If anything should be clear from the range of creatures that I write about at Tet Zoo - think caecilians, borhyaenoids, imaginary giant owls and rhynchosaurs - it's that there's an almost infinite amount of technical information on obscure creatures 'locked away' in the technical literature. Among the most remarkable of mammals are, without doubt, the winged cats or pantheropterygines, yet for all their fame and notoriety, most of the information on these creatures has remained widely scattered in the literature and a good synthesis is absent.
Winged cats were first brought to scientific attention in 1891 when Richard Lydekker named Pantheropteryx anglicus for the famous London Zoo specimen. Known to her keepers as Minxy Cottonsocks, she'd been discovered in the Banister, Walton & Co. builder's yard in Trafford Park, Manchester, in 1880, and then sent to London Zoo [Minxy is shown above]. Though Minxy became a popular attraction (well known for her fantastic 15-metre leaps), the deaths of several keepers at her vicious claws made the zoo keen to dispense with her. In 1882 she was purchased by P. T. Barnum at the same time as Barnum obtained Jumbo the elephant; Jumbo and Minxy were actually close friends, and when Jumbo was killed by a train on arrival in New York, Minxy went into mourning and hid for weeks in a pigeon loft. After the sale of Barnum's exotic cat collection in 1889 (Cherrio the famous cherry-coloured cat, Manglox the five-legged leopard and Parsley the vegetarian lion were also included in the lot), Minxy ended up back at London Zoo, and here she lived until her suspicious death in 1890.
P. anglicus specimens had, of course, been known long prior to the publication of Lydekker's paper (Lydekker 1891) - Athanasius Kircher famously illustrated one as early as 1667 [his illustration is shown here] - but they'd generally been dismissed as one-off freaks or even as hoaxes. Well, ha ha ha, how we laugh at those naÃ¯ve suggestions now. A remarkable succession of fossils, and the discovery that the dropgorgons of ancient Eurasia are members of the pantheropterygine clade, have shown that winged cats are one of the most remarkable and divergent lineages within the carnivoran radiation.
Winged cats have, of course, gotten some attention in the popular media (who can forget the 13-part TV series Skypuss!), but the full story behind the palaeontology, developmental genetics and molecular phylogeny of Pantheropteryginae has gone unreported outside of the technical literature. Karl Shuker has provided a series of extremely useful reviews of winged cat history (e.g., Shuker 2008), but his work is marred by his interpretation of winged cats as normal members of the domestic cat lineage afflicted by a hypothetical condition dubbed feline cutaneous asthenia (FCA) (see also Patterson & Minor 1977). The FCA hypothesis has become the bane of research on extant winged cats, allowing some misguided animal carers to surgically remove the wings of their cats. I'd compare this to snipping the wings off angels and cherubs, except those things don't exist.
Pantheropteryx = surprisingly anachronistic and conservative!
It is rather ironic that P. anglicus was the first winged cat taxon to be discovered, as phylogenetic analyses show that it represents one of the oldest divergences within the winged cat clade (Ramsburger et al. 2001) [simplified version of the Ramsburger et al. phylogeny shown above]. Furthermore, it belongs to a strikingly conservative lineage and seems to have undergone little change over its
30 60 many millions of years (Graur & Martin 2004) of evolution. The discovery of additional pantheropterygines has demonstrated that the wings of Pantheropteryx are proportionally small compared to those of other species, and are only stiffened internally by cartilage. Indeed, P. anglicus is so similar to true cats of the domestic cat lineage that members of its species have (when mutilated or injured) often been mistaken for feral domestic cats.
