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Pleistocene Big Cats
The Cave Lion (Panthera spelaea and Panthera fossilis)
The Cave Lion (Panthera spelaea and Panthera fossilis)
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The Cave Lion (Panthera spelaea and Panthera fossilis)
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I gathered published studies on cave lion evolution after the newest Barnett et al (2016) study and modified Sabol's (2011) excellent figure to incorporate all the findings together.
Original Sabol (2011) model:  *This image is copyright of its original author Multi-study model:  *This image is copyright of its original author I opted to have an ancestral Eurasian cave lion to split into P. spelaea and P. fossilis to incorporate the 3 migration waves theories of Hanko and Korsos (2007) and Sotnikova and Foronova (2014). Barnett et al. (2016) served to calibrate the split point at 1.89 mya and the P. fossilis of Isernia la Pineta for this species' first appearance. Thoughts? 
What does the Panthera "leo" sp. section mean?
Does it include the geographical clade of West African/Asiatic lions today, or the previous generation lions (1.2 - 1.4 MYA) born from P. leo fossilis, that eventually gave rise to P. leo atrox? By the way, the American Lion resembles quite a bit more like the former European Cave Lion (fossilis) than the newer European Cave Lion (spelaea) in terms of skeletal structure. Fossilis and Atrox both had a more elongated facial structure and, from speculation, a less-robust build than Spelaea. So it does make sense that Fossilis would be the direct ancestor of the American Lion.
"Leo" in quotes means the species is not Panthera leo but as it has not been discovered yet (and given a proper taxonomic name), the authors will call this (currently missing) link "Panthera "leo"" to have some ID to refer to. "ssp." means subspecies. So that point on the tree is indicating a subspecies of the ancestral Panthera "leo" was isolated in Eurasia and became the cave lion, while the original African Panthera "leo" became the modern lion.
The modern lion split is not shown in the tree. The P. fossilis evolution into P. atrox in the Americas (not shown correctly as a dichotomy due to spatial constraints) is shown. Correct on the relationship between P. fossilis and P. atrox, Sotnikova and Foronova (2014) very well describe such. The paper is in the library post on the first page. Postcranial-wise I do not remember seeing any robusticity difference between the American and Cave lion forms. Cranially, P. spelaea was quite more robust. 
How about both leo and spelaea/fossilis being directly evolved from shawi?
The African subspecies of shawi had been evolved into leo, while the Eurasian subspecies had been evolved into spelaea/fossilis. Is really necessary to put a transitional form between shawi and leo-spelaea-fossilis? Also, isn't the Beringian lion just a subspecies of the Cave lion? The Eurasian subspecies is usually the nominated one known as Panthera spelaea spelaea, while the Beringian subspecies should be Panthera spelaea vereshchagini.
Correct on the Beringian lion, that's why I showed a double arrow to show synonymy.
On P. shawi, the chronology and location of its record complicates things. All specimens attributed P. shawi can tentatively be considered chronological stages of the species, but one must remember that the P. shawi of Bolt's farm is around a million years older than the most recent P. shawi-attributed fossils. This leaves the possibility of the latter fossils being a unique species, although the lack of spatial movement in these specimens would reduce the possibility. Nonetheless, if we recognize 1 millions years of range grounds for speciation was already need a transitional form after P. shawi. There is also the location to be acknowledged. P. shawi is South African. The cave lion lineages will have to be from a population that originated in the upper half of Africa. This could result in the cave lion parent population not being exactly P. shawi due to the distance and thus the lack of genetic interchange between populations. We can dive into the issue with this scenario. If cats originated from east Asia, the first cats would have arrived and settled in North Africa. From there they would have moved south, and P. shawi may have been one of the feldis to evolve while the northern pantherine went on its own trajectory. A population of the northern pantherine would then enter Eurasia to became the cave lion. Thus in this scenario, P. shawi is actually never directly part of the cave lion lineage but rather a sister or aunt of sorts to the cave lion group.
