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.
Is it not possible that the leopards in Italy were feral escapees from the Roman games? I don't know when animals started to be used and transported, or what species were used, but it's a thought. Perhaps you could blog about it at some point?
Presumably that wouldn't account for the Ukrainian records, though.
Isn't there good evidence of lions in SE Europe much more recently than 5000 yr ago? The Greek historian Herodotus specifically mentions lions attacking pack animals in King Xerxes' Persian army marching through what's now Bulgaria/north Greece on its way to conquer Athens. I vaguely remember that he states that lions were formerly widespread but were now confined to one particular area of Greece, but I'll have to dig out the book and check that.
Lions are also mentioned repeatedly in the Iliad, so they were obviously familiar animals to the Greeks of Homer's time.
The extinction of lions in Europe isn't too hard to understand, as an open-country big cat would be relentlessly persecuted by herdsmen and its habitat taken over by agriculture. However I think it's more of a puzzle why leopards didn't last longer in Europe. As forest/hill country cats you'd think they would still have had plenty of habitat and prey until well into mediaeval times at least, and the climate here is obviously no problem for a cat that can survive in Siberia and northern China.
"We began by looking at Homotherium latidens, sometimes called the scimitar cat, scimitar-toothed cat or dirk-toothed cat."
I may be wrong, but isn't Homotherium considered a scimitar-toothed cat (i.e. shorter and broader upper canines with a lighter build than the "dirk-toothed" cats)? At least that's what I gathered from Martin, et al.'s paper "Three Ways to be a Saber-Toothed Cat."
Darren - do you know if there has been any publication on the buffalo killing lion = cave lion descendant theory recently (I presume that von Buol ref is along those lines)?
I recently read Ghosts of Tsavo which pushed the theory quite heavily but a) suggested publication was a way off and b) was a very unscientific read (whilst being quite interesting).
The biggest question to me is why tigers and leopards only occur at the eastern end of Siberia... is the rest of Siberia so different?
Your reference to Mazak 1970 contains three mistakes: Zeitschrift für Säugetierkunde.
Reading Sommer and Benecke (2006), I'm not convinced that they were saying that leopards are known from Italy in both the 1st Century AND the early Middle Ages. My reading of the relevant paragraphs (pasted below) is that remains are known from the Ukraine which could be imported, and there is also one isolated record from Italy associated with the Colosseum. Looking at the map the Italian record is right on top of Rome, so I think there's only these imported animals known from Italy at this time.
"The imprecisely dated Mesolithic record from northern Spain (Altuna, 1967) and the early Holocene bone remains from Greece (Symeonidis, Bachmayer & Zapfe, 1980) are two indicators that the leopard could have survived into the Holocene in Europe. In the later Post-Glacial periods, P. pardus appears first in the sub-Atlantic in the subfossil deposits of the Ponto-Mediterranean region.
Finally, the lack of accurately dated subfossil records in Europe requires to be taken in their interpretation. The Iron Age records from the Hellenistic period could possibly be imported individuals of P. pardus from Asia Minor. From this point of view, it is remarkable that all remains of the leopard discovered in the Hellenistic Olvija in the northern Black Sea region are assigned to a period in which Olvija was a colony of Greece. The conclusion of Zuralev (1999), that the bone remains of the leopards of Olvija were from free-living individuals, is not strongly supported. The early Middle Ages remains of P. pardus from the Colosseum in Rome (De Grossi Mazzorin, 1995) are obviously a matter of import (J. De Grossi Mazzorin, pers. comm)."
Of interest is Michel Meurger's paper in Fortean Studies 1, which discusses European leopard stories from a folkloric viewpoint. Unfortunately I don't have it to hand so can't remember the specifics, but he definitely doesn't think that leopards survived in the Alps until the Middle Ages, and I believe links them to stories of mountain devils attacking pilgrims.
Meurger, M. 1994. The Leopards of the Great Turk: mystery felines in medieval France. Steve Moore (ed.). Fortean Studies 1, John Brown Publishing, London, pgs. 198-209.
> There are also leopard remains from the Middle Ages of
> Rome (De Grossi Mazzorin 1995), but these animals almost
> certainly represent imported individuals that were used
> in the Roman games.
The games persisted into the Middle Ages? I understand that many kinds of animal-baiting were practised well into the 20th century, and are probably still practised nowadays in some corners of the world, but I think the roman games in the classical sense ceased in the fourth or fifth century AD for both religious and economical reasons.
According to Wikipedia (I know not the best source, but there is a reference given (Claridge, 1998), which I haven't seen) animal hunts continued at the Colosseum until at least 523 A.D. The Early Middle Ages are usually thought to start approximately 500 A.D., with the fall of the Western Roman Empire, which apparently (Wikipedia again) is usually dated to 480 A.D.
