo-environmental forces, among others, have impacted developmental tendencies
Whether these changes are due more to genetics, or to what degree and extent environmental factors
have played in the outcome, have not been fully explained.-
In one sense, it is easy to assume that genetics are responsible, after all, biological -
structure is primarily genetically based, although the influence of environment may be more important then is currently believed.-
Nonetheless, in our view, genetic changes
, resulting in physical differences among these cats are intrinsically subtle
, and therefore, difficult to analyze
Therefore, until the scientific community clearly differentiates these factors and accurately defines their interaction
(s), some conclusions, as they relate to developmental variability
, remain fragmented, and more theory then fact.-
Fragmentation, of course, has complicated a definitive genetic picture and taxonomy
Because of this, some experts suggest, that until the ALC subspecies question(s) is resolved, individual species names
would be more appropriate or logical, rather then the current listing(s). However, when one reviews the literature on this subject, which, incidentally, is very limited, such "premise" (with its associative inferences and assumptions), is not
entirely supported by current evidence.-
So, until there is more definitive genetic information, the current taxonomy
is more then adequate for classification purposes.-
Nonetheless, since questions remain regarding evolutionary, genetic, and physical variability, the following represent points of inquiry
that the scientific wildcat community should consider in their quest to resolve some of the ALC subspecies controversy
- Should each ALC subspecies be considered a unique singular species based on regional differences or geographic variation?
- In the presence of limited genetic information, is the current subspecies listing valid?
- Has prolonged isolation, particularly, in remote areas or habitats, been long enough to influence or impact genetic change (see speciation), which in turn, has led to variability in physical appearance, and if this is true, has this been of sufficient magnitude to warrant subspecies recognition (Yu and Wozencraft, 1993, as presented in IUCN/SSC Cat Specialist Group web site, 1996; Rabor, 1986; Nowell and Jackson, 1996)?
- Is there enough genetic information to clearly differentiate relationships between subspecies?
- Should specific "genetic markers" for physical characteristics be the primary source, or starting point, to build an overall species/subspecies classification?
- Is there a significant genetic heritage or link to other species of "small wild cats" (primarily to those found in the geographic regions of southeastern Asia)?
t is worth repeating, that the debate, surrounding specific aspects of ALC taxonomy and its subspecies, can never be fully resolved until a concise genetic description
and geo-evolutionary history
is completed. In a sense, without
an accurate genetic map
as a guide, and in the absence of verifiable information
, the questions surrounding subspecies status and classification
will be difficult to answer with any degree of certainty, if at all
. As reported by the International Cat Specialist Group (1996-2006):
" With regard to subspecies, there is considerable debate on definition,
--and even whether the traditional taxonomic concept is valid in the light
--of contempory knowledge of population biology and genetics. It is generally
--agreed that too many subspecies of cats have been described in the past
--on the basis of very slim evidence. "
s an addendum, one could reasonably assume, it would be relatively easy to obtain additional data
that would explain, and hopefully, resolve, the issues, questions, and developmental events that have impacted the ALC?-
But, presently, no serious attempt has been made to gather such information. It is true, some "field research
" and radio-telemetry studies
have been conducted in the past (Rabinowitz, 1990; Izawa et al., 1991), but such investigations, although helpful and necessary, have identified and yielded only estimates
, movement/migration patterns
, population densities
, and behavioral characteristics
. In all fariness, however, gathering the most important data, which in this particular case, would be genetic information
, could prove to be difficult. Here, blood, tissue, and similar samples would need to be obtained and carefully examined using molecular/genetic techniques and instrumentation (not an easy undertaking). As well, one must also consider present world events
, such as terrorism
in some Asian countries; you would certainly not want to put field-research
personnel "in harms way."-
Likewise, there are unforeseen geo-political
, resource/finanical allocation
, and geo-topographical
constraints, which also complicate such ventures. Because of these factors, among others, we believe relevant genetic information will not
be forthcoming anytime soon.