But from what I read before, the Cave lion group was supposed to evolve from the Panthera shawi-like group in East Africa around 3 million years ago.
Since all Panthera shawi population in Africa remained some sort of genetic exchange, it did prevent them to diverge into different species. Let's say all Panthera shawi population from South Africa to East Africa remained subspecific for one million since 3 mya, but in 2 mya, the East African population suddenly migrated to Eurasia, and in just a brief period, the migrating population had diverged into a different species due the genetic isolation. And the South African population remained in Africa and dispersed to the rest of Africa, and finally evolved into the modern lions that we known today.
P. shawi is so poorly defined due to the records being teeth and a broken humerus that in most cases, one can get away with calling any old African pantherine from that time period P. shawi. Would you remember where that was read? I may have missed it or forgotten.
In terms of population movement, usually it's a portion of a population that migrates and not the entirety. So the northern form of P. shawi (or whatever we'd like to call the stem group) that stayed in Africa could have been the parent of modern lions as well.
It was from some quite old sources which I couldn't even remember the title and author properly.
But I think Panthera shawi should be the basal species/common ancestor for all lion species we known. It was the genetic isolation that created the interspecific status of the Eurasian/American Pleistocene lions. The modern lions look mainly descended from the southern group of Panthera shawi, but when they dispersed to the rest of Africa, they could have managed to absorb the leftover of the northern group. From some anecdotal observation, I've seen the canine teeth of East African lions bear some resemblance with the Cave lion canine teeth. Maybe some modern lion population could show some closer genetic affinity with the Cave lion group than the rest.
A gigantic P. fossilis
Reading through records, no one noticed Sabol's (2014) giant. The paper: Panthera fossilis (REICHENAU, 1906) (Felidae, Carnivora) from Za Hájovnou Cave (Moravia, the Czech Republic): a fossil record from 1987 – 2007 The reason may be that this specimen is nothing more than a humerus in which about a 3/5 of it remains distally. However, the width measurements caught my eye. Unfortunately, there are no measurements in other literature that correspond to what Sabol took. Nonetheless, Sabol presented a beautiful image with a 3 cm scale bar which was almost 99% accurate allowing measurements of more common humeral dimensions. Here is probably the largest humerus of a pantherine we will see for a long while:  *This image is copyright of its original author For comparison, the widest P. atrox humerus had a greatest distal width of 111.3 (and a total length of 396 mm). The longest P. atrox specimen (humeral length = 409 mm) had a distal width of 107.6 mm. Using the ratios of the P. atrox femurs, this P. fossilis humerus would be 441 mm in total length. Then we have the midshaft width comparison. This specimen midshaft traverse width of about 45 mm exceeds that of the two largest P. atrox humeri (length = 396 mm, midshaft width = 41.4; length = 409 mm, midshaft width = 38.2 mm). Using the P. atrox midshaft widths we get a total humeral length of 455 mm for this P. fossilis specimen. Comparing the P. fossilis fragment to a few cave lion and American lion femur it look to be about 450 mm in total length. Therefore, the massive width dimensions alongside the predicted length put this specimen in the candidacy for the largest cat on fossil record to date. Comparing the 43 mm tranverse width at the point of least circumference to a 32.5 mm wide, 187 kg lion bone puts this specimen at 450 kg in my estimation, probably outsizing the giant 465 mm ulna and the 192 mm MT3 (which may be proportionately longer in P. fossilis looking at Marciszak et al. 2014 data).
That's difficult to say due to the poor correlation, but using the humerus and the trend that P. fossilis probably had a longer skull anyhow, 450 mm-500 mm may be an okay range.