Some experts even suspect that the type specimen for the domestic cat Felis catus - Carl von LinnÃ©'s pet moggy Maximilian III - might really be a P. anglicus specimen with cropped wings (for a review of LinnÃ©'s well documented and rather sickening obsession with cropping the wings off winged cats, see French (1980)) [von LinnÃ© and Maximilian III shown in the adjacent image; both pics from wikipedia]. Kitchener (1988) proposed that, if Maximilian III does turn out to be a mistakenly identified P. anglicus specimen, the International Commission on Zoological Nomenclature (ICZN) will be petitioned to elect a new type specimen that actually really does belong to F. catus. Brunet & McGee (1990) argued that it might be quicker to circumvent the slow, tortuous ICZN route, and that both Maximilian III, and the complete published works of Linnaeus, should be discarded in a ceremonial bonfire. An early plan to analyse Maximilian III's DNA was scuppered when Lewis (1999) pointed out that LinnÃ© used distilled flying cat oil to embalm his taxiderm specimens. I asked a colleague how widespread the use of flying cat oil was in the 18th century taxidermy business. "Pretty widespread", he said.
Anatomy of a cat wing - - oops, I mean a cat AIDO
For some time the mechanism behind the evolution of what appeared to be a third set of limbs in a tetrapod was regarded as an irresolvable enigmatic mysterious uber-problem. Observations of winged cats in captivity and in the wild show that they do use the wings to glide; they can actually increase the length of a leap by holding the wings outstretched (Femes et al. 1991).
Because the use of the term 'wing' has - to some - implied homology with limbs, some workers have argued that we really should use a different name for the structures. Verbosio (2002) proposed the term aerodynamic integumentary dorsal organs, or AIDOs, for the 'wings', in order to circumvent this issue. Cartilaginous rods within the AIDOs provide stiffness and, combined with the stiff, feather-like organs dubbed feloplumous integumentary structures (or FISs), produce an aerodynamic effect. The wing cartilages represent neomorphs and are known variously as AIDO spar sections or AIDOSSs and neomorphic cartilaginous rods or NCRs. A big question remains: did the AIDOs and their associated FISs evolve first (perhaps for use in display, intimidation or thermoregulation), with the AIDOSSs and NCRs being opportunistic structures that evolved later on (if this is true, then the role of AIDOs as aerodynamic organs could be an exaptation)? Or... were AIDOSSs and/or NCRs there from the start, in which case AIDOSSs, NCRs and FISs drove the evolution of AIDOs all along? There's a load of evo-devo literature on this but it bores me to tears so I don't want to talk about it [P. anglicus AIDO shown in the adjacent image; Dropgorgon AIDO shown below].
Flapperailurus shows that AIDOSSs and NCRs had appeared within the AIDOs by the Pliocene (Verbosio (2002) provided yet another new term - neomorphic ossified rods or NORs - for these), and it's inferred that simple bones like those seen in this taxon became elaborated over time, eventually giving rise to the branched NORs present in the wings of Roflailurus, Slooflirpapuss and the dropgorgonids.
The remarkable dropgorgonids
Without doubt the most remarkable pantheropterygines - and arguably the most remarkable of all mammals - are the dropgorgonids (often just called dropgorgons) of Pliocene and Pleistocene Eurasia and Africa. Of the several known taxa, Tnihevissam of Romania, Chinnychinpuss of China, and Dropgorgon (known from several species that occurred across Europe, Asia, the Middle East and northern Africa) seem to form a morphological sequence in which body size increased, the anterior dentition became hypertrophied, the cranium became increasingly bulbous and the brain more enlarged, and the limbs became more robust (KurtÃ©n 1956). The culmination of this trend was the incredible, facultatively bipedal Dropgorgon, the surreal skull of which was originally assumed to be pathological (Dawkins & Sanford 1869) prior to the discovery of additional specimens [skull shown here, from Dawkins & Sanford (1869)].
Dropgorgon giganteus was a truly formidable beast. A large male would be more than 3 m tall when standing bipedally, 5 m long, have a wingspan exceeding 6 m, and weigh over 200 kg. The frequent occurrence of its remains in caves and steep-sided ravines indicate that it really inhabited such places, and chewed bones found in these dropgorgon 'lairs' show that it ate goats and humans. Dawkins & Sanford (1869) surmised that it dropped down on prey from above in drop-bear-like style, and this inferred behaviour explains the name they chose for it (incidentally, this demonstrates that drop-bears were known to Europeans long prior to the 21st century!). Dawkins & Sanford (1869) included D. giganteus among the felids, but a link with the other pantheropterygines wasn't realised until Chinnychinpuss gansuensis from China and the good Dropgorgon postcranial remains from Turkey were discovered in the 1980s.
