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So, on to the contents of my BCiB talk (see previous article for preamble). We began by looking at Homotherium latidens, sometimes called the scimitar cat, scimitar-toothed cat or dirk-toothed cat. H. latidens is one of several Homotherium species that inhabited North America, Eurasia and Africa during the Pleistocene: the different species varied in body size, skull shape, the proportional length of the forelimbs, and in other features. It’s repeatedly been suggested that H. latidens might have survived in Britain until as recently as 11,000 years ago (close to the start of the Holocene): that’s an incredibly recent date given that most other evidence has indicated that homotheres didn’t survive in Europe beyond the Middle Pleistocene (beyond about 300,000 years ago).
This idea of late British survival for H. latidens has all been based on the presence of a single upper canine tooth discovered at Robin Hood Cave in Derbyshire in 1876. Unfortunately though, the Robin Hood Cave tooth has a hole drilled in its base, a fact demonstrating that it had been transported by people (it was presumably worn on a necklace), and isotope studies published in 1980 showed that it differed chemically from all other British homothere fossils (Oakley 1980, Charles & Jacobi 1994, Yalden 1999). These factors cast so much doubt on its origin that this tooth is no longer regarded as a valid record of H. latidens in Britain, and in fact it’s even been suggested that the specimen was planted in much the same way as were the Piltdown man specimens (Yalden 1999).
Having said all that, we do now know from a lower jaw dredged from the bottom of the North Sea [shown in adjacent image: borrowed from here] that homotheres did persist in northern Europe until about 30,000 years ago – in other words until late into the Late Pleistocene (Reumer et al. 2003). What’s particularly interesting is that the age of this jaw matches the age of the famous Isturitz statuette, discovered in south-west France in 1896 (Mazak 1970). These records – the North Sea lower jaw and the Isturitz statuette – probably don’t represent the last homotheres in Europe ever, so it’s actually pretty exciting to think that these sabre-toothed cats survived until surprisingly recently: that is, until late in the Late Pleistocene [for more on European homothere survival, see the article on this subject at Tet Zoo ver 1 here].
Homotheres might not have survived in Europe until the Holocene but another large cat – this time a species that’s still with us today – definitely did, and this is the lion Panthera leo. Lions occurred throughout Europe during the Middle and Late Pleistocene, and conventionally these European lions have been regarded as a type of lion – the so-called Cave lion – which was distinct from modern lions. Cave lions were more robust than living lions, with stockier limbs and broader, shorter-muzzled skulls, they are larger – up to a third larger than the biggest modern lions – and cave art [see below] shows that male cave lions did not have large or luxuriant manes.
It’s mostly agreed among experts that cave lions were not distinct enough (genetically or anatomically) from other lions to warrant recognition as a separate species (e.g., Hemmer 1974, 1979, Turner & Antón 1997, Patterson et al. 1999, Barnett et al. 2006, Sommer & Benecke 2006) [for an opposing point of view see Sotnikova & Nikolskiy 2006]. Burger et al. (2004) showed that cave lions represent an old lineage that diverged from modern lions during the Middle Pleistocene. The modern lion populations of Africa and Asia diverged from one another more recently, perhaps 200,000 years ago or so, and none of the modern populations have any strong genetic link with cave lions. So cave lions were a dead-end: they did not give rise to Asiatic or east African lions, as has been suggested on occasion (von Buol 2000).
Lions survived until surprisingly recently in parts of Europe: they made it into the Holocene in northern Spain, in Bulgaria and the Ukraine, and were definitely still present in south-east Europe as recently as about 5000 years ago (Sommer & Benecke 2006) [and then some: see comments]. They survived in Asia Minor until the 19th and possibly the 20th century. Which of these lions were cave lions and which were modern-type lions still remains somewhat uncertain, but it’s likely that the younger ones were more closely related to Asiatic and north African lions (both of which are more closely related to each other than to lions from southern and eastern Africa (Barnett et al. 2006)) and weren’t cave lions. As a particularly large, often social, cat that will often prey on livestock, lions have of course been heavily persecuted by people, so within the European region it’s not surprising that they were only able to survive in very wild and remote regions.