(01-09-2017, 10:52 PM)tigerluver Wrote: That's difficult to say due to the poor correlation, but using the humerus and the trend that P. fossilis probably had a longer skull anyhow, 450 mm-500 mm may be an okay range. Since Panthera fossilis got proportionally longer skull than both Panthera atrox and Panthera spealea, then I assume they are leaning toward the upper end. And the 450 kg estimation is based on the proportion of a modern lion, but the actual proportion of Panthera fossilis could be much more robust. With a GSL about 500 mm with a body mass up to 500 kg, what is your regard about this Pleistocene giant lion? @peter 
AMATEUR SPECIALISTS
In the days I visited one museum after the other, I met a number of people interested in Pleistocene big cats. Some of them had collected a lot of information over the years. Every now and then, we discussed Pleistocene big cats and those of today. At times, conservators joined and showed us a number of great skulls. Based on the measurements I had, we concluded that European lions had been larger than the largest wild big cats measured in the last 2 centuries or so. We never got to a satisfying conclusion on size because of a lack of data, but the bones and skulls I saw were large, dense and heavy. Most were found in southern and eastern parts of Europe. Judging from the info recently posted by Tigerluver, those found in central Europe could have been even larger. Many of these amateur specialists, all of them well-educated and well-informed, thought the size of European lions had declined during the Pleistocene. Climate change could have been a reason. The changes had a profound effect in that forests replaced plains. The result was the large herds moved south and east. Some lions followed. Lions remaining in the northwestern part of Europe showed a greater decline in size than lions who followed the herds. They also hunted different animals. Bones of European lions found in the west of Europe and in the sea that separates Europe from the UK are smaller than those found in southern and central parts of Europe. Based on what I read and heard, however, males still reached impressive dimensions. One of the few who tried to get to a bit of insight in dimensions concluded that the male found in the western part of Germany some decades ago reached a length only seldom seen in the big cats of today (200-210 cm. in head and body in a straight line). THE SIZE OF PLEISTOCENE EUROPEAN LIONS It's difficult to imagine a big cat with a greatest skull length ranging between 370-500 mm., let alone get to a decent guesstimate on body dimensions. If we want to give it a try, we have no option but to use information about big cats shot between 1850-1950 in Africa and Asia. The reason is there are enough records and details. The problem is we have no clue as to the correlation between size and mass in Pleistocene big cats. As they could have been more robust than big cats of today (likely), we could contemplate adding reliable info about brown bears (if we know how they compare to big cats). The problem is there is no study. Furthermore, there are significant differences between big cats and bears. Big cats are full-time hunters, whereas bears are omnivores. For a hunter, weight is important. The limit, apart from individual variation, depends on the local conditions. Today, a big cat of 400 pounds is a large animal everywhere. The only regions that would qualify are India, Nepal, parts of Indochina, Russia and some parts of Africa. If we raise the bar to 450, India, Nepal and, perhaps, a few hotspots in Africa remain. If the bar is raised to 500, not one region, except for a few hotspots, would qualify. For this reason, I propose to use 475 pounds as the limit of today (averages for adult males). Based on what we have, only some parts in India and Nepal would qualify. How use what we have to get to a guesstimate for European Pleistocene lions? here's some ideas. ASSUMPTIONS, PROPOSALS AND CALCULATIONS We know that the largest representatives of a species can be 30-40% larger than average (big cats). This means the expected 'normal' maximum would range between 602,5 (30% of 475) and 700 (40% of 500) pounds. Any details known about these giants? Yes. Extra-large tigers nearly always are longer (referring to total length and not head and body length) than average, but not to the degree seen in mass. An extra length of 10-15% seems to result in an extra mass of 30-40%. This means that mass, when length is increased, increases by a factor 2, if not more. Have lions of 600 pounds or thereabout been shot, measured and weighed? Yes. Same for tigers ranging between 650-700 pounds or even a bit over? The answer again is affirmative, meaning the hypothesis above can't be rejected directly. Anything known about the greatest skull length of extra-large individuals? Yes. The longest tiger skulls most probably reach 16 inches or a bit over. Lions can reach 16,5-17,0 inches with less mass. Anything known about the correlation between greatest skull length and mass or other dimensions in tigers? Yes. Although some individuals, like the Hasinger tiger of 10.7 straight in total length who had a