Radiocarbon dating has shown that D. giganteus survived in central and eastern Asia well into the Holocene and to as recently as the 5th century. The relatively recent existence of this enormous, monstrous, long-winged predator has not been lost on folklorists and classicists who have drawn attention to the many similarities this animal has with griffons (Griffen 1997). We're never really going to know whether the griffons of mythology really were based on dropgorgons, but I agree that this is a very real possibility. Griffen (1997) also drew attention to the strong similarity that dropgorgons have with various creatures from African folklore, notably the Sasabonsam from Ghana, and the Kikiyaon and Guiafairo of Senegal. If this is correct, it raises the exciting possibility that one or more dropgorgon species awaits discovery in the zooarchaeological record of tropical Africa.
Due to their remarkable morphology and behaviour, dropgorgons and other pantheropterygines have been important animals in the history of zoological discovery, and their story is an incredible one. Where else could you read about a controversial zoo specimen from the 19th century that once belonged to P. T. Barnum, about the evolution of entirely novel wing-like organs, and about obscure monsters from African folklore? Yet I've hardly scratched the surface of the pantheropterygine literature; hopefully this brief introduction has piqued your interest.
Refs - -
Brunet, J. J. & McGee, T. 1990. Thinking the unthinkable: a neat and tidy proposed resolution to an inextricable problem of historical taxonomy concerning the first winged cat. Proceedings of the American Winged Cat Society 67, 45-60.
Dawkins, W. B. & Sanford, W. A. 1869. The British Pleistocene Mammalia. Part III. British Pleistocene Felidae. Palaeontographical Society, London.
Femes, B., Dorchap, E. & Gerwih, H. 1991. They seem to glide. Are there aerodynamic effects in leaping winged cats? Zeitschrift fur Morphologie und Catologie 79, 37-385.
French, G. W. 1980. Carl von LinnÃ© was one sick mother: a secret obsession with cat mutilation revealed through pictograms and retrogressive astrology. The Smudgener 1, 5-8.
Graur, D. & Martin, W. 2004. Reading the entrails of chickens: molecular timescales of evolution and the illusion of precision. Trends in Genetics 20, 80-86.
Griffen, M. 1997. The discovery of dropgorgonid cats brings an end to the griffon mystery. Pokemonia 34, 38-46.
Kitchener, B. 1988. What a sodding mess: the status of the type specimen of Pantheropteryx anglicus and why YOU should care. Taxonomy, Taxonomy, sigh, Taxonomy 250, 129-140.
KurtÃ©n, B. 1956. A case of Darwinian selection in dropgorgon flying-cats. Evolution 10, 412-416.
Lewis, O. 1999. Carl von LinnÃ© used distilled flying cat oil to embalm his taxiderm specimens. Taxidermy Now 46, 10-12.
Lydekker, R. 1891. On an aeluroid carnivore from the United Kingdom, notable for its wings. That is correct, I said wings. It is a freakin' winged cat for crying out loud. This is one paper that you really should read, you know. Proceedings of the Zoological Society 20, 883-900.
Patterson, D. F. & Minor, R. R. 1977. Hereditary fragility and hyperextensibility of the skin of cats. A defect in collagen fibrillogenesis. Laboratory Investigation 37, 170-179.
Ramsburger, D., Slice'n'dice, M. & Groovy, E. 2001. Holy shit!! A phylogeny for winged cats reveals amazing stuff, hence this two-page paper. Science 308, 1112-1113.
Shuker, K. P. N. 2008. Dr Shuker's Casebook. CZF Press, Bideford.
