Belated welcome to a 'new' clouded leopard.. named in 1823

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The naming of any new large mammal species is always an exciting event, and within the past few days you've probably heard much in the news about the formal recognition of a new species of extant big cat: the Indonesian clouded leopard Neofelis diardi (that's not its formal common name by the way, but it'll do for the time being). However, I confess to being somewhat confused...

You'd think from some of the global media that (1) N. diardi has only just been discovered, and (2) the study announcing its discovery has only just been published. On the first count, I note that the wording used by many journalists is in fact accurate in that they are taking care to state that new studies have shown that long-known populations have only just been realized to be worthy of species status, not that the species itself is a brand new discovery. Like many tetrapod taxa conventionally regarded as a subspecies, N. diardi was originally described as a full species. This happened way back in 1823 when George Cuvier coined Felis diardii in honour of his student Pierre-Medrad Diard*, and F. diardii was widely used later on for, apparently, all clouded leopards by authors including Desmoulins, Blyth, Jerdon and Sterndale (Guggisberg 1975).

* I say bring back the noble Cuverian tradition whereby supervisors name new taxa after their students. Hint hint.

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Sir Thomas Stamford Bingley Raffles had beaten Cuvier to it, however, by describing clouded leopards in his descriptive catalogue of new specimens from Sumatra (Raffles 1822). Cuvier and Raffles must have known what the other was up to, as Diard and Raffles travelled together to Sumatra in 1818. Raffles didn't name the species however: the first to do this was Edward Griffith in 1821, and this is where Felis nebulosa was first coined. In 1867 John Edward Gray decided that the clouded leopard needed its own genus and coined Neofelis for it, hence the modern binomial Neofelis nebulosa.

A few things confuse me here. Firstly, Raffles' volume is always cited as 1822 which, if correct, raises doubts as to whether Griffith's publication really appeared in 1821 as always said. Maybe Griffith saw a pre-print? Or maybe one of the dates is wrong? As many of us have learnt from bitter experience, the 'established' citation date of any given taxon is not always correct (one of the best known examples is Dinosauria, long given as Owen, 1841 when it was actually not published until 1842). Secondly, Griffith stated that the specimen described by Raffles came from Canton in Guangdong, China, and not from Sumatra as you might assume given the title of Raffles' volume. Thirdly, I've never seen Cuvier's original reference, but some sources (e.g., Guggisberg 1975) give Cuvier's original spelling as Felis diardii. There's an awful lot of inconsistency as to whether specific names are spelt with a double i at the end or not, and I still can't recall what the rule is... if there is one (for now I'll use N. diardi because everyone else is).

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Anyway, as Griffith's naming of Felis nebulosa became better known during the 19th century, F. diardii and two other taxa, F. macrosceloides (endemic to Nepal and the adjacent parts of India, Bangladesh and so on) and a Taiwanese taxon first named as Leopardus brachyurus - today it's N. nebulosa brachyura - became regarded as subspecies of F. nebulosa. In the case of N. diardi at least, we're back where we started: a taxon originally named as a distinct species turns out, after all, to be a distinct species. Is this a case of over-zealous lumping burdening us, yet again, with an incorrect understanding of taxonomic diversity? Or should we be more prosaic and agree that good, solid science has tested one hypothesis (viz, that N. diardi is a clade within N. nebulosa) and found it wanting? That's meant to be rhetorical question by the way (for previous discussions of over-lumping in mammals see The many babirusa species: laissez-faire lumping under fire again). The recognition of N. diardi as a valid taxon obviously has great significance in terms of conservation: like it or not, 'species' are rated as more important and deserving of effort than 'subspecies', and it's clear that conservation work on Borneo is about as deserving as it could possibly be (note that N. diardi isn't endemic to Borneo: it's also present on Sumatra and elsewhere in Indonesia).

I'm not sure what all of this means for the supposed subspecies N. nebulosa macrosceloides and N. n. brachyura. The latter is supposed to be extinct, with the most recent sightings being from the 1960s, and even they are little better than rumours. But it always sounded to me like an unusually odd clouded leopard, being smaller, more brightly coloured and shorter-tailed than the others.