Claridge, A. 1998. Rome: An Oxford Archaeological Guide. Oxford University Press, Oxford, UK, pp. 276-282.
Lots of comments:
- lions are usually said to live until 10.cent. A.D. in Balkans, vide all Greek myths related to lions.
- I wonder again if any specimens exist of Balkan lions, to test their relationships as Persian/Cave lions?
- What is known about lions in N Spain?
- I wonder if you checked Russian literature about lions and leopards in in Caucasus and Ukraine area? Geographically it is Europe. I'm pretty sure leopards lived in northern Caucasus until last decades.
A leopard was photographed in Georgia in 2004:
So the species is still hanging on at the fringes of Europe at least.
Thanks to all for comments, I'll address more later, but briefly: my reference to 1st century leopards was a mistake, I meant Greece, not Italy! (have changed this). Furthermore, '1st century' is a misinterpretation on my part: I based this on Sommer & Benecke (2006), who give records from Greece and the Hellenistic Olvija in the northern Black Sea region as '1000 BC-0'. My numerical problems somehow led me to interpret this as 1st century AD, which it isn't (unless you want to count the year 0 as part of the 1st century). Sorry! To spoil things further, the Olvija remains (they were described by Zuravlev (1999)) are probably not of wild animals, but of imported ones (Sommer & Benecke 2006, p. 14: see Richard's quotation above).
PS - one for David: Zuravlev has a caron on the Z but I can't find the html for it.
Ref - -
Zuravlev, O. P. 1999. Säugetierfauna und Umwelt am Südlichen Bug während der Antike. Archäologie in Eurasien 6, 389-407.
From Book 7 of the Penguin English translation of "The Histories" by Herodotus, who lived between approx. 485-425 BC:
Referring to the march of Xerxes' Persian army through Thrace (roughly modern Bulgaria) and Macedonia, in 480 BC:
" It was during this march that his pack-camels were attacked by lions, which came down from their haunts at night and never molested either the men or any of the other animals, but only the camels. I am puzzled at what it could have been that made the lions ignore every other living creature and set only upon the camels - beasts which they had never seen, or had any experience of, before. This part of the country- namely the region between the River Nestus which runs through Abdera and the Achelous which runs through Acarnania - abounds with lions, and also with wild oxen, which have those enormous horns that are imported into Greece. There are no lions anywhere in Europe east of the Nestus, or in the continent west of the Achelous; they exist only in the country between those rivers."
Lions hunting aurochs in southeastern Europe - what a tragedy we can't see that today. It would be a lot cheaper than a trip to the Masai Mara!
So #142 it is then. The Windows character table says the shortcut is Alt+0142, I tried that, and it worked.
Lowercase (): #158.
David: the area of Russia above the Amur is actually called the Russian Far East and is very different from the taiga and tundra of Siberia proper to the north and west. It has milder winters (though a lot of snow), deciduous trees, and many species from eastern and southeastern Asian ecosystems, such as tigers, leopards, musk deer, raccoon dogs, paradise flycatchers, and ginseng. It is like the area south of the Amur once known as Manchuria, only (so far) better- preserved.
But did Herodotus's Greek contemporaries import just the horns or the wild oxen they were attached to?
Indian leopard (P. p. fusca)...Javan leopard (P. p. melas)
I'm not absolutely sure, but I don't think these can both be correct: they should either both be masculine (fuscus, melas) or both feminine (fusca, melaena).
The complicating factor here is that the genus and species names differ in gender (Panthera is feminine, pardus masculine). This is OK since they are both nouns, but I haven't been able to find out whether the subspecies name is supposed to match the genus or the species. Subspecies of the other mixed-gender Panthera species (P. leo and P. tigris) generally appear as feminine. David, have you read anything about this?
(The other subspecies names listed are all either nouns, genitives of names, or adjectives that have the same form in the masculine and the feminine.)
David: the area of Russia above the Amur is actually called the Russian Far East and is very different from the taiga and tundra of Siberia proper to the north and west.
That's true, but this biome extends west all the way to Europe, doesn't it?
David, have you read anything about this?
All "species-group names", which means names of taxa at ranks that belong to the species group of ranks, which includes species, subspecies and "species group" (...which is sometimes called "superspecies", but not in the ICZN!), are supposed to match the genus according to ICZN Article 31.2.
However, Greek adjectives, when not latinized, are not changed to agree in gender with anything according to Article 31.2.3, which lumps Greek with Tibetan (Tibetan lacking anything similar to gender). People are officially not supposed to know Greek anymore.
this biome extends west all the way to Europe
Plant-wise I mean.