Verbosio, H. 2002. Patterns of variation in the 'wing' morphology of dropgorgonid carnivorans: developmental constraints, size, and phylogenetic history. Journal of Experimental Zoology (Mol Dev Evol) 303B, 135-147.
I think P. anglicus should have more teats (probably 8) instead of a pair of breasts! 0:-)
Oh My God. I thought I knew a lot about fossil mammals but I have never heard of this group before. I note that they are excluded from Turner and Anton's book _The Big Cats and their Fossil Relatives_.. is this because these authors don't regard pantheropterygines as cats??
Its a "well known fact" that Alan Turner and the indomitable Ramsburger had a large disagreement about the phylogenetic placement of Pantheropteryx. This boiled over into personal animosity, which resulted in the exclusion of the taxa from "Big cats..."
Its a shame, as I would love to see some reconstructions by Anton.
"I asked a colleague how widespread the use of flying cat oil was in the 18th century taxidermy business. "Pretty widespread", he said."
The "best pers. com." ever!
the dropgorgonids (often just called dropgorgons)
If they're extinct, couldn't they also be called dropdeadgorgonids?
The _Dropgorgon_ skull always struck me as highly strange. As Dawkins and Sanford (19869) describe, perhaps the strangest thing is that the zygomatic arch reach well anterior to the orbit, resulting in the orbit positioned next to the posterior part of the arch. VERY odd! Perhaps unique? Then again, some rodents do a similar sort of thing and we already have reason to think that dropgorgons had a hystricomorphous arrangement of jaw muscles.
I very pleased to see this article!!
Wow, thanks for that review! The chupacabras is often displayed as having wings, is there any possibility that it is actually a Caribbean dropgorgonid?
However, notice how AIDOSS 1 would have been useless without AIDOSS 2, 3 and 4. Clearly irreducibly complex; clearly evidence for Intelligent Design!
We need to do more to control Pantheropteryx populations. Feral cats are bad enough on native bird populations but when Pantheropteryx are introduced as well, avian extinction follows apace.
I adore the lovingly crafted reference list. It's clear it took you time to find some of those obscure citations, but (as everyone knows from Wikipedia) comprehensive citations are absolute proof of veracity.
I'm getting pretty bloody sick of this: every survey of the dropgorgonids seems to parrot this 200 kg mass estimate for Dropgorgan uncritically, even though it wasn't based on any kind of modelling. Even so simple a technique as isometrically scaling up from a 100 kg, 2 m tall man gives a mass estimate of 338 kg for Dropgorgon, and in truth its robust proportions suggest a mass closer to 500 kg. Darren, I know maths isn't your strongest area, but you really should know better than to propagate the older, lower figure.
It's a generally good write up, but I'm disappointed you didn't further explore the contrasting origin of wing hypotheses; personally I have always favored the ground-up scenario.
love, love, love the bibliography!!!
Isn't it possible that the dropbears were named after/in honor of the dropgorgons?
and thank you for posting such a lengthy and educational post about a clade that gets far too little academic attention.
have nice days and be well.
Then again, some rodents do a similar sort of thing and we already have reason to think that dropgorgons had a hystricomorphous arrangement of jaw muscles.
As did... who was it? Thylacosmilus?
love, love, love the bibliography!!!
Graur & Martin (2004) is not cited in the text. But it fits the rest of the bibliography so well that you just had to include it, I think. :-)
Unbelievable. . . Jumbo was killed by a train? The other stuff I knew. I actually own a copy of the article "Seriously, I know what a bat looks like, and I'm certain it wasn't a squirrel. What kind of bat purrs?" The Jumbo thing is quite a revelation.
Though i confess i'm a little dubious about the picture at the top of the article. Based on the photo alone, i'd consider it an image of the equally rare (if not more so) 3-tailed cat.
Wait a minute, so all these "feral cats" throughout the islands of the world are actually secondarily flightless pantheropterygines that have lost their AIDO's? Wow, too long have I been blind. Here I was thinking that they were brought on ships. I'll never listen to Wikipedia again.