Moving on.... regarding the second count (the implication from the media that the study announcing the species-level distinction of N. diardi has only just been published): well, the relevant publications appeared... not this week, nor last week, nor last month, but last year. In the December 2006 issue of Current Biology, Valerie Buckley-Beason and an army of co-authors showed how a substantial amount of genetic difference observed between the clouded leopards of the Asian mainland with those of Borneo indicate that the two should be recognized as distinct species-level taxa. Indeed the amount of variation observed is equivalent to, or exceeds, that present between various of the Panthera species. Buckley-Beason et al. therefore suggested that N. diardi might warrant recognition as a separate species, and noted that morphological studies should be carried out to confirm their results (Buckley-Beason et al. 2006).

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Exactly this was done by Andrew Kitchener and colleagues, who published their morphometric data on clouded leopard coat patterns in the same December 2006 issue of Current Biology. From coat pattern differences, they also concluded that the clouded leopard includes two species (Kitchener et al. 2006). So, both molecular and morphological data agree in indicating that N. diardi is a valid species, distinct from N. nebulosa. All of this caused quite a bit of excitement and discussion in the zoological world, particularly among those working on carnivoran conservation. If you don't believe me, check out this December 2006 page from Carnivore where felid specialists brought attention to the two studies, noting how this means that we should all update our species lists [adjacent range map of clouded leopards from here].

The fact that the results of Buckley-Beason et al. (2006) and Kitchener et al. (2006) have only now been released as 'new' is a bit odd then, but I'm not down-playing these studies, or their announcement, by any means. While it's standard for press releases to go out at the same time as the publication of a new study, there are also many occasions when news is held back for one reason or another. So it seems that the big press release wasn't made in December 2006, but now, in March 2007, instead.

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As any 'good' species should, N. diardi does look quite different from N. nebulosa: in the article here, the top two images show N. diardi while the third shows N. nebulosa. Some sources imply that Indonesian clouded leopards are also different in their behaviour from other forms: Rabinowitz et al. (1987) found Bornean clouded leopards to be predominantly terrestrial, for example, which is odd given the many specializations that all clouded leopards have for scansoriality. Unlike most other felids, clouded leopards have particularly flexible ankle joints that allow them to descend head-first down a tree (Hemmer 1968), and to hang by their hind-feet alone. They are even claimed to be able to hang by one foot alone (Kitchener 1991). They have the longest tail of any cat; it accounts for 45-50% of their total length and is used to aid balance, and reportedly as a rudder when they leap, and they also have particularly long hindlimbs, apparently enhancing their leaping ability (Gonyea 1976). However, whether clouded leopards of any population are really as arboreal as sometimes thought is open to debate: the sad fact remains that we know very very little of the ecology and behaviour of these cats in the wild. Adjacent pic of clouded leopard teeth from here.

On another point associated with discussion of N. diardi in the media, I am - as usual - frustrated by the fact that I've now heard several people on TV state such things as 'the discovery of a new large mammal is a very rare event, and not likely to happen again for a long while'. Comparatively speaking, yes, new big mammals are rare: if you compare their discovery rates with those of new insects, fish, lizards or rodents.


But they aren't really rare, and certainly not the 'once in a lifetime' or 'once in a century' event that some journalists (and scientists) state or imply. Saola Pseudoryx nghetinhensis, named 1993 [shown in adjacent pic]. The new muntjacs Megamuntiacus vuquangensis, named 1994, Muntiacus truongsonensis, named 1998, and M. putaoensis, named 1999. Dingiso Dendrolagus mbaiso, named 1995. The new brocket Mazama bororo, named 1996. Arunachal macaque Macaca munzala, named 2004. And so on and so forth. There are also several new taxa of extant large terrestrial mammals - now represented by specimens housed in collections - that await official description and naming, including the giant peccary from the Amazon and, apparently, two new big cats from Peru (sigh, when is Peter Hocking ever going to publish?). Exactly the same sort of mistake was made when a new European rodent, the Cypriot mouse Mus cypriacus, was described last year. Rather than being the 'first new European mammal in 100 years' as some journalists said, it was, like, the 33rd new European mammal in 100 years! (go here: The first new European mammal in 100 years? You must be joking).

