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THE LEOPARD CAT
"Prionailurus bengalensis"
(A discussion of the SMALL WILD CAT of central and southern
Asia, often referred to as the Asian Leopard Cat - ALC
)




PREFACE
---Throughout our presentation, some abbreviation was used to reference the Leopard Cat and its hybrid cousin, the Domestic Bengal Cat -(i. e., -ALC - for "Asian Leopard Cat,"- SWC - for "Small Wild Cat,"- and- DBC - for "Domestic Bengal Cat"). - Such usage was necessary, not only to simplify the overall context, but, as well, to enhance and express our own views. We have also included a brief introduction to "wild cats" that relates to their evolution, number, and taxonomy. This inclusion, it is hoped, creates a better perspective of the species, realizing, of course, that there are many unanswered questions and debate concerning their history and development.- In this regard, the use of all facts, historical or otherwise, was always in accordance with current knowledge.- However, we do not include any technical or "in depth" scientific data, as an example, on details regarding genetic transformation, structure, or felid evolutionary theory . Because our articles are fairly long, we have created two files: the present document, consisting of information on "felidae," including the ALC, and a separate article on the Domestic Bengal Cat. -If you want more information, we list major references in the bibliography, and, of course, their are important "online" INTERNET LINKS throughout the present document.

----Concerning the number of ALC subspecies, just remember, this subject is controversial (there is some argument concerning their "status" and taxonomy).- Similarly, specific questions, regarding evolutionary/genetic change and species/interspecies relationships, have not been clearly differentiated, answered, or defined. Therefore, some conclusions and inferences, drawn by the scientific/biological "wild cat" community, are based more on singular events and circumstantial evidence, rather then by direct observation and verifiable data. In other words, there is a lot that is not known about the ALC and its subspecies.

----Since this writing is not a precursor to a thesis or a doctoral dissertation, we did not feel obligated to acknowledge, credit, or list all of our sources in the standard way (i. e., in the precise format of a research paper). - You will also notice that we have included many pictures of the Asian Leopard Cat. Unless they are specifically listed with a "copyright © notice,"- these on-line images are in the "public domain"- and can be used freely by anyone ( see our reference/copyright statement ).- Lastly, we are not so naive to believe the opinions presented in this document reflect the current thinking of the day. Inevitably, the underlying context, shaped by our conceptual ideas and perspectives, are more a reflection of an on-going process, tempered by experience and knowledge, and as such, is a "work in progress." -Nonetheless, we do hope that the information presented here, will, in some small way, increase your interest, your knowledge, and your understanding of these most unusual and beautiful cats.





(THE EVOLUTION OF FELID SPECIES)


---An account of the "family" of cats, know as "felidae,"- is hidden by the subtlety, and perhaps, more abstractly, in the fabric of evolutionary events.- According to what has been learned from the analysis of fossil remains and through theoretical modeling, their development occurred rather quickly over a few million years (Johnson and O'Brien, 1997; Nowell and Johnson, 1996). - However, exact points of divergence into lineages (i. e., into the species and sub-genera of today's wild cats), to global spread, and to primary continental migration patterns, are less clear (such occurrences are sometimes referred to as felid radiation). - Nonetheless, despite unexplained developmental and historical gaps, evolutionary time periods for the majority of mammalian species, including felids, have been described in the scientific literature (Johnson, et al, 2006), and it is generally agreed, with some degree of certainty, that the fossil record is fairly accurate in this regard (as an example, the Oligocene and Miocene epochs, as part of the Cenozonic era.- See Hunt, 1996; Nowak, 1999; Johnson, et al, 2006).

