Pumas of South Africa, cheetahs of France, jaguars of England

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In the previous article (required reading) we looked at European leopards. But the leopard wasn't the only big spotted Panthera species that lived in Europe during the Pleistocene: it was joined by a second, far less well known animal: Panthera gombaszoegensis (originally Leo gombaszoegensis Kretzoi, 1938). This cat seems to have been very jaguar-like and in fact the name 'European jaguar' is often used for it. In fact, it may actually be a jaguar - that is, a member of the species Panthera onca - and some cat experts classify it as an extinct Panthera onca subspecies (Hemmer et al. 2001, 2003, 2005) [note however that this is not universally accepted: Agustí & Antón 2002, O'Regan & Turner 2004, O'Regan et al. 2002)].

The European jaguar makes its first appearance in the fossil record about 1.5 million years ago, where it's recorded from Italy, and it then persisted into the Middle Pleistocene, at which time it's known from Germany, Spain, France and Westbury in England: in fact, some of the best fossils of this subspecies come from the cavern site of Westbury-Sub-Mendip in Somerset (Bishop 1982) [see map below, from here]. European jaguars and leopards lived alongside one another during the Middle Pleistocene: remains of both species have been reported to occur at the same stratigraphic levels in the Czech Republic, France and Germany (Agustí & Antón 2002, García & Virgós 2007).

i-f84216e228e93cee89023cd0e7856896-Panthera gombazoegensis range.jpg

The idea of jaguars in Europe might be surprising given that we generally think of this cat as a South American species, and one that also occurs in Central America and in some southern parts of the USA. But this present distribution is a historical artefact: jaguars were originally Old World cats, originating in Eurasia or Africa, and later moving across eastern Asia and, at the end of the Pliocene, crossing Beringia into the Americas (Kurtén & Anderson 1980, Hemmer et al. 2001). Fossils from the Late Pliocene or Early Pleistocene of India, originally identified as early leopards, have more recently been suggested to be jaguars (Turner 1990); both Panthera toscana and Panthera gombaszoegensis of the European Pliocene and Pleistocene are best regarded as jaguar subspecies; and it is the North American Pleistocene subspecies P. onca augusta that gave rise to living jaguars (Hemmer et al. 2001). Pleistocene jaguars were not only bigger than living ones (15-20% bigger or more), they also had proportionally longer limbs and feet. Kurtén (1973) and others suggested that the particularly short, stocky limbs of recent and extant jaguars demonstrate increasing specialisation for life on broken ground and hilly terrain.

During the Pleistocene, jaguars seem not to have occurred in the same areas as lions, possibly because the two avoided competition: in Pleistocene North America, jaguars are absent from Rancho La Brea for example (where lions were abundant), but numerous in Florida, Texas and Tennessee (where lions were scarce or absent) (Kurtén & Anderson 1980, Turner & Antón 1997). Genetic data indicates that living jaguars originated in northern South America during the Middle Pleistocene, with even living North American jaguars being descended from these South American ones (Eizirik et al. 2001). This is similar to the pattern inferred for modern pumas: they also seem to have become extinct in North America prior to the reinvasion of the region by South American founders (Culver et al. 2000).

The pumas of Europe (!), Asia (!!) and Africa (!!!)


Most people know that Europe was formerly home to leopards and lions, everybody knows that lynxes inhabit Europe... but very poorly known is the fact that Europe was also inhabited by pumas during the Pleistocene. Pumas today are of course exclusively American but a European species from the Pleistocene is now regarded as a European puma. This cat has had a confusing history. It was originally described in 1846 as Felis pardoides (later becoming Panthera pardoides) and was thought to be a leopard-like species - in fact it was still being described as leopard-like as recently as the 1980s. It has sometimes been called Owen's panther, after Richard Owen, its describer. In 1954 what was thought to be a totally distinct species, Panthera schaubi - Schaub's panther - was described from the Pleistocene of France [reconstruction and life restorations of Schaub's panther shown in adjacent image. By Velizar Simeonovski].

A reanalysis of this species published in 1965 showed that it wasn't a member of the genus Panthera but was instead far more similar to pumas, and Schaub's panther was now given its own, new genus: Viretailurus Hemmer, 1965. Hemmer (1965) drew attention to puma-like features in the skull, and a few authors were to later remark on this similarity: Turner & Antón (1997), for example, noted of Viretailurus that 'suggestions have included a possible link with the American puma' (p. 63). Sotnikova (1976) was the first worker to bring specific attention to the fact that Viretailurus was probably allied to Puma, and also to note that the Viretailurus fossils from France probably represented an animal closely allied to (or the same as) the 'Felis sp.' fossils from the Pliocene or Early Pleistocene of Beregovaya and Shamar in northern Mongolia.