Anyway, I need to stop there and get back to the editorial work I'm supposed to be doing. More on big cats in future, including on the diversity of lions and... whence the onza? I'll also have to write more on clouded leopards in future: there's the fact that they have extraordinarily long upper canines for one thing, leading some workers to regard them as 'living saber-tooths'.

Refs - -

Buckley-Beason, V. A. Johnson, W. E., Nash, W. G., Stanyon, R., Menninger, J. C., Driscoll, C. A., Howard, J., Bush, M., Page, J. E., Roelke, M. E., Stone, G., Martelli, P. P., Wen, C., Ling, L., Duraisingam, R. K., Lam P. V. & O'Brien, S. J. 2006. Molecular evidence for species-level distinctions in clouded leopards. Current Biology 16, 2371-2376.

Gonyea, W. J. 1976. Adaptive differences in the body proportions of large felids. Acta Anatomica 96, 81-96.

Guggisberg, C. A. W. 1975. Wild Cats of the World. David & Charles, Newton Abbot.

Hemmer, H. 1968. Studien zur Ethologie des Nebelparders Neofelis nebulosa (Griffith 1821) und des Irbis Uncia uncial (Schreber 1775). Veroeffentlichungen der Zoologischen Staatssammlung Muenchen 12, 155-247.

Kitchener, A. C. 1991. The Natural History of the Wild Cats. Christopher Helm, London.

- ., Beaumont, M. A. Richardson, D. 2006. Geographical variation in the clouded leopard, Neofelis nebulosa, reveals two species. Current Biology 16, 2377-2383.

Rabinowitz, A., Andau, P. & Chai, P.P. K. 1987. The clouded leopard in Borneo. Oryx 21, 107-111.

Raffles, T. S. 1822. Descriptive catalogue of a zoological collection, made on account of the Hon. E. I. C. in the island of Sumatra and its vicinity. Transactions of the Linnean Society of London 13, 1-239.


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You're complaining about overlumping in large mammals? Give it a break! The entire mammalian radiation, were it taxonomized like an insect clade, would represent a single genus.

Excellent and thorough post (definitely puts mine to shame). I too was puzzled, upon learning when it was actually determined that the Borean Clouded Leopard was a valid species, why the story was held back. As well, I found it frustrating that the popular news outlets were heralding this as a "discovery" rather than some taxonomic redefinition (although I am glad it showed up in the news at all). In any event, wonderful stuff, thanks!


The double i in "diardii" comes from Cuvier first Latinizing Diard's name to "Diardius", then putting the Latinized name in the genitive case. This was common practice for a long time.

While modern convention favors adding the (second-declension) genitive singular suffix -i directly to the name, I'm not sure that the standard can be applied retroactively, and my guess is that the spelling "diardii" should stand.

By Nick Pharris (not verified) on 16 Mar 2007 #permalink

There's an awful lot of inconsistency as to whether specific names are spelt with a double i at the end or not, and I still can't recall what the rule is... if there is one

Nope. We're just supposed to stick with the original spelling.

The only rule about this is that if two species names in the same genus differ only in this respect, they are considered homonyms.

By David Marjanovi? (not verified) on 16 Mar 2007 #permalink

Great and very thorough post. Do you have an opinion on the molecular evidence that defines the two species? As I mentioned to someone earlier today, I don't think that being able to investigate genomes has made it any easier to define a species, it has simply given us another definition.

Thanks to all for their comments. On -i vs -ii endings, I did sort of remember that we should try to stick to the original spellings. This is particularly relevant for a big review paper I currently have in press, where I deliberately used original spellings: the result being that some species names end in -i, and others in -ii. To be consistent, I had to be highly inconsistent.