---Briefly, felid-like carnivores first appeared approximately 34 million years ago (mya) during the Oligocene epoch (Hunt, 1996; Pecon-Slattery, et al, 2004; Nowak, 1999).- From a structural perspective, at least, anatomically, many were compact and muscular, but, they were also not as long or as tall at the shoulder.- The wild-cats of today, on the other hand, are relatively long-legged and lithe in comparison (Cox, B., et al, 1999; Turner, A., 1997). Interestingly, it has been suggested, based on what has been learned from fossil remains found in Australia, that some of them were more lion-like marsupials, in some ways, similar in appearance to kangaroos (Turner, A. and Anton, M., 2000.- See PREHISTORIC-CAT WEB SITE for an excellent overview of ancient felids).- Nonetheless, the global spread and development of "present-day" species can be traced to Miocene- times ~ 23 mya, and continued into the periods,- Pilocene ~ 4.5 my- and - Pleistocene ~ 1.8 my- to- 10,000 y- (Hunt, 1996; Pecon-Slattery, et al, 2004; Johnson, et al, 2006).-

---As an example of earlier felids, perhaps the "Saber Tooth Tiger" is the best known (see picture, left).- However, based on fossil evidence, this cat, as well as other ancient and prehistoric felids, are not genetically related to today's species, but, like many other mammals, evolved along separate, parallel paths (Thornhill and Ussery, 1999-2000; Cox, B., et al, 1999; Turner, A., 1997; Hunt, 1996; Nowak, 1999).- Such evolution,- according to some authorities, has, from a theoretical perspective, increased the genetic propensity for specific traits in many of today's plants and animals, and may account, at least in our view, for transformational trends in subspecies development (Merrell, D. J., 1994; Johnson, G., 2008).- These changes, of course, have been impacted by geological events which have lead to the formation of major continental landmasses, and, ultimately, to the creation of unique ecosystems (e. g., as exists in the delta and basin areas of southeastern Asia, in the surrounding regions of Indonesia, and in Australia and New Zealand.- See the work of Johnson and O' Brien, 1997 and Nowell and Jackson, 1996).-Interestingly, it is believed, that under the right conditions and over evolutionary time periods, "nature," set in motion a sequence of changes that has favored certain organisms, to the extent, that unrelated species have assumed "like" characteristics in response to habitat and to genetic influences (see the blog, "Laelaps," for an account of evolutionary developments). -Chance or coincidence alone, however, can not explain this occurrence, but rather, when viewed within the context or framework of scientific inquiry, suggests some form of natural selection and convergent evolution (see the writings of Dr. George Johnson, online, 1998-2008).- When considered together, these theories imply how species can sometimes exhibit strikingly similar physical features, similar adaptations and behaviors, yet, at the genetic level, remain distinct (Gould, S. J., 2002; Berger, et al, 1979).- Among mammals, one of the best examples of convergence can be seen in placental animals and the marsupial species of Australia (pictured to the right, in the next paragraph).- In the plant kingdom, this phenomena can be seen in the adaptive characteristics of flora growing in desert ecosystems (e. g., the catus, milkweed, and euphorbia families; see explanation at bottom of page under "convergence of plants" - Cornell Univ., tutorial course, Intro. Bio. 105/106, 2008).- It is noted, that these theories do not explain the large number of felid subspecies, nor, do they delineate, developmentally, an exact or definitive relationship between genetics and exogenic forces (see ecological niche factors, as an example).- Nevertheless, they do offer a partial rendering and explanation for the proliferation and wide variety of organisms observed across environmental systems, and as importantly, the immence biodiversity seen in the world today.

---In terms of hereditary influences, questions remain regarding evolutionary change, genetic relationships, and the scientific categorization of felids.- Here, at least in a comparative sense, an accurate baseline, grouping, and description of ancient cats, and any genetic link to today's species, has not been firmly established or defined.- Consequently, there is some disagreement within the biological wildlife community and among paleontologists concerning:- taxonomy, subspecies order, evolutionary processes, and ancestral characterizations.- Likewise, there are issues relevant to statistical inference, and how the correlation of genetic data and fossil remains should be used to explain felid evolutionary development (e. g., Bayesian posterior probabilities for DNA;- J. P. Hulsenbeck, el al, 2001, "Bayesian inference of phylogeny".- Also, see the work of Johnson, et al, 2006, and the genetic supplemental data for more detail).- To date, these questions, including unbiased explanations for genetic evolution, interspecies relationships, and to matters of "Darwinism,"- have not been resolved, and it is doubtful, at least in our opinion, that there will be a consensus, anytime, soon (since some of these issues become entanged in the precepts of religious philosophies, theological considerations, and creationism).- However, a better understanding of "convergent evolution" and "natural selection,"- a closer examination of fossil remains, and more precise DNA studies, will eventually untangle the "web" of confusion surrounding these subjects, and perhaps, create a more perceptible and unifying picture of the relatedness of species, both living and extinct (Johnson, et al, 2006; Nowell and Jackson, 1996; Nowak, 1999).