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he scientific name for the ALC is prionailurus bengalensis
(Pocock, 1917), but the original name, felis bengalensis
(Kerr, 1792), is sometimes used. Such description is in the "binomial" form of "Genus + Species
In this "Latin
" classification system, developed by the 18th century Swedish botanist and physician, Carolus Linnaeus (see Linnaean taxonomy
), the organism is composed of its "genus" name, with the first letter always "capitalized,"-
followed by a "species
modifier" in small letters, and in some cases, if there are more then one "subspecies,"-
or geographic descriptor (this nomenclature still stands as the basis for most scientific classification and identification systems). Of course, such usage can be shortened with the letters "p" and "b" (e. g., "Prionailurus bengalensis chinensis," is the scientific name for the ALC subspecies
found in regions of China, and is abbreviated as "P. b. chinensis
this is sometimes referred to as trinomial
description). Not surprisingly, any particular Gensus
(its grouping), -
can have one or more SPECIES
. As an example, the Flat-headed Cat, the Fishing Cat, and the Asian Leopard Cat, although different species, all belong to the same genus, "Prionailurus" (i. e., "small wild cats" in regions of Asia).
any aspects of "taxonomy" and "classification"-
were addressed in a major update and review of "wild cats" by Wozencraft (1989, 1993). Concerning the ALC, enough phylogenetic information is present to clearly list this cat
in the genus
along with the Fishing Cat
(p. viverrinus), the Flat-Headed Cat
(p. planiceps), and the Rusty-Spotted Cat
(p. rabiginosus). Such placement, according to Wozencraft, not only defines its status
as the most common and widely distributed SWC, but, as well, reflects a close relationship
to felines of similar genetic background and geographic location.
s examples of ALC's in the classification debate, those of Java, Sumatra, and Borneo have been recognized as distinct subspecies
(p. b. sumatranus, p. b. borneoensis, p. b. javaenis), but not
the cats of the Philippine islands (p. b. minuta), which have not been adequately described ( see IUNC web site, Cat Specialist Group, 1996, descriptive and behavior section). Likewise, the Iriomote Cat, found only
on the island of the same name (on the southern most tip of the Ryuku island-chain), near Taiwan, is a unique species
(Imaizumi, 1967), although others consider it to be a subspecies
of the ALC (Masua et al., 1994; Suzuki et al., 1994). Similar questions exist for the Amur Leopard Cat, subspecies, p. b. euptilurus (found in the Amur river region of south eastern Russia), and the Tsushima Cat (found on Tsushima island, off of Japan). It has even been suggested that there may be as many as 15 ALC subspecies. However, there is currently not enough genetic information to support such claims (see articles and cat information in the IUCN Cat Specialist group news letters, 1990).-
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KINGDOM: animalia.- PHYLUM: chordata.-SUBPHYLUM: vertebrata.- CLASS: mammalia.
- ORDER: carnivora.
FAMILY: felidae.- GENUS: prionailurus. -SPECIES: bengalensis
(the subphylum category refers to all species with vertebra, that is, the fishes,
reptiles, birds, and mammals, but it is often not included in many taxonomic listings)
* * F.(P.) b. bengalensis ---- - The regions of India to Indo-china and Yunnan.
* The number of subspecies and their classification is much debated (THIS ISSUE HAS NOT BEEN RESOLVED AS OF 2008). Further, there is some question regarding the Iriomote Cat of Japan. Some feel it is a subspecies, while others consider it a rare and separate species. Until more supporting genetic data is found, it is, in our view, a genetic variant belonging to the genus, Prionailurus, and its status remains undetermined (although it is often listed in the literature as a subspecies of the ALC, as Prionailurus bengalensis iriomotensis). There are also similar questions regarding the Tsushima Cat and other sub-genera of Prionailurus.
----F.(P.) b. borneoensis ----- Borneo.
----F.(P.) b. chinensis--------- China and Taiwan region.
* * F.(P.) b. euptailura ------- Far eastern regions of Russia and Siberia.
----F.(P.) b. horsfieldi -------- Parts of Kashmir to Sikkim.
----F.(P.) b. javaenis ---------- Java and Bali.
----F.(P.) b. iriomotensis------ Iriomote-jima, Ryukyu island chain of southern Japan.
----F.(P.) b. manchurica ------ The region of Manchuria (the largest subspecies).
----F.(P.) b. minuta ---------- - The Philippines (the smallest subspecies).
----F.(P.) b. sumatranus------- Sumatra and outlying regions of the Indonesia basin.
----F.(P.) b. trevelyani------- - North Kashmir to South Baluchistan, Pakistan.
--The Abbreviation of subspecies, as presented in the scientific literature, is usually listed with a lettered prefix(s): -F. - felidae/felis (i. e, the cat family), -B. - bengalensis (species designation), -P. - prionailurus (its genus - as a small Asian wild cat); in most context, the F prefix is not needed. Such description, whether it is in binomial-or trinominal-form, is based on Latin (see Big Cats Online history and evolution section, which gives a short, but concise account of taxonomy).