Well, Viretailurus is indeed puma-like and in fact it's so puma-like that the most recent work on this cat (Hemmer et al. 2004), has shown that Viretailurus actually is a puma, and is the same thing as Owen's panther (Panthera pardoides). So... Viretailurus schaubi and Panthera pardoides are the same thing, and are part of the genus Puma, so the correct name is Puma pardoides.

What we know of Puma pardoides suggests that it was similar in appearance to modern pumas - certainly its short-faced skull is puma-like [see picture at very top], and with an estimated mass of 40-45 kg, it was similar in size to typical modern pumas. Old World puma records are now known from the Transcaucasian region of central Asia and Mongolia, and Hemmer et al. (2001, 2004) suggested that Tanzanian and South African fossils from the Pliocene, identified as leopard, might actually be puma remains: 'it now emerges from the haze of the highly fragmentary nature of the specimens, that the Pliocene African cats originally identified as leopards are not related to Panthera pardus, but rather to Puma. These African animals seem to foreshadow the later Eurasian pumas' (Hemmer et al. 2004, p. 220) [map below, from here, shows Eurasian range of Puma pardoides].

i-1052fcf29123319f6dd75e44db88bb20-Puma pardoides distribution.jpg

One of the great mysteries of the American puma has always been the fact that, in the fossil record, it appears suddenly about 40,000 years ago in the Late Pleistocene and yet doesn't have an obvious American ancestor. The discovery of pumas in eastern Asia, and of older puma records in Europe and Africa, has now led to the suggestion that pumas originated in Africa, were widespread across the Old World during the last couple of million years, and crossed the Bering land-bridge during the Late Pleistocene to invade North America, then giving rise to the American puma Puma concolor (Hemmer et al. 2004). When looked at within the broader context of the phylogeny of the puma clade however, this scenario might be problematical.

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Johnson et al. (2006) found that cheetahs, jaguarundis and pumas form a clade: the fossil American cheetah Miracinonyx is also part of this group, and apparently the sister-taxon to Puma* (Barnett et al. 2005). Given that jaguarundis, American cheetahs and American pumas are all, well, American, it is more parsimonious to posit an American ancestry for the clade, with Old World cheetahs and Old World pumas being invaders from the Americas (Johnson et al. 2006). However, the outgroup to the puma-cheetah clade (the lynx clade) is of ambiguous biogeographical origin (Johnson et al. regarded lynxes as American, but they only included the four extant species and not the additional fossil ones). Furthermore, adding Puma pardoides results in one additional 'Old World score'. So, for Old World we have (1) possibly the outgroup (it's lynxes in Johnson et al.'s phylogeny), (2) Acinonyx and (3) Puma pardoides. For the Americas we have (1) Herpailurus, (2) Miracinonyx and (3) Puma concolor. So, neither biogeographical option is definitely more parsimonious. Maybe members of this clade hopped in and out of the Americas several times (see Stewart & Disotell 1998: 'Primate evolution - in and out of Africa', for a nice discussion of this sort of thing) [adjacent picture is Mauricio Antón's life restoration of the American cheetah Miracinonyx inexpectatus. I found this picture on the web - someone has naughtily scanned it from Turner & Antón's The Big Cats and Their Fossil Relatives].

* Puma sensu stricto, as Johnson et al. (2006) found the jaguarundi Herpailurus yagouaroundi close enough to Puma to be included in the same genus.

It might not surprise you to hear that cheetahs were present in Pleistocene Europe as well as pumas: during the Early and Middle Pleistocene, the giant cheetah Acinonyx pardinensis (about 50% bigger than living cheetahs) was found in Germany, France, and also in China and India. European cheetahs occurred alongside jaguars and leopards at some Middle Pleistocene localities, and it is possible that competition between the three contributed to the cheetah's decline (García & Virgós 2007). And I will have to finish there, but I need to do more on cheetahs at some time, not least because I've been promising an article on onzas for some time now. One Day I Will Deliver.

More soon.... (and, coming very soon: aquatic proto-people).