---Although it is probable that the progression of evolutionary genetic change among felids was set in motion as early as 34 Million years ago (mya), the beginning of significant transformation, culminating in the "wild cats" of today, did not occur until later (Johnson and O'Brien, 1997; Pecon-Slattery and O'Brien, 1998).- Suggested time intervals are from 23 mya, especially between 10.8 ~ 6.2 mya (Pecon-Slattery, et al, 2004; Johnson, et al, 2006; Mckennan and Bell, 1997; Hunt, 1996).- More specifically, at least according to many mammal/evolutionary biologists, major species radiations, including global spread and anatomical variability, were impacted by:- (1) "rapid" speciation events, (2) the affects of genetic drift and adaptive radiation on isolated, and in some cases, small or niche populations, (3) exogenic forces, that is, geo-enviornmental/ecological variables (e. g., the structure or makeup of habitat, weather patterns, glacial oscillations, changing sea levels, tectonic plate shift of land masses, among others), and, according to some authorities, (4) the beginning, the differentiation, and the genetic expression into dominate species through a process called natural selection (Mattern and McKennan, 2000; Nowell and Jackson, 1996; Hunt, 1996).- Naturally, some of the aforementioned events and their interaction are inferred from "theoretical models,"- but according to the work of Johnson (2006), as based on the interpretation of updated fossil data, sudden changes in sea levels, especially, at low levels, corresponds fairly well to major movement across land-bridges and the establishment of lineages from Miocene times; with major continental migrations from locations in Eurasia, primarily, the central Asiatic regions (as an example, the Panthera and Lynx lineages and subspecies, thereof. Click on the picture, upper-left, to see a larger composite map of time intervals, sea levels, and the appearance of felid species).

---From a purely developmental perspective, some interesting questions have been raised regarding rapid evolution relative to ecological/environmental response, and whether the attributes of such systems affect the outcome and speed at which genetic change occurs (Hairston, et al, 2005; Gingerich, P. D., 2001, 1993; Thompson, J. N., 1998).- Theoretically, it is resonable to assume, based on current knowledge, that the make-up of the external environment probably altered the course of evolution, and perhaps, changed the character and pace of biological /anatomical variability of all living things, especially, vertebrate species and the genera of mammalian subspecies (Thompson, 1998; Johnson, et al, 2006)?- Such premise has created debate among evolutionary biologists regarding the interaction of these variables and their effects.- However, finding definitive answers to these questions is difficult, simply because the very nature of evolution is often clouded by an incomplete fossil record and our interpretation of such information (including the components, complexity, and statistical data associated with it). In this regard, the scientific research seems to imply that the field of genetics and analysis, thereof, can best describe such relationships.- On the other hand, others believe that exogenic forces have significantly impacted, and thus, are intrinsically linked to such events.- Of these, one of the most important, in our view, is the study of ecological dynamics (see the scientific letters of Hairston Jr., et al, 2005).- Here, do sudden changes in environmental conditions influence the transformation of genetic traits?--As well, does "rapid" evolutionary change correspond to significant ecological performance, and do these changes occur on similar time scales (Hairston, 2005, 1986; Gingerich, 1983, 2001; Hendry and Kinnison, 1998; Grant P. R. and Grant, B. R., 1995)?- There is evidence to support such claims and investigation because environments are never static.- Likewise, the components of ecological systems can vary at dramatically different rates, influenced, as an example, by rapid bio-ecochemical fluxes, as compared to the more measured progression of global climate change (Hairston and Walton, 1986; Hairston, 2005).- It will be interesting to monitor, as more geogenetic information is unearthed, how science unravels such complex relationships and how this may alter our understanding of evolutionary processes.