Sarda: despite the fact that there are now twenty-something different species concepts, I think most biologists can agree that the concept of 'the species' is effectively subjective and how much variation we're prepared to accept differs from group to group. On molecular data, so long as there is a general agreement as to how much variability is acceptable, then fine, even if this means that a 'species' within one group may encompass ten times as much variation as a 'species' within another. This problem has been characterised as the 'frog's eye view vs the bird's eye view' (in the former group, 'species' exhibit a wide range of genetic variation while in the latter 'species' are more narrowly defined).

I dunno if what I'm saying makes any sense, but once we get over the idea that a 'species' is an entirely artificial unit (which it is, given the fact that evolution happens), all we have to do is agree that a 'species' is essentially whatever is convenient. Cue that John Ostrom quote..

Fascinated by the hint there may be two Peruvian big cat species to be described... can you give any info?

Excellent post, as always.

Sigh...can't the media at least do 5 minute's worth of fact checking? I bet some poor mixed-up kid is already using this of "proof" of a Cryptozoological success!

On that subject, is this the same Hocking who described the giant black panther, speckled "tiger", striped "tiger", jungle wildcat, jungle "lion" and "leopard-spotted" jaguar mentioned on the Strangeark Guide to Cryptids? Are any of these the two cats that apparently have specimens? Is there any resource for these cats not published in a defunct journal? This all sounds incredibly fascinating, and I'm wondering why those with Cryptozoogical sympathies seem to ignore that which is (more) probable and focus on the improbable.

I'm still hopelessly addicted to your blog, best on the web for sure. And a potential onza post? I'll be in Lord Geekington heaven.

Somewhere(*) recently I've read that new mammal species are being described at a rate of several dozen a year, and that the number of recognized extant mammal species is up to about 5400, having been in the low 4000's a few decades ago. What wasn't stated, however, was what PROPORTION of new species represented discoveries of new ... I'm not sure how to word this ... new and previously unrecognized populations of mammal differing at species level from those previously known, and what proportion represent splittings of previously recognized species. If I remember your post on Mus cypriacus correctly, it was (like the Indonesian clouded leopard) a case of the latter sort: everyone knew there were mice there, but it was assumed that they were conspecific with those somewhere else. I think the largest new mammal (Omura's (?) whale) is also a case of that sort: they were known and hunted, but assumed to be (I think) Bryde's whales.

There are probably cases where it's hard to classify the sort of discovery involved: new species of mouse from somewhere in Brazil, indistinguishable to a layman from seventeen others, from a locality not previously sampled but not too far from sampled ones inhabited by another, similar-looking, species: do we call this a new animal discovery or just a splitting of THIS locality's mice from those of the neighboring valley? So there may not be any statistics available, or even easily extractable from the mammalogy literature. Do you have any sort of off the cuff gut feeling about what the proportions would be, though?


(*) I guess it was in a book review, possibly in "Journal of Mammalian Evolution", of a newly published checklist of extant and recently extinct mammals.


(Oh. Old business: can you give me any sort of pointer to information on the suspension-feeding placodont you mentioned in responding to my "comment" on your sebecosuchian post? I'm a bit of a Christine Janis fan (she strikes me as someone who isn't afraid to tackle big questions even if they can sound unfashionably vague) and I'm going to try to interlibrary-loan the Ancient Marine Reptiles book to see what she (& co-author) had to say even if it WAS based on a false premiss.

By Allen Hazen (not verified) on 16 Mar 2007 #permalink

Thanks for comments. On onzas and the new Peruvian cats, rest assured that I have the goods and will deliver, one day. Yes Cameron, you have the right Peter Hocking in mind. Some of his cryptids are not all they're cracked up to: I discovered that the yana puma (aka giant black panther) is actually an ethnic name for the spectacled bear, for example.