Refs - -

Agustí J. & Antón, M. 2002. Mammoths, Sabertooths, and Hominids. Columbia University Press, New York.

Barnett, R., Barnes, I., Phillips, M. J., Martin, L. D., Harington, C. R., Leonard, J. A. & Cooper, A. 2005. Evolution of the extinct sabretoths and the American cheetah-like cat. Current Biology 15, 589-590.

Bishop, M. J. 1982. The mammal fauna of the early Middle Pleistocene cavern infill site of Westbury-Sub-Mendip Somerset. Special Papers in Palaeontology 28, 1-108.

Culver, M., Johnson, W. E., Pecon-Slattery, J. & O'Brien, S. J. 2000. Genomic ancestry of the American puma (Puma concolor). The Journal of Heredity 91, 186-197.

Eizirik, E., Kim, J.-H., Menotti-Raymond, M., Crawshaw, P. G., O'Brien, S. J. & Johnson, W. E. 2001. Phylogeography, population history and conservation genetics of jaguars (Panthera onca, Mammalia, Felidae). Molecular Ecology 10, 65-79.

García, N. & Virgós, E. 2007. Evolution of community composition in several carnivore palaeoguilds from the European Pleistocene: the role of interspecific competition. Lethaia 40, 33-44.

Hemmer, H. 1965. Studien an "Panthera" schaubi Viret aus dem Villafranchien von Saint-Vallier (Drôme). Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen 122, 324-336.

- . & Kahlke, R.-D. 2005. Nachweis des Jaguars (Panthera gombaszoegensis) aus dem späten Unter-oder frühen Mittelpleistozän der Niederlande. Deinsea 11, 47-57.

- ., Kahlke, R.-D. & Keller, T. 2003. Panthera onca gombaszoegensis (KRETZOI, 1938) aus den frühmittelpleistozänen Mosbach-Sanden (Wiesbaden, Hessen, Deutschland) - Ein Beitrag zur Kenntnis der Variabilität und Verbreitungsgeschichte des Jaguars. Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen 229, 31-60.

- ., Kahlke, R.-D. & Vekua, A. K. 2001. The jaguar - Panthera onca gombaszoegensis (Kretzoi, 1938) (Carnivora: Felidae) in the late Lower Pleistocene Akhalkalaki (South Georgia: Transcaucasia) and its evolutionary and ecological significance. Géobios 34, 475-486.

Johnson, W. E., Eizirik, E., Pecon-Slattery, J., Murphy, W. J., Antunes, A., Teeling, E. & O'Brien, S. J. 2006. The Late Miocene radiation of modern Felidae: a genetic assessment. Science 311, 73-77.

Kurtén, B., 1973. Pleistocene jaguars in North America. Commentationes. Biologicae Societas Scientiarum Fennica 62, 1-23.

- ., Anderson, E., 1980. Pleistocene mammals of North America. Columbia University Press, New York.

O'Regan, H. J. & Turner, A. 2004. Biostratigraphic and palaeoecological implications of new fossil felid material from the Plio-Pleistocene site of Tegelen, the Netherlands. Palaeontology 47, 1181-1193.

- ., Turner, A. & Wilkinson, D. M. 2002. European Quaternary refugia: a factor in large carnivore extinction? Journal of Quaternary Science 17, 789-795.

Sotnikova M. V.1976. The Upper Pliocene Carnivora of Central Asia. Izvestiya
Akademii Nauk SSSR, Seriya Geologicheskaya
11, 133-137 [in Russian].

Stewart, C.-B. & Disotell, T. R. 1998. Primate evolution - in and out of Africa. Current Biology 8, 582-588.

Turner, A. 1990. Late Neogene/Lower Pleistocene Felidae of Africa: evolution and dispersal. Quartärpaläontologie 8, 247-256.

- . & Antón, M. 1997. The Big Cats and Their Fossil Relatives. Columbia University Press, New York.


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The Puma pardoides reconstruction appears to have been done by Velizar Simeonovski, who has some art at the following link:


Aside from Acinonyx pardinensis, is there any possible evidence that like the leopard, the modern cheetah Acinonyx jubatus may have had a more widespread range in the Late Pleistocene? It's tempting to imagine cheetahs roaming the mammoth steppe of Eurasia, chasing saiga and gazelle (Are gazelles known from Late Pleistocene Europe?).

A provincial question - are any of these big cats (not counting lynxes) known from Scandinavia?