---Ironically, it has been suggested that a large portion of felid evolutionary history may not be represented in the fossil record (Johnson, et al, 2006), yet, the formulation of present "theoretical models" are often based on such findings.- Ultimately, resolution of this paradox will require an analysis of large, multigenic samples (Johnson, 2006), and a clearer understanding and definition of the developmental relationships between genetics and the natural environment, relative to evolutionary processes.- However, in a review of the literature, very little data currently exists on this subject (at least, in meaningfully large enough data sets to have significant statistical relevance and correlation).- There is, of course, considerable debate regarding these matters and to many aspects of phylogenetics and tree of life issues (some of these are philosophic, while others are grounded in the arguments/complexity of empirical science, theology, and, according to some, the pseudoscience of "intelligent design").- So, if you decide to review the scientific literature, bear in mind, that although the fossil record is generally clear,- some "theory" is usually involved in all explanations of evolution.- Therefore, regardless of prevailing thought and conjecture, it is our opinion, that the correlation of ecological conditions to rapid evolutionary genetic change, as well as estimates of respective time intervals, dates, and the like, are, nonetheless, fairly accurate representations of actual events.- More philosophically, or perhaps, from a pragmatic perspective, since man holds stewardship over the animal kingdom, including the protection of worldwide ecosystems and habitats, an understanding of the relationship between exogenic forces and evolutionary change, regardless of species, has far-reaching implications.- With such knowledge, man will be better equipped to find logical solutions for the beneficial use and management of earthly resources, and more profoundly, how he can best live in harmony with the natural environment, hopefully, without destroying it, first!




(THE PRESENT-DAY -"FELIDAE"- FAMILY)



-- Many wildlife authorities and mammal biologists identify 37 species of "felidae" worldwide (Corbett and Hill, 1993; Honacki, et al, 1982; Guggisberg, 1975; Green, 1991; Alderman, 1998), and this has been, until recently, the most widely accepted view.- However, based on the interpretation of genetic evidence from felid fossil remains by Warren Johnson, et al, in 2006, there may be as many as 40, all descended from a common ancestor 10.8 million years ago.- Similarily, 60 % of the known modern species of cats probably developed within the last million years (Mott, 2006). Nevertheless, according to present evolutionary, genetic, and morphological data (e. g., biological and geological), a more comprehensive and representative update suggests 36 SPECIES ( Felid Taxonomic Advisory Group, 1996-2006;
Wilson and Reeder, 1993-2005).-Among these, the larger cats, as an example, the lions, tigers, and leopards of the genus, "panthera,"- are believed to be the oldest species, while many smaller felids, such as those belonging to the genus, "felis" and "prionailurus,"- are considered the youngest. Of the wild cats, the Cheetah, in our opinion, is one of the most interesting, its long body supported with powerful legs, built for speed, and paws that are almost doglike. These characteristics, among others, distinguish it from other cats, and because of its unique qualities, it is the only feline in the taxonomic group, "Acinonyx," with five subspecies (although the number of sub-genera is argured). Regarding anatomical size, the largest of cats is the "Siberian Tiger,"- while the smallest is the "Rusty-spotted Cat" of Indian and Sri Lanka (consult the following web-sites for more in depth information regarding FELIDAE:-Big Cats Online, the Cat Specialist Group, and Animal Diversity web sites).