While restricted today to Africa and Asia, the leopard P. pardus was formerly a European species: like the lion, during most of the Pleistocene, leopards occurred across Europe. Phylogenetic trees compiled from DNA data indicate that modern leopards have African ancestry, and they seem to have migrated into Eurasia at the same time as members of our species first did the same thing (Uphyrkina et al. 2001). However, the oldest fossils claimed to represent leopards are from the late Pliocene and Early Pleistocene of India and Africa (Hemmer 1979)*: these and some other Old World Pleistocene leopards are inferred to be basal to crown-group leopards given that all extant leopards seem to have diverged from one another during the Middle Pleistocene and Late Pleistocene.
* Note that these fossils are controversial and have more recently been suggested to belong to jaguars or pumas.
Like the lion, the leopard did survive beyond the end of the Pleistocene and into the Holocene in Europe, as there are Holocene remains from Spain and Greece. In fact exactly how recently leopards remained in Europe isn’t entirely clear. Leopard bones from Greece and the Ukraine have been dated to the 1st century AD (Sommer & Benecke 2006) and might indicate that leopards survived in these areas until modern times. There are also leopard remains from the Middle Ages of Rome (De Grossi Mazzorin 1995), but these animals almost certainly represent imported individuals.
The Leopard is a highly diverse species: in fact it’s one of the most diverse cat species, varying to an incredible degree in body size, habitat choice, ecology, coloration and patterning. 27 subspecies of leopard have been described. Some of this naming of subspecies does reflect genuine variation within the species, but several recent studies (Miththapala et al. 1996, Uphyrkina et al. 2001, 2002) suggest that it might be more realistic to recognise just eight or nine subspecies: African leopard (P. p. pardus), South Arabian leopard (P. p. nimr), Persian leopard (P. p. saxicolor), Indian leopard (P. p. fusca), Sri Lankan (P. p. kotiya), Javan leopard (P. p. melas), South Chinese leopard (P. p. delacouri), North Chinese leopard (P. p. japonensis) and the Far Eastern or Amur leopard (P. p. orientalis).
As we’ve seen at Tet Zoo before, one of the persistent frustrations about diverse polytypic species like the leopard is that, while there might be nine subspecies, the vast majority of us have only even seen images or specimens of three, maybe four, maybe five, of these subspecies. Most of them are very poorly represented – not only in the literature, but also in zoological collections and on TV. We tend to only really see images or specimens of African and Indian leopards, both of which are relatively short-haired, robust, large-bodied leopards. European Pleistocene leopards have, generally without exception, been imagined to look like this, but given that modern leopards are far more diverse in appearance, is it possible that European Pleistocene leopards also looked quite distinct? What exactly might European Pleistocene leopards have looked like, and how were they related to living leopard populations?
Molecular phylogenies indicate that African leopards are the most primitive leopards, that those from western and central Asia have evolved more recently from African ancestors, and that the leopards of northern and eastern Asia are the most recently evolved (Miththapala et al. 1996, Uphyrkina et al. 2001, 2002). How do European Pleistocene leopards fit into this? Unfortunately, at the moment we can’t really answer this question because nobody’s yet studied the DNA of fossil leopards to see how they compare to living ones. Old European Pleistocene leopards might have been primitive leopards outside of the modern leopard clade, whereas others (specifically some from Rubeland, Germany) have been suggested to belong to the extant subspecies P. p. saxicolor (Hemmer 1979).
We do have one Pleistocene cave painting of a European leopard (it comes from Chauvet, France: shown here. The larger animal is a spotted hyaena) and it’s interesting in showing the leopard as having a particularly prominent white belly. Most living leopards don’t have this (they’re spotted on the belly), so maybe some European Pleistocene leopards would have looked distinctive to us. However, with just one cave painting to go on, this is all very speculative.