By Andreas Johansson (not verified) on 13 Mar 2008 #permalink

Pumas crossed to North America 40,000 years ago... But people didn't cross for another 20,000 years or more? Was this two separate appearances of the Beringia 'land bridge'? Or do I have the dates wrong?

Rosie Redfield: The Bering land bridge has been intermittently present through much of the Late Miocene, Pliocene and Pleistocene. Various groups of animals would have made the crossing at different times.

AFAIK, brown bear Ursus arctos is the only large animal species thought to have crossed over from Eurasia to North America at about the same time as humans; others like lion, steppe bison, moose, wapiti, grey wolf and woolly mammoth would have been somewhat more ancient immigrants.

You may have spelled the jaguarundi's scientific name wrong; "yaguarondi" and "yagouarundi" both appear in recent peer-reviewed articles, but it appears that "yagouaroundi" is the most commonly accepted form. I'm still dubious putting the species into "Puma", the supplementary data from Barnett et al. indicated it was rather divergent and Johnson et al. didn't include Miracinonyx.

When you say "% bigger", does that mean by linear measurement or by predicted mass? Saying that P. atrox is 1/3 as the size of P. leo could mean more than a doubling in mass (which appears to be the case) - but is the giant cheetah really that giant?

This might be the place to ask a question that involves pumas and cheetahs...

I read in one book that the cougar/puma/mountain lion is the only "other" cat that purrs. I read elsewhere that cheetahs do, too. Is purring primitive to the cats, or restricted to one branch? Is there a definitive list of purring species, and does it match a phylogenetic grouping?

Purring is usually described as expressing satisfaction, but my experience is that it means the cat wants something. Do we know anything about the social function of purring in the bigger cats that indulge?

By Nathan Myers (not verified) on 13 Mar 2008 #permalink

Many thanks to all for comments, and to links to Simeonovski's artwork. Here are responses to points raised so far..

-- prehistoric range of modern cheetah: I haven't yet read anything indicating that Acinonyx jubatus occurred outside of its historical range in the Pleistocene. Would love to hear otherwise (by the way, 'historical range' includes northern Africa, entire Middle East, Transcaspian region and India).

-- fossil felids from Scandinavia: you'd need to ask an expert on the region. So far as I can tell the area has only had lynxes and wildcats.

-- correct name for jaguarundi: I've also noted different spelling used in recent publications so checked original sources. The species name was (so far as I can tell) originally spelt yagouaroundi when coined by Geoffroy in 1803. It has frequently been mis-spelt since.

-- what exactly does '% bigger' mean? I have no idea, I simply copy other authors. Numbers mean - almost literally - nothing to me. I can no longer remember where the '50% bigger' figure came from, but this doesn't look inappropriate given that living cheetahs reach 60 kg, and giant cheetahs are said to have been as big as small lions (which would be about 120 kg). Feel free to look into this properly!

-- finally, I screwed up in saying that Johnson et al. regarded lynxes as Old World. In fact they didn't: with Canadian lynx and bobcat at the base of the lynx clade they regarded lynxes as American, but this fails to account for the fossils. Whatever, I've changed the text in the article to (hopefully) better reflect the ambiguity about this.


"There is good reason to assume that all species of the Felidae, with the exception of the six pantherine species Lion, Leopard, Jaguar, Tiger, Snow Leopard and Clouded Leopard, have purring."

G. Peters (2002) Purring and similar vocalizations in mammals. Mammal Review 32 (4),245271

Here's the link: http://tinyurl.com/2l7dyb

Apparently "purring" is a somewhat nebulous term that encapsulates a lot of similar vocalizations such as rumbling or growling, and this is where a lot of the controversy comes from.

Thanks, Alec! Following the link as far as it will go (i.e. to the US$39 paywall), I find that Viveridae also purr: civets and genets, in particular. Meerkats too?, I wonder.

By Nathan Myers (not verified) on 13 Mar 2008 #permalink

With regards to the movement of cats into and out of europe it really depends on whether you give more weighting to the fossil evidence or the phylogenetic/biogeographic evidence. Looking at johnsons paper it does seem more parsimonious to conclude that Acinonyx evolved from a stem American group and crossed back into the old world (this phylogeny wasnt available when we published on Miracinonyx). But if you look at the fossils you can see pretty early puma and cheetah like cats in europe before anything comparable in the americas, which would suggest an entry into the new world for the puma group and subsequent diversification.
Whats interesting to note is how contracted the jaguars range became after the pleistocene. The used to occur as far north as Nebraska and Washington state and even south to tierra del fuego before losing ground to the north and south in the holocene.