----Interestingly, one possible explanation for differences in the "numeric count" of felid species may relate to the "status" of the Iriomote Cat (sometimes referred to as "Felis iriomotensis" in the literature, and found only on the island of Iriomote-jima of southern Japan, in the Ryukyu island chain, east of Taiwan; see map, left).-This feline is believed to be either:-(1) a subspecies of the Leopard Cat (listed as "Prionailurus bengalensis iriomotensis"), (2) a close relative of the Leopard Cat, but a genetic variant (listed as "Prionailurus iriomotensis"), or, (3) a distinct species (in which case, it would be the sole member, as "Mayailurus iriomotensis").- If it is eventually proven not to be a subspecies of the "leopardf cat,"- it is probably one of the most ancient and rarest of cats (see Cat Specialist Group, 2007), in which case, if you follow the "numeric debate,"- the actual count of felid species would be 37.- However, based on current phylogenetic evidence, the concensus among many wild cat specialists indicates three primary sub-familes (i. e., FELINAE, PANTHERINE, and ACINONYCHINAE), with 18 GROUPS and 36 species, therein (Wilson and Reeder, 1993-2005; Wozencraft, 1993; Shoemaker, 1996; Felid Taxonomic Advisory Group, 1996 and 2006; IUCN Cat Specialist Group, 2007; review the breakdown of the "felidae" family to the right). We wish to point out, this particular representation of wild cats, although the current "popular" view, is subject to change as more information becomes available. It is, therefore, not conclusive, but in our opinion, is the most logical based on current genetic data and taxonomic description ("subspecies order" and numeric accounts, not surprisingly, continue to be debated by wildlife and taxonomic experts). Additionally, wild cats are normally grouped into five geo-enviornmental areas (i. e., the continental regions where they are found):- Sub-Saharan Africa, North Africa and Southwest Asia, Tropical Asia, Eurasia, and the Americas (see the Cat Specialist Group for more in depth information and a breakdown of current species vulnerability).


----Upon close inspection, disagreements (at least, regarding the identification, number, and taxonomy of "felidae"),- have complicated the formulation of a more clearly defined felid classification. This has been, to some degree, a source of annoyance and argument among academia and the general wildcat community for many years, and continues to be so. More specifically, current taxonomic information suggests, that the majority of debate relates to differences in the number of "taxa" within the family of cats, the logical placement of species within groups, and the hierarchical ordering of subspecies. Likewise, there is the underlying inclusion and interpretation of ongoing research to consider, and, as importantly, the lack of pertinent and corroborative genetic information (which has added another layer of complexity and confusion to the debatable landscape).- Therefore, just be aware, if you browse the academic literature on the web, or study other sources related to "felid taxonomy,"-scientific or otherwise, you will find more then one explanation and opinion, especially, issues related to subspecies. Nonetheless, the combined work and research of Wozencraft (1989, 1993, 2005), Wilson and Reeder (1993, 2005), the Cat Specialist Group (1996, 2007), The Felid Taxonomic Advisory Group (2006), and the protected status of cats as monitored by CITES (2007), are often referenced as the standard for current felid species (see legal status of felidae). Concerning the domestic cats, Felis silvestris catus, they are currently listed under the African subgroup, Felis silvestris lybica, which, itself, is one of three distinct varieties belonging to Felis silvestris, known as the -"Wild Cat" (see Animal Diversity Web for more information).- However, there is, even here, disagreement regarding such order and designation.


---- Although evolutionary change has created variability in overall physical appearance among felids, behavioral differences, shaped by environment, are also noted across populations.- Still, within the general framework of taxonomy, the "family" of cats is fairly well established, but phylogenetic questions related to origin, to subspecies status, and to interspecies relationships, are less clear.- However, some of these issues were addressed in a fairly recent and important publication, The Late Miocene Radiation of Modern Felidae: A Genetic Assessment (Warren E. Johnson, et al, in SCIENCE, vol. 311, January 6, 2006. You can view this document on our web site, make sure you have the Adobe Reader plugin installed in your web browser).- Briefly, this study proposes an updated biogeographic hypothesis of felid evolution based on the statistical analysis of fossil remains and on geological events. The authors suggest that past intepretation of genetic information has underestimated the first occurrence of "lineages" by an average of 76 %.- Further, Y-chromosome segments are more informative than mitochondrial DNA, X-linked, or autosomal genes in identifying genetic relationships, and lends itself to a better understanding of "felid radiation" (and the rapid extent, thereof).- From a purely phylogenetic perspective, their research more closely defines causality, intercontinental migration patterns, and ancestral tree structure, which, intrinsically, increases our understanding of genetic diversity within felid populations. Nonetheless, although there are unresolved issues related to taxonomy and to evolutionary events, -all species of cats, for the most part, are differentiated and easily recognized in their natural environments (Wozencraft, 1993; the Cat Specialist Group, 2007; see tropical Asia felid species).