Good article. I personally think that the cave lion would be a biologically separate species when compared to modern lions, even given their shallow divergence. What we know of cave lion external morphology would make them unlikely to be behaviourally compatible with modern lions. i.e. how would an African female recognise a male cave lion in the absence of a mane?
I have a hard time believing that the Holocene lions from Europe are refugial cave lions. It doesnt really make sense when compared to the extinction patterns of other large mammals of Europe. If anyone had well preserved late lions it would be really easy to DNA test whether they were cave or african lions.
Interesting thing about the Robin Hood scimitar tooth is that it is only partially perforated, so whoever was trying to turn it into a necklace didnt quite make it. (do a GIS on "robin hood homotherium) to get an image of it.
From the ICZN:
34.2. Species-group names. The ending of a Latin or latinized adjectival or participial species-group name must agree in gender with the generic name with which it is at any time combined [Art. 31.2]; if the gender ending is incorrect it must be changed accordingly (the author and date of the name remain unchanged [Art. 50.3.2]).
45.1. Definition. The species group encompasses all nominal taxa at the ranks of species and subspecies (see also Article 10.2).
I guess that means the Javan leopard should be Panthera pardus melaena.
Ah, xpost with David and Ross.
Thanks Ross. Quick responses...
how would an African female recognise a male cave lion in the absence of a mane?
We now know that the mane is not a universal character of male P. leo: more on this when I get round to talking about Tsavo lions, buffalo lions and so on.
I have a hard time believing that the Holocene lions from Europe are refugial cave lions
So does everyone else. I don't think anyone is advocating this (although there's always Loren Coleman's theory that American black panthers are modern-day P. leo atrox!).
I'm just glad I saw 10,000 BC before I read this-I'm sure
I would have been terminally depressed to discover that
Smilodon was long gone by the time the Atlanteans landed
in Northern Egypt. ( Fun movie, but leave your brains at
the Concession stand...)
Is mating among Lion that dependant on visual signals? I
would have thought pheremones would have been more key.
I guess that means the Javan leopard should be Panthera pardus melaena.
No, because it's not "Latin or latinized" (your quote of 34.2).
Is mating among Lion that dependant on visual signals?
For what it's worth, dogs don't seem to care at all what a potential partner looks like. They just sniff all the time.
For what it's worth, dogs don't seem to care at all what a potential partner looks like. They just sniff all the time.
And then you have peacocks.
No, because it's not "Latin or latinized" (your quote of 34.2).
You are quite correct. I assumed that because melaena is Latinized melas would be considered also Latinized, but 31.2.3 is explicit that it is not.
I'm wondering how widespread leopards were across Europe, and whether they would have been more typical of humid, warmer interglacials, just like hippopotamus, macaque and straight-tusked elephant, or whether these Pleistocene European leopards would have been just as comfortable hunting in the snow alongside woolly mammoths and reindeer.
If these leopards turn out to have done just fine even when glaciation was at its peak, then it's likely that they might have been cold-adapted, the same way the Amur leopard and North Chinese leopards are. On the other hand, if they were dependent on somewhat warmer climate, they might have been more like Persian or Caucasus leopard.
Just how tolerant to temperate climates are extant leopards, anyway?
However I think it's more of a puzzle why leopards didn't last longer in Europe. As forest/hill country cats you'd think they would still have had plenty of habitat and prey until well into mediaeval times at least, and the climate here is obviously no problem for a cat that can survive in Siberia and northern China.
That's a good question. Hell, I've heard leopards can and do live very close to people in Africa and Asia...
Such a shame P. atrox apparently didn't have a mane, because I was otherwise completely comfortable with the notion of a half-ton social cat roaming around North America - especially with females that look like "black panthers" for some reason...
Several years ago I saw the very last non-introduced european big cat. It was in a little museum at Samos, Greece. Among many interesting pleistocene mammal fossils they had a stuffed leopard from the beginning of the 20th. century, which did swim from Turkey to Samos. Further info here: http://hellas.teipir.gr/Thesis/Samos/english/tdk160.html
If I have time (perhaps in some weeks) I´ll go again to the museum of Schloss Hohentübingen, which owns not only many striking archaological relics, but also some world famous master pieces of the ice age, among them the little figurines from the Vogelherd-Cave. They are made of ivory and show little animals, not as big as a finger. But their acuracy is amazing, you immediately see what animals they are. There are lions and something which is most probably a leopard. I have somewhere on a cd still photos I made at my last visit, but I don´t know where. But if I find them or get new ones, I can send them to you. There is also another amazing and comparably big figurine, a human with the head of a (female?) lion. Lions seems to have had a big impact on humans.
And then you have peacocks.