And there endeth part I. Next: jaguars, pumas, cheetahs!
Refs – -
Barnett, R., Yamaguchi, N., Barnes, I. & Cooper, A. 2006. The origin, current diversity and future conservation of the modern lion Panthera leo. Proceedings of the Royal Society B 273, 2119-2125.
Burger, J., Rosendahl, W., Loreille, O., Hemmer, H., Eriksson, T., Götherstrom, A., Hiller, J., Collins, M. J., Wess, T. & Alt, K. W. 2004. Molecular phylogeny of the extinct cave lion Panthera leo spelaea. Molecular Phylogenetics and Evolution 30, 841-849.
Charles, R. & Jacobi, R. M. 1994. The Lateglacial fauna from Robin Hood Cave, Cresswell: a re-assessment. Oxford Journal of Archaeology 13, 1-32.
De Grossi Mazzorin, J. 1995. La fauna rinvenuta nell’area della Meta Sudans nel quadro evolutivo degli animali domestici in Italia. Padusa Quaderni 1, 309-318.
Hemmer, H. 1974. Untersuchungen zur stammesgeschichte der Pantherkatzen (Pantherinae). Teil III. Zur Artgeschichte des Löwen Panthera (Panthera) leo (Linnaeus, 1758). Veröffentlichungen der Zoologischen Staatssammlung München 17, 167-280.
- . 1979. Fossil history of living Felidae. Carnivore 2: 58-61.
Mazak, V. 1970. On a supposed prehistoric representation of the Pleistocene scimitar cat, Homotherium Farbrini, 1890 (Mammalia; Machairodontinae). Zeitschrift für Säugetierkunde 35, 359-362.
Miththapala, S., Seidensticker, J. & O’Brien, S. J. 1996. Phylogeographic subspecies recognition in leopards (Panthera pardus): molecular genetic variation. Conservation Biology 10, 1115-1132.
Oakley, K. 1980. Relative dating of the fossil hominids of Europe. Bulletin of the British Museum (Natural History), Geology 34, 1-63
Patterson, B. D., Yamaguchi, N., Dubach, J. M. & York, D. 2005. Molecular genetics and morphological variation of lions (Panthera leo). African Lion News 6, 17-23.
Reumer, J. W. F., Rook, L., Van Der Borg, K., Post, K., Mol, D. & De Vos, J. 2003. Late Pleistocene survival of the saber-toothed cat Homotherium in northwestern Europe. Journal of Vertebrate Paleontology 23, 260-262.
Sommer, R. S. & Benecke, N. 2006. Late Pleistocene and Holocene development of the felid fauna (Felidae) of Europe: a review. Journal of Zoology 269, 7-19.
Sotnikova, M. & Nikolskiy, P. 2006. Systematic position of the cave lion Panthera spelaea (Goldfuss) based on cranial and dental characters. Quaternary International 142-143 (2006) 218-228
Turner, A. & Antón, M. 1997. The Big Cats and Their Fossil Relatives. Columbia University Press, New York.
Uphyrkina, O., Johnson, W. E., Quigley, H., Miquelle, D., Marker, L., Bush, M. & O’Brien, S. J. 2001. Phylogenetics, genome diversity and origin of modern leopard, Panthera pardus. Molecular Ecology 10, 2617-2633.
- ., Miquelle, d., Quigley, H., Driscoll, C. & O’Brien, S. J. 2002. Conservation genetics of the Far Eastern leopard (Panthera pardus orientalis). The Journal of Heredity 93, 303-311.
von Buol, P. 2000. ‘Buffalo’ lions. A feline missing link? Swara: the Magazine of the East African Wildlife Society 23 (2), 20-25.
Yalden, D. W. 1999. The History of British Mammals. Poyster Natural History, London.
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