Skulls of the 'speckled tiger' and 'spotted tiger' have both been procured and photographs are available in the literature. Matt Bille showed these photos to a big cat worker (I forget whom) and got some pretty positive results, which he published. Matt reads this blog, so if you're reading this Matt and have any further comments, feel free to contribute. Regardless, I'll blog on the Peruvian cats - and onzas - soon. In fact I'm aiming to get some new information on the 1986 onza specimen at the up-coming big cat meeting: stay tuned.

Allen: IIRC the current discovery rates for mammals are a steady 15 or so per year, though that's a date I recall from the early 1990s and the current figure is probably higher (yes, HIGHER). You're dead right that a lot of 'new' mammal species result from taxonomic revisions, not necessarily from the finding of new taxa in the field. I still get the impression that half or more of all newly recognized taxa do stem from new discoveries in the field however: this goes for the saola and other Vu Quang hoofed mammals, dingiso, the multiple new Amazonian primates discovered by Mark van Roosmalen and colleagues, all the new SE Asian rodents, and - among cetaceans - Perrin's beaked whale and the Lesser beaked whale. Specimens of Omura's whale were recognized as new when first obtained in 1978, but it took a while for the material to be properly studied and published.

On placodonts, I'll try and find you the relevant pdfs. I'm a great fan of Janis' work on hoofed mammals, though I never enjoy what she says at conferences about bird origins...

The TV newspiece I saw on this held another point of interest - at one point it mentioned, IIRC, that the Bornean clouded leopard was one of "53 new species of animal described from Borneo in the past year". As an invertebrate worker, my response to this was "Only 53?". My experience is that most people are genuinely amazed to learn just how many undescribed species there are out there (whatever your preferred 'species' concept).

... except that the Ostrom quote didn't originate with Ostrom. I think you probably got that from me, so sorry for the misinformation. I was told that my attribution was wrong in the review that rejected a paper that quoted. Having now looked into it a little, it's clear that it's been around a long while, and attributed to various people, including Art Cronquist and G. G. Simpson. But the oldest attribution I've seen for it is Regan 1926, where it's apparently stated as "a species is what a competent taxonomist says it is" and that seems to be correct.

Regan, C. T. 1926. Organic evolution. Report of the British Association for the Advancement of Science. 1925, 75-86.

Mike: thanks for the comment. Sorry, I knew that the quote didn't originate with Ostrom; rather, that various people have used it through the years. However, on the DML Tom Holtz once mentioned Ostrom's use of it, and I conclude that most of the people who read this blog will think of it as 'that Ostrom quote'. I recall Simpson using it (I think in The Structure of Evolutionary Theory), but didn't know that it originated with Regan. Thanks for the extra information.

"a species is what a competent taxonomist says it is"

"A language is a dialect with an army and a navy."

By David Marjanović (not verified) on 19 Mar 2007 #permalink

Nice blog covering many bases, Darren. Thank you & looking forward to reading more. :)
Referenced by 'Pharos' on , and added to by myself, there.
*nods* diardii should be retained rather than diardi, IMO, even though the first illos. for Diard's cat (from Java, not Sumatra or Borneo, of course) in the UK were of Marbled Cats, prior to that species actual "discovery" several years later. Those have long been questions of mine as to exactly why Cuvier pinned diardi(i) to Java and where that type specimen is... I have access to neither the original volume nor that specimen, alas.
I loved Guggisberg's comments relating to later authors asserting the primarily arboreal nature that "nobody remembered what Raffles and Jerdon had written". How often do non-fieldworkers actually bother to go back to primary sources, check out the original species descriptions, type specimens, etc., rather than merely repeat "received knowledge"?

By David Witts (not verified) on 19 Mar 2007 #permalink

I think Clouded Leopards are really cool, and to find out that there has been a new species found is really cool. I think they are one of the most coolest animals ever.

I meet N. diardi in jungle west java

By edi junaedi (not verified) on 28 Jul 2009 #permalink

Edi: It's amazing that they can survive in a place as densely populated as west Java, and astonishing that you got to see one. How did it happen?

By Nathan Myers (not verified) on 28 Jul 2009 #permalink