* * It is noted, that p. b. euptailura, is listed as a subspecies of the ALC by some authorities. However, this is debated by wildlife biologists, and, to date, this issue has not been resolved by the scientific community. Nonetheless, this cat is endangered and is approaching extinction. The subspecies, p. b. bengalensis, of India and Indo-China regions is endangered. The current status of these two cats is according to the CITES listing (2008).
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ctual field observations describe the ALC as shy, reclusive, and nocturnal
, although there are documented reports of movement and foraging behavior during the daylight hours (Rabinowitz, 1990). They are intelligent, accomplished swimmers, agile climbers (arboreal in some habitats), and like all wild cats, are primarily carnivores
. Their overall size, at maturity, is approximately 5-8 pounds, as seen in the southern parts of their range, to as much as 18-20 pounds in the northern geographic regions, and males, not surprisingly, are larger then females (Izawa et al. 1991; Rabinowitz, 1990).- Ten subspecies
have been identified, each of which show variation in overall markings and body color, but as previously mentioned, this number
is controversial (See IUCN/SSC web site, 1996). The general concensus, however, suggests seven to ten subspecies. Additionally, our research indicates their physical and behavioral makeup is dependent
not only on the "genetic footprint" of the individual, but as importantly, by environmental influences (i. e., climatic conditions and the topographical features of the particular regions where they are found). Still, no two ALC have exactly the same look
, since there can be both subtle, and sometimes, striking differences in appearance (see pictures throughout this document). Such physical divergence and development is relative to, and influenced by: time-duration
, specific topographical habitat
, and more importantly, to genetic interactions
, which, to date, have not been clearly defined.
esearch and data reported in the literature suggest ALC's are very adaptable
, and can: (1) live in diverse
environments, (2) adapt
quickly and readily to sudden or abrupt changes
in habitat (in a sense, to what we define as environmental complexity
), and (3) easily adjust and supplement
their dietary needs in order to survive (since they can eat almost anything).
In other words, you can infer these cats are probably "survivors
" based solely on their adaptability, despite the fact that they are often hunted for their fur and as a food source in many countries (esp. in China and India). As an example of their ability to adapt to the presence of man, it has been reported that villagers, in some Asian countries, keep ALC and Leopard cat hybrids in their homes (as pets
, and to hunt/control rodent populations, such as mice and rats). So, although the ALC is classified as a wild cat
, it appears that under certain conditions or circumstances, they are trainable, and therefore, could be
considered good candidates for domestication
, whether it be as an adult, that is raised from a kitten, or through selective breeding programs that produce hybrids
(as seen in the domesticated Bengal Cat of Canada, the United States, and Europe). Conversely, there are reports and evidence that suggest the opposite, that even if carefully "hand-raised," -
they can not
be domesticated, because they will revert back to their wild and shy behavior as they mature.
lthough there will always be similarities between cats, many characteristics, nonetheless, remain quite variable
(Heptner, V. H. and Sludskii, 1972, 1992). Such observations are especially evident in the various colors seen in their coat (i. e., background colors and the foreground markings). These traits tend to be darker
in the northern geographic areas, and lighter
in the southern regions (Goa, Y. T., et al., 1987; Pocock, R. I. 1939). As examples of coat differences, the ALC of Java and Bali have somewhat dull coloration, the Sumatran subspecies have fewer and smaller markings as compared to cats of the Asian mainland, the Pakistani subspecies coat is quite gray, while the cats of Borneo have rufous and brightly colored coats and markings. In terms of size, f. b. manchurica, is the largest subspecies, while f. p. minuta, of the Philippines, is the smallest (Gao, Y. T., et al., 1987; IUCN Cat Specialist Web Site, on-line, 1996; The Cat Survival Trust Web Site, on-line, 1996).