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-Prionailurus Bengalensis -


THE LEOPARD CAT
SCIENTIFIC-LATIN-NAME: "Prionailurus bengalensis" (Pocock, 1917); sometimes listed as "Felis bengalensis" (Kerr, 1792).
COMMON-ENGLISH-USAGE:- Leopard Cat, Asian Leopard Cat, Bengal Leopard Cat, Spotted Bengal Cat.

CURRENT TAXONOMIC LISTING
KINGDOM: animalia.- PHYLUM: chordata.- CLASS: mammalia.- ORDER: carnivora.
FAMILY: felidae.- GENUS: prionailurus.- SPECIES: bengalensis




--- When you review the scientific literature, taxonomic data, and relevant information from various sources, especially those "online,"- you will notice that many authorities group 28 to 30 of the known felid species into a category called, "small wild cats" (SWC).- The Leopard Cat, "Prionailurus bengalensis" (Pocock, 1917; Wozencraft, W. C., 1989; Wilson and Reeder, 1993, 2005), originally defined as "Felis bengalensis" (Kerr, 1792), and placed in the genus, "Prionailurus,"- by Pocock (1917), is listed in this category (see its current grouping, in green, above, right, and its taxonomic listing, below). It is referred to, in English, as the Leopard Cat or Asian Leopard Cat in many publications (abbreviated in the present document as ALC). They were first observed near the "Bay of Bengal,"- in India (see the accounts of Pocock, 1917, 1939), hence, the scientific designation "bengalensis," and the English derivative, "Bengal,"- appears in the literature. Colloquially, this name is also used by domestic cat organizations and the general cat "fancy"-to denote the hybrid form, widely referred to as the Domestic Bengal Cat.- (As an addendum, there is disagreement over an appropriate non-scientific or common name.- Here, Bengal Leopard Cat and Spotted Bengal Cat, among others, are sometimes used. However, ASIAN LEOPARD CAT-is more logical, since they appear to be endemic to the central areas of southeastern Asia, especially in the southern regions, and because this name is more closely aligned to its scientific description.- Regarding the SWC category and ALC subspecies account, whether inclusive to a particular group or linked to a more generalized classification, continues to be debated).

----Wildlife experts consider the ALC and its subspecies to be the most common of small-wild-cats (Alderman, 1998), although they are not widely known in western cultures (the Bob Cat, Lynx, and Ocelot, as examples, are some of the more familiar SWC's).- Still, they exist in large numbers over a wide geographic area in 21 Asian countries, primarily, those of the central region (Santiapillai and Suprahman, 1985; Johns, 1989; Rabinowitz, 1990). The largest concentrations, however, can be found in southeastern Asia (i. e., east of India and south of China), especially, subcontinent locations and island nations within the Indian Ocean, South China Sea, Java Sea, Bay of Bengal, and the general area of Indonesia and Malaysia (excluding Australia and some island groups of eastern Pacific proper, sometimes called Oceania). Some authorities believe they are indigenous to the central Asiatic mainland, but from a phylogenetic and historical perspective, an exact point of origin, to date, has not been clearly established (See picture, left,- shaded green represents the primary range of the ALC. CLICK on image to see a larger map).- Habitable environments are many, and include: mountainous areas, up to 3000 meters (Habibi, 1977; Feng et al., 1986), tropical rainforests, temperate broadleaf coniferous forests, deforested riverines, shrub forests, semi-desert regions, agricultural land, low elevation mountain forests, jungles, and successional grasslands, among others (Heptner and Sludskii, 1972; Feng et al., 1986). Their overall geographic range encompasses the following areas, but may also include adjacent land and unknown habitats in bordering countries (Nowell and Jackson, 1996):



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(THE SUBSPECIES AND EVOLUTIONARY DEBATE)

----Interestingly, 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 -"subspecies status."- 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.