greatest skull length of 14 inches only (about average), seem to point in another direction, reliable data from Cooch Behar showed a positive correlation between greatest skull length, body dimensions, age and mass. The Cooch Behar tables I posted a year ago said males averaged 461 pounds and 295 cm. in total length 'over curves' about a century ago. I propose to assume they averaged about 14 inches (about 353 mm.) in greatest total skull length. I also posted a table which had information on long-skulled male tigers. Same region. These males averaged 375 mm. in greatest total skull length, 493 pounds in weight and 10 feet in total length (measured 'over curves'). They were about 3% longer than an average male in total length (measured 'over curves'). In greatest total skull length and weight, they had about 6-7% on an average male. RESULTS If we use the data from Cooch Behar to get to an assessment on the size of Pleistocene big cats, we have to assume that European lions compared to Cooch Behar tigers in all respects. This, most definitely, was not the case, but we have no other option for now. I propose to use an average greatest total skull length of 450-460 mm. for starters. This means the Pleistocene European lions had about 30% on Cooch Behar tigers. It also means they were about 15% longer (total length measured 'over curves') and 35% heavier. If we use everything we have, an average Pleistocene male European lion would have averaged 600-620 pounds and 339 cm. in total length 'over curves'. If we assume that extra-large individuals, weightwise, also had 30-40% on an average male (of 610 pounds), just like today, they would have maxed out at 780-860 pounds, maybe a bit more (the outcome of the calculations was based on very conservative assumptions). Giants of that weight, if they had been similar to Cooch Behar tigers in all respects, would have been 375-385 cm. in total length measured 'over curves' (12.4-12.8). Measured in a straight line, they could have reached 11.6-12.0 (350-366 cm.). DISCUSSION There's no doubt that Pleistocene European lions and Cooch Behar tigers were very different animals. If we include everything we know about density, mass and size, the European lions might have been a bit shorter than what was assumed above (11.6-12.0 in total length in a straight line), but more robust and, as a result, less agile. Not saying they would have compared to brown bears of today, but they would have been very robust. This is what the bones and skulls suggest. It's difficult to imagine. A true cat of 800 pounds and over would have needed large herbivores close to home to stand a chance. On the other hand. When the British entered India and started hunting big cats in earnest, they, not seldom, were amazed at what they saw. How many writers reported about tigers reaching 11 and even 12 feet in total length? Did all of them, as many seem to think, escape from an asylum where they had been taught that a foot has 11 inches? Were they released on the condition they would move to India and target tigers only? Did those who had a position in life agree to cheat everyone at every possible opportunity? This in the days a position and responsability were connected and still had some meaning? Why is it the calculations above produced results that, to a degree, support the information about size in the countless reports about extra-large tigers shot in the 18th and the first half of the 19th century? Why is it that Nepal male tigers lost 4 inches in total length when they had been hunted for a decade only? And why is it that two of the few males actually weighed still bottomed a 600-pound scale (Nepal has about 200 tigers only)? As to the size of Pleistocene European male lions. My guess is they didn't need 780-860 pounds to produce skulls with a greatest total length of 450-460 mm. (at similar size, lions produce longer skulls than tigers). Those who got to 500 mm., however, might have reached or exceeded the weights mentioned. Also remember the weights were based on conservative assumptions. Tigers living in regions that have big herds of large herbivores are more robust than tigers elsewhere. Some males seem to be out of this world. In these hotspots, males could reach an average of 500 pounds. What would happen when the reserve would be tripled in size and humans would be removed for a long time? If we add serious protection and severe legislation, anything is possible. Protection and good conditions apparently had an effect on the size of tigers in the Imperial Hunting Reserve in the extreme east of China (see 'The Tiger's Claw'). Not saying an average male would exceed 600 pounds In Assam in a decade from now, but they could surprise us. Size is a bit of a mystery in big cats. Returning to Pleistocene European lions. The bones found recently suggest they might have been among the largest big cats ever. Similar to the Pleistocene Java tigers or even larger? Could be. Bones of jaguar-like cats found recently also point in the direction of a very large cat, quite a bit larger than his relatives in South America. Europe wasn't the cradle of big cats, but it could have produced the largest big cats ever.