enerally, the hair, fur, texture, and soft covering of the coat, called the pelage
, and its background and foreground coloring, appear as yellowish-brown, from pale-yellow to orange (with a mix of reddish hues), in the southern climates, and greyish-brown, with reddish brown markings on a yellowish-gray background, in the northern regions (Gao, Yao ting, et al., 1987; Pocock, R. I., 1939). Cream colors with an orange-tinge mix, occassionally dark orange, are quite common, but pigmentation is not overly dark
; the orange hues are referred to an rufous
However, as previously mentioned, color is variable and is determined by hereditary influences, diet, extremes in temperature, the amount of sun exposure, and topographical features (e. g., intermittent elevation differences
and the amount of terrestrial vegetation). The pelt is dotted with dark spots (varying in size and shape), which are sometimes solid, sometimes rosetted, and mixtures of these. The underside of the chin is usually white (sometiimes as off-white color), with no markings. The under-belly is also white, with dark spots, which are usually solid, although rosetted type patterns sometimes appear. The tail is banded (with black rings toward the tip), is moderately long (from 10 to 14 inches), and unlike the "domestic cat," does not come to a point at its end (it has a more rounded tip
). The pads of the feet are dark, usually dark brown or shades of black (never light or with skin-tone colors). Their overall body length, form head to tail, is longer as compared to the standard domesticated cat. Full maturity is reached at approximately 18 months, but sexual/breeding behavior can begin as early as 8-10 months (especially in the tropics). They have been know to live for 15-20 years. Other characteristics of the ALC, as reported in the literature (see IUNC-SSC web site, 1996), and by field observations, are:
(1) The head is relatively small in size, and is more rounded, but not triangular (as viewed from the front). The space between the ears, the upper skull, has a rounded, dome-type shape, but is not flat (see side view, below). The chin is strong with no over-bite, and the under-chin is essentially white. The muzzel is slightly long and somewhat narrow, but not overly long
. The ears are relatively small with rounded tips and are positioned at a slight angle from the mid-point of the head (they are not vertically placed
). The whisker pads are prominent, and the eyes are large with varying colors from amber to grey. Their are usually four
vertical band-type stripes running from the forehead to the back of the neck (variable, as per subspecies), and a degree of white coloration as a background to contrast these stripes (many of these characteristic markings are also observed in the domestic bengal, but are especially evident in early generation hybrids, F1 to F3). The pictures, below, depict various head profiles of the ALC:
-- -- -- --
(2) ALC's utilize habitat uniformly, with a marginal preference for riverine and off-road areas. Of course, this is very much dependent on the topography of the terrain. They adjust quickly to environmental changes, and therefore, are highly adaptable animals.
(3) Behavior is fairly consistent between subspecies, with a demeanor of shyness and wildness, but are non-aggressive. They remain solitary, except during the active breeding season, and generally, have nocturnal feeding habits.
(4) In certain habitats, where there is restricted or reduced movement due to thick ground-vegetation, they are, to some degree, arboreal, living in the heights of trees (Rabinowitz, 1990). Similarly, they live in close proximity to water (Gao et al., 1987), like most wild cats, and avoid deep snow cover as found in some northern climates (i. e., greater then 10 centimeters as reported by some experts). Their diet consists of rodents and other small animals (Heptner and Sludskii, 1972; Robinowitz, 1990), but they will also eat birds, fish, reptiles, and an assortment of insects.
(5) Kittens are born after a gestation period of approximately 65-70 days, with 1-4 young (usuallly 2-3) born in a hollow tree, rocky crevice or burrow. In the nothern climates kittens are born in May, while in the southern regions kittens are found at all times of the year.
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nformation obtained from the internet and other sources, concerning captive
Asian Leopard Cats, indicates there are approximately 30 to 40 of these animals in structured breeding programs
; most of these are found in the European zoological community
. There may even be a larger number
, held privately
, in the United States, although an accurate numeric count, whether it be in the home environment
(as pets), in on-going breeding programs, or in registered commercial enterprize
, is not known. Similarly, despite the loss of some habitat due to human expansion
, with its resultant destructive deforestation
agricultural/plant practices, suggests they are in no immediate danger of extinction
(at least, at the present time). However, many conservation
groups consider them to be at risk
because of over-hunting
and the massive fur trade in cat skins
; this is especially true in China (e. g., The Species Survival Commission, The World Conservation Union, The Nature Conservancy, The Carnivore Conservation Organization, The Small Cat Conservation Alliance, among others). Based on these concerns, the ALC have been placed in the protected categories
(the "Convention on International Trade in Endangered Species of Wild Flora and Fauna
," 1974). This organization monitors the international trade of animal and plant species, many that are endangered (those at great risk
are carefully monitored and placed in Appendix-I
of their guidelines and regulations). At last count, 100 countries, including the United States, follow the "conventions" of this worldwide agency. The ALC are listed in Appendix-II
, as "not endangered
, but could become so if the trade is not regulated"
(of these, only the subspecies, p. b. bengalensis and p. b. euptailura
are listed in Appendix-I
(p. b. euptailura, commonly called the Amur Leopard Cat, is nearing extinction
, at least according to some wildlife experts). Currently, all
ALC, regardless of subspecies "status" (as an example, the Iriomote and Tsushima Cats of Japan), are afforded some protection under the CITES guidelines.
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