This thread is like memegenesis. Although I guess they could all just be ropens....
Whoa. So by 1990 the Proceedings of the American Winged Cat Society was up to 67 volumes. Man, I have some catching up to do. Thanks for the intro to the lit Darren.
You have an error in Femes & al. Z. Morph. Cat. was founded by Leibniz (a great helofelophile, himself, as it happens - and a 'kitten'* of his favourite floggie - Differentiala - was given by him to a young Kant. Some even claim that the purring, little Dingensig was an inspiration to him in his work) so obvious by 1991 it'd clocked up more issues. You want the int. ed. - as should also be obvious from the title.
*Yes, I know that technically they're pups, but I refuse to bow to pressure. You won't find my calling Ununbium anything but Copernicum either.
Shouldn't this have been posted in http://scienceblogs.com/hexapodzoology/?
If you ask me, hexapodia is the key insight.
Ah, this makes Droparctos venatokoala just another example of convergent evolution from Australia!
Have you never read B. Ueberschlau's magnificent "Prodromus zu einer systematischen Erfassung der gefluegelten Katzen nebst einer Synopsis ihrer Beutethiere und Bemerkungen ueber Flugsaurier" (Verlag von Petrus Schueler, Unterambach, 1812)? Ueberschlau already introduced (the etymologically more correct and still valid nomen)"Pterailurini" for winged cats.
Kircher was confused: In his "Mundus subterraneus" he repeatedly refers to "Catus volans" but I see little evidence that he was actually referring to Pantheropteryx anglicus. Indeed, Ueberschlau thought that Kircher frequently confused bats and winged cats - a common problem after significant absorption of winged cat oil due to handling of taxidermy specimens in his Vatican museum.
By the way, Lizzardo, Fu and Peters (PLoS One, in press) have recently argued that Dropgorgon was a marginocephalian, presumably impressed by the nuchal shelving.
"Oh My God. I thought I knew a lot about fossil mammals but I have never heard of this group before. I note that they are excluded from Turner and Anton's book _The Big Cats and their Fossil Relatives_.. is this because these authors don't regard pantheropterygines as cats??"
I heard someone found that pantheropterygines form a clade with barbourofelids in one cladistic analysis. Turner and Anton must be following that interpretation of the pantheropterygines.
April fools? Haha.
"Brunet & McGee (1990) argued that it might be quicker to circumvent the slow, tortuous ICZN route, and that both Maximilian III, and the complete published works of Linnaeus, should be discarded in a ceremonial bonfire." ... awesome.
And the paper titles in the references are great too... especially "On an aeluroid carnivore from the United Kingdom, notable for its wings. That is correct, I said wings. It is a freakin' winged cat for crying out loud. This is one paper that you really should read, you know." (Hmm, I wonder if I could find a copy of that one...)
Interesting, I always thought the 'wings' were dorso-hyperextendible flattened supernumery ribs attached, anchored and powered by hyperpneumatic vertebral air sacs. Whoops!
Now that it's April 2nd, I feel obliged to mention that Graur & Martin (2004) is real. It's about the fact that many molecular dating studies published till then had made horribly naive assumptions (up to and including circular logic in some cases!) and had thus come to laughably unreliable conclusions.
B. Ueberschlau's magnificent "Prodromus zu einer systematischen Erfassung der gefluegelten Katzen nebst einer Synopsis ihrer Beutethiere und Bemerkungen ueber Flugsaurier" (Verlag von Petrus Schueler, Unterambach, 1812)
I heard someone found that pantheropterygines form a clade with barbourofelids in one cladistic analysis.
Ha! That tells us nothing! We don't even know whether barbourofelids are carnivorans!!! A recent phylogenetic analysis that included them was based on a matrix with eleven characters, and IIRC not even all of these were parsimony-informative.
Thanks to all for comments, much very appreciated - and I'm pleased that so many of you could contribute sensibly to the discussion (rather than pretend that this was a hoax of some kind). A few select responses...