Me and @tigerluver we both tried to specialize on the limited materials available to us.
Canine teeth: Based on what we discuss so far, a 500 mm lion skull is pretty much producing the largest upper canine teeth for any pantherine cat, comparable to the few obscure massive tiger canine teeth I just posted. When both lion and tiger have the same canine teeth, tiger turned out to be the smaller cat. So the owners of these huge tiger canine teeth are arguably smaller than this super giant lion. Skull and limb: @tigerluver made his own research and found out that these available measurements has reached the theoretical max for the big cats. They were not only longer, but also much thicker than the late Pleistocene lions. I guess the middle Pleistocene was warmer, so the middle Pleistocene lions didn't need to spend too much energy to against the harsh winter that frequently occurred in the late Pleistocene. Although my conclusion sounds amateur-ish, but everything seems to match perfectly, all puzzles fit in its place perfectly.
This thread is a good one, Grizzly. There's no question that you and Tigerluver do a great job on Pleistocene big cats. I enjoy reading threads in which extinct big cats feature.
When you asked for an opinion about the size of the owner of the enormous bones found in central Europe, I first decided to read the notes I had made during discussions with amateur specialists many years ago. I added 'amateur' to make it clear that their ideas had not been tested in any way. This means that it's hard to say if these ideas are valid or not. I do know I consider them as very well-informed and dedicated specialists. Their opinion on Pleistocene European lions is very outspoken in that most think that they could have been the largest big cats ever. As they didn't agree on a guesstimate, I had no other option but to use data of big cats shot and measured in the last two centuries. I used tigers, because of the number of records. The results of the calculations I did were extrapolated to Pleistocene lions. I know the method used is a poor one. I also know that the assumptions were incorrect. In spite of that, the results of the calculation can be considered as a start. If the Pleistocene European lions would have compared to Cooch Behar tigers shot a century ago, males would have averaged just over 600 pounds and reached a length of just over 11 feet 'over curves'. Exceptional animals could have reached 780-860 pounds. This regarding male lions with a greatest total skull length of 450-460 mm. (not those approaching 500 mm.). The remark on canine length you made is important. If Pleistocene European lions would have compared to lions today in the canine department and produced canines similar in length to those of the largest Pleistocene tigers, chances are they would have outsized these tigers by a margin. Lions have, and most probably always had, shorter canines than tigers. A Pleistocene lion with canines which compared to those of the largest tigers, for this reason, had to be larger. The question is how much larger. The largest tigers I'm aware of are the Pleistocene Indonesian tigers. The bones of the Pleistocene lion found in Europe, however, were larger and more robust. If some Pleistocene tigers really exceeded 400 kg. (883 pounds), chances are the owner of the lion bones found in central Europe, as you suggested, could have reached 1000 pounds or even more. The people mentioned above (I consider them as authorities, which, regarding extinct big cats, is not true for yours truly) concluded Pleistocene lions differed from the lions of today in that they were not only longer and taller (by about 10-20%), but relatively more robust. This, they thought, was needed to get to the agility, speed and power needed to hunt some of the extra-large herbivores that surrounded them. It's a guess only, but it is possible they could have been an improved version of the tigers in northeastern India today. Assam tigers are not longer or taller than average, but more robust and big-skulled. With 'big', I mean they have relatively wide and heavy skulls for their length. Robustness seems to be an adaption to very big prey animals. Before posting the calculations (referring to the previous post), I took a tape and made a few points on a wall in order to get an idea of the size of Pleistocene European lions. I was flabberghasted by what I saw, but have to admit that the information you and Tigerluver posted more or less fits the picture that emerges from the bones. |
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