Holy crap, you're Ross Barnett! (of the Barnett et al. cited above everyone). I had no idea. Well, I'm honoured to have you as a site visitor :)

The pumas of Europe (!), Asia (!!) and Africa (!!!)

Surely that should be "The pumas of Asia (!), Europe (!!) and Africa (!!!)"?

Great post anyway. Sorts a lot of things out, and finally gives me the Johnson et al. reference (I was sure I had downloaded the article long ago, but I don't have it, and I didn't remember any part of the reference). :-)

Human immigration to the Americas seems to have been a complicated affair. It is clear now that the Clovis culture people were not the very first. There's a 17,000-year-old site somewhere in the eastern USA and even older ones in South America. Probably humans crossed at every opportunity in the last 50,000 years or so.

By David Marjanović (not verified) on 14 Mar 2008 #permalink

Nice post!

Always wondered, how reliable is distinguishing jaguar vs. leopard bones. Could there be misinterpretation?

Im just glad that someones read the papers! I read TetZoo every day!


Welcome to the gab fest. :)

On Purring.

The strictest description of purring I've seen says that purring is what you get when the animal can make his larynx vibrate while inhaling as well as exhaling. That requires a non-ossified hyoid bone. Lions et al have an ossifed hyoid, so they cannot, strictly speaking, purr.

As for way small cats purr, it depends on who or what they're purring for. Queens purr for their kittens because it calms them down. Domestic cats purr for their people because, from the cat's point of view, we purr to them and cats are courteous beasts.

BTW, here's a bit of video featuring a lion greeting old buddies he hasn't seen in over a year. :)

David Marjanovic wrote:
Human immigration to the Americas seems to have been a complicated affair. It is clear now that the Clovis culture people were not the very first. There's a 17,000-year-old site somewhere in the eastern USA and even older ones in South America.

There's a review article on the current state of knowledge on initial human settlement of the New World in the latest (14 Marh 2008) issue of "Science." The writers don't seem to have any confidence in dates much more than 16,000 years ago: still safely a few thousand before Clovis, but not as early as the dates (30,000+) for modern humans in Siberia would make one suspect. They seem to think the first settlers used boats a lot and went down the Pacific coast... so the remains of any REALLY early settlement might now be under water, given post-glacial sea-level rise.

By Allen Hazen (not verified) on 14 Mar 2008 #permalink

The strictest description of purring I've seen says that purring is what you get when the animal can make his larynx vibrate while inhaling as well as exhaling. That requires a non-ossified hyoid bone.

Are you sure? I can... well... growl when inhaling just like when exhaling, and my hyoids are ossified and fused in the middle (pers. obs.). Am I doing something else?


I'll check out the latest Science tomorrow, thanks!

By David Marjanović (not verified) on 16 Mar 2008 #permalink

David wrote:

Are you sure? I can... well... growl when inhaling just like when exhaling, and my hyoids are ossified and fused in the middle (pers. obs.). Am I doing something else?

I can't be sure, since I've never heard David growl (much less X-rayed or endoscoped his larynx while he was doing it), but I suspect it's his false vocal folds (AKA ventricular or vestibular folds) that are doing the growling.

Neat trick making them vibrate when you inhale, though!

By Nick Pharris (not verified) on 16 Mar 2008 #permalink

R.E. lions and Jaguars in North America; both are present at Rancho la Brea, although neither are common. Lions outnumber jaguars, but neither really occur in large numbers (certainly <50 lions and <10 jaguars). Puma and Homotherium also occur but are fairly rare. Jaguars are much more common in the South East during the late Pleisocence, but lions are also found there. There are good records of P. atrox and P. onca from Texas, Tennessee, and Florida among other places.

It is hard to think that big cats like lion, juguar lived in britain and the rest of eurasia in ice ages.

It is hard to think that big cats like lion, juguar lived in britain and the rest of eurasia in ice ages.

When prey is available, the predators will come. They have fur, they can stand the cold.

Interesting? or rubbish?

There was a paper in the Journal of Vertebrate Paleontology a few months ago that found no evidence for Panthera atrox being a particularly close relative of P. leo and said it might be closer to the jaguar, P. onca, instead, though it didn't find much evidence for that either.

By David MarjanoviÄ (not verified) on 18 May 2010 #permalink