I wasn't generalizing about "animals", I was generalizing across Carnivora. That seems fairly defensible.
David: I am no botanist. But I think arid ecological zones make the deciduous zones very discontinuous across Asia, as in the US (there is no deciduous zone in the Rockies or the Great plains in North America, only in the East.)
In Kazakhstan there IS a deciduous tree ecosystem in the Tian Shan foothills, and it includes species found in the Russian Far East, but there is not a continuous connecting "band" AFAIK, unless it used to exist in part in agricultural China.
The leopard remains coming from a V_VI centuries AD layers (no middle age). The last "venationes" was held in the Colosseum in the middle of the 6th century (see Cassiodoro, var, V, 42, ep. 142)
"However I think it's more of a puzzle why leopards didn't last longer in Europe. ...climate here is obviously no problem for a cat that can survive in Siberia and northern China."
it can be a mistery indeed, but what is important in the case of leopard - the cat has little problem adapting to cold itself - just like any pantherine cat of to-day including lions ( the do prosper in the Novosibisk Zoo open exibits, in the climate of middle Siberia with temperature as low as minus 45 in January).
BUT - unlike long-legged tiger and lion, for a leopard the snow cover depth is critical.
When it snows heavily so the depth of snow is more the 25 cm for over a week - the spotted cat is doomed.
Leopard has absolutely no adaptations for stable successful hunting in deep snow. It can tolerate much colder winters than the Southern "Siberia" (actually it is southern Maritime Region of the Russin Far East, not Siberia in technical terms) but leopard starves in winter in deeply snowed areas.
I am sure that middle Age Cental, Northern and Western Europe were all areas of deep snow cover for quite a prolonged period of times. Add here that after dying out of european lions, there were no effective ecosystem regulators for populations of gray wolf. And the latter is a deadly enemy of leopard especially in a conditions when the cat is weakened with hunger and has problems trevelling in the snow.
In Southern Europe the reason for exntinction of leopard well could be direct exterminations by people. It has always been a densely human-populated zone and unlike India and Africa, Southern Europe would have VERY low level of wild ungulate population for millenia. So local leopards HAD to come into constant conflicts with southern-european heardsmen.
"The biggest question to me is why tigers and leopards only occur at the eastern end of Siberia... is the rest of Siberia so different?"
Yes, the rest of Siberia is VERY different what what you in the West call "Eastern Siberia".
Those are two different zoo-geographical regions of Russia, with the Far East being the actual pleistocene-type refugium with ecoton-type ecosystems (many different landscapes, different vegetation and different animal species mixed in small territory).
The Siberia itself (roughly - from the north of the Eastern Mongolia and westward up to the Urals) - is a dead land. The land is covered mostly with the so-called "dark-conifer forests" and endless cold swamps to-day. In the absecence of rhinoceronce and mammoth and bison herds, it is not the rich steppes and parklands anymore, unlike its pleistocene state.
Neither lion, nor tiger, and NOR leopard have much to eat there. :-(
Even though young migrating Siberian tiger males do come to the eastern fringes of the "true Siberia" from time to time they have big problems with finding enough prey to sustain themselves and also - with horrible local poachers.
As for the leopard - read the above about the crucial role of snow in life of this cat in the Northern areas.
I hope you read comments to earlier posts.
What CGI exist of Homotherium?
Am I right that BBC produced CGI stills of Homotherium as either white or grey-spotted animal for "Walking with Beasts", but no moving animation was ever published?
Regarding the cave painting, it's interesting to note that the Hymalayan snow leopard (now included in Panthera as P. uncia) does have a white belly. Considering that 1) Glacial Europe wouldn't be that different from its modern habitat and 2) leopard remains in Europe are awfully scarce and partial, couldn't it be possible the European remains, or at least those from the Pleistocene, actually belonged to the Hymalayan species?
Hi Spanish Guy - that's an interesting idea, but, alas, it's just an idea. You'd need fossil evidence to support it. So far, all those European fossil leopards do indeed seem to be leopards, not snow leopards. The latter species is quite distinctive skeletally; to date, no evidence for it in Pleistocene Europe (so far as I know). Incidentally, it's certainly plausible that the Snow leopard did reach the far eastern fringes of Europe once upon a time. It's far from restricted to the Himalayas, occurring (historically) as far west as Kazakhstan. It's not as well known as it should be that many large mammals typically associated with eastern Asia once (or still) occur in central Asia too.
Re the cave painting of the hyaena and the leopard: the cat with the white belly doesn't seem to have rosettes, it has spots. Since these artists were very detailed in their paintings of these animals, are you confident that this is indeed meant to be a leopard? The coat pattern looks more like that of a cheetah, though the body of the animal looks more like a leopard. Any thoughts?