-- Mike (comment 11): it's true, I blindly followed previous estimates in the literature. An original version of this article included some discussion of dropgorgonid mass (they have usually been regarded as light for their size, owing to their low sg), but I lost a huge chunk of text in amusing yet frustrating circumstances. American lions have been estimated by some to be as heavy as 500 kg (Anyonge 1993), so it would ordinarily seem conceivable that Dropgorgon giganteus would match or exceed even this.
-- David (comment 15): hystricomorphy has been proposed for barbourofelids (Naples & Martin 2000). Based on the paper, this looks pretty reasonable to me, but I have no experience with these animals and would be interested to know what specialists think of it.
-- Hans (comment 23): all I say can is that I bow to your superior knowledge of the literature.
For previous articles that, entirely coincidentally, were also published on April 1st, please see At last, the rhinogradentians (part I), Amphisbaenians and the origins of mammals and Recently discovered late-surviving carnivorous reptiles probably explain the origin of the dragon myth.
Refs - -
Anyonge, W. 1993. Body mass in large extant and extinct carnivores. Journal of Zoology 231, 339-350.
Naples, V. L. & Martin, L. D. 2000. Evolution of hystricomorphy in the Nimravidae (Carnivora; Barbourofelinae): evidence for complex character convergence with rodents. Historical Biology 14, 169-188.
So which of the references are ... um... accessible?
I am surprised that the well-known feud between Lydekker and Seeley didn't manifest itself in the field of pantheropterygines given the latter's interests in flying amniotes.
Loved it, much better than last years April Fools article!
Now you just need to follow it up with an article on real "winged cats", though it would be hard to out do the one at
Distilled flying cat oil? Hilarious! :)
I made it through nearly the entire post before I caught myself.
(I didn't catch on until the dragon skull, I'm ashamed to admit.)
Disturbing...kind of like the mob mentality at play in a more passive way but disturbing none the less. That's what modern technology does to you - fight it people!
A correction: the title of the Kitchener paper is:
Kitchener, B. 1988. What a sodding mess: The status of the type specimen of Pantheropteryx anglicus and why should YOU care.
Rodents and carnivores have different functional constraints: I wouldn't think porcupines need anything like the extremely wide gape angle of sabertooth "cats" (quotation marks to include Barbourofelines and Thyylacosmilus), so the convergent evolution of the hystricomorph contition in Barbourfelinae comes as a REAL surprise: the Naples & Martin paper, however, is NOT an April Fool's joke.
Darren got you twice. Winged cats belong to the pseudo mammal group, which are archosaurs and thus reptiles. The winged cat is a case of convergent evolution, being a member of the false mammals, which are most closely related to the wyrms; which are archosaurs like crocodiles, dinosaurs, and dragons.
The winged cat is most closely related to the griffin, and has be called the little griffin at times. Like the griffin they are related to the Sphinx and to the lung wang (earth dragon), but less closely related to the pegasus. The close resemblance to felines is thanks to similarities in lifestyle and habitat.
As with other false mammals the flying cat undergoes pseudo-lactation, producing a milk like substance from two glands on the chest. The flying cat, being descended from a common ancestor of the griffin, is a live bearer, having one kit at a time. The third set of limbs (also found in dragons as well as the pseudo mammals and the mammalian manticore) is now known to arise from a mutation allowing for the expression of the genes governing forelimb development a second time.
And that's my contribution to Darren's thesis. Do I get co-author credit in the eventual publication in the scientific literature?
I await the first LOL-pantheropterygine captioned pictures.
"OH HAI. I maded you a nice roasted poisson d'avril, and sent it par avion d'chatte. Did you get it?"
I noticed the mention of Roflailurus. Is it the same animal vernacularly known as the LOLcat ? It would be interesting to elaborate on its genealogical affinities, and on the origins of its typographical peculiarities.
Someone ought to inspecty this unique specimen...