----As 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:


----Although 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 (see allopatric),- 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"-world.


----Not 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
--molecular analysis, including genetic analysis, and most classically described
--cat subspecies are not valid based on current genetic information."


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(SUBSPECIES VARIABILITY)

--- If- you examine pictures of the ALC and its subspecies, you will notice that no two cats look alike (see pictures throughout this writing).- Discounting genetic change, such development, in our opinion, is due, in part, to 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 (Heptner, V. H., 1971; Johns, A. D., 1989; Yu, J., and Wozencraft, 1993), with attributes, that, for the most part, are easily recognized (e. g., differences in color, markings, and in some cases, overall size).

--- From a phylogenetic perspective, the primary mechanisms responsible for physical variabiity among ALC subspecies remain unexplained. However, speciation events,- random occurrences, genetic drift, undefined and subtle genetic interactions, and geo-environmental affects (the influence of climate and topography, as examples), among others, are major factors that have impacted change and developmental tendencies in these cats. These, in turn, have intensified physical variability among subspecies.- Whether such events are due more to genetics, or, to what degree and extent environmental factors have played in the outcome, is not known. In one sense, it is easy to assume that genetics are responsible, after all, biological -"animalian"- structure is primarily genetically based, although the influence of environment may be more important then is currently believed. Nonetheless, in our view, any theory that relates to changes in outward physical appearance 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 more theory then fact. Still, fragmentation, in whatever form, 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:

--- It 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. "

----As 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 of territorial, 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, academic, logistical , 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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(TAXONOMIC DESCRIPTION)


--The 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,"- a regional 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).

---Many 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,-"Prionailurus,"- 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.

--- As 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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ALC TAXONOMY
KINGDOM: animalia.- PHYLUM: chordata.-SUBPHYLUM: vertebrata.- CLASS: mammalia.
- ORDER: carnivora. FAMILY: felidae.- GENUS: prionailurus. -SPECIES: bengalensis
(the vertebrata category, under subphylum, referes to all species with vertebra, that is, the fishes,
amphibians, reptiles, birds, and mammals, but does not appear in many taxonomic listings)


*ALC SUBSPECIES

* * F.(P.) b. bengalensis ---- - The regions of India to Indo-china and Yunnan.
----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 number of subspecies and their classification is much debated; THIS ISSUE HAS NOT BEEN RESOLVED AS OF 2007). 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.

--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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(CHARACTERISTICS)


----Actual 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.

-- Research 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.

--- Although 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).


--- Generally, 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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(PROTECTION)

-- Information 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 and poor agricultural/plant practices, suggests they are in no immediate danger of extinction (at least, at the present time). However, many conservation and animal-watch 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 of CITES (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 as endangered (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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REFERENCE STATEMENT and DISCLAIMER

The context of this article, including page layout, was designed and written by IMAGINIQUE BENGAL CATS, Inc.- Its content is copyrighted, and use of the material, in any way, is strictly prohibited (unless permission is granted from the author).- Some of the information was obtained from journals, text books, scientific/academic web sites, the "Bengal Bulletin" (March 1995), and the Cat Specialist Group of the World Conservation Union (1971). Also, condensing all of the information into a definitive and concise form was accomplished through many hours of research and hard work. Therefore, if you wish to use any of our "text" in an article, or, on your own web site, you are granted permission to do so, but only if you give credit and "list" us as a reference in your writings (primary sources for our article can be found on the REFERENCE page). Lastly, this document is an EDUCATIONAL RESOURCE, and has been edited for content and in accordance with current knowledge. However, we cannot guarantee the accuracy of all data from other authors. Likewise, all species of "felidae" are not covered in this document, nor does it list the latest scientific information or accounts, thereof (for this, the reader is referred to the literature).


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