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

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).

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].

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.

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.