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The Cave Lion (Panthera spelaea and Panthera fossilis)

Canada GrizzlyClaws Offline
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#46
( This post was last modified: 01-10-2017, 01:48 PM by GrizzlyClaws )

@tigerluver is more about skull/limbs, while I am more about the skull/canines.

Together, we are trying to create the grand picture from our own perspective and the domains that we are most specialized with respectively.

I made a comparison with African lion skull and Barbary lion skull, I found out that the Barbary lion as a more solitary modern lion subspecies, they tend to have proportionally longer canine teeth than their African cousins. Same observation also applies to the Pleistocene lions, since they lived in smaller group, their canine teeth were also more developed, and the ratio is in between that of the modern lions and tigers.

I initially thought this fossilized canine tooth was a Pleistocene tiger's, but later I figured out it was a lion lower canine tooth. It is almost 40% longer and 50-60% wider than the lower canine teeth from a 40 cm-ish African lion skull. Overall, the proportion of this lower canine tooth also matches those 50 cm-ish Pleistocene giant lion skulls.

The upper canine teeth would be no less than 16 cm, and its diameter is also no less than 4.5 cm, very close to the magnitude of the largest tiger canine teeth. Only a lion with this type of canine teeth would produce a 50 cm skull.



*This image is copyright of its original author



*This image is copyright of its original author
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Netherlands peter Offline
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#47
( This post was last modified: 01-10-2017, 09:46 PM by peter )

I was aware of the previous posts, which were very interesting. I think both of you did a great job. It was a textbook example of cooperation. The conclusions no doubt will be of interest to those involved in extinct big cats. Keep it coming.

I'll deliver a number of tables in a few months. Some of these will be about lions. The tables are instrumental for the papers we want to publish. All of them will have individual entries and quite a few details. I will also post a number of photographs of skulls I consider typical for African and Indian lions. Indian lion skulls are different in that the occiput in male skulls is quite similar to those seen in skulls of Javan and Bali tigers. 

The problem with wild big cats is individual variation. You could be right regarding the canines of lions leading a more solitary way of life, but we need large samples to get to solid conclusions. The problem is only few museums have skulls of and extinct big cats. I'll do my best to find out a bit more.
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India brotherbear Offline
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#48

https://www.wired.com/2011/11/cave-lion-diet/ 

Ancient Cave Lion Bones Reveal Big Cats’ Diet 

BY ANDREW CURRY, ScienceNOW

A quarter larger than today’s lions, the European cave lion was one of the biggest cats around 12,000 years ago. Now, an unusually sophisticated analysis of its bones is revealing what these creatures ate—and why they may have disappeared. 
Although they were certainly massive cats, the term “cave lion” is a bit of a misnomer. Unlike today’s lions, males probably didn’t have manes, and they appear to have been solitary hunters. What’s more, though their bones are best preserved in caves, they probably lived in the open. But they did have one thing in common with their modern relatives: they appear to have worried humans. The big cats show up in ice age cave paintings and in ivory figurines, suggesting that they were a major concern for our ancestors. 

To figure out what these lions hunted, biogeologist Hervé Bocherens and colleagues at the University of Tübingen in Germany, analyzed bone samples from 14 cave lions—found in four caves in France and central Europe—that lived between 12,000 and 40,000 years ago. The team focused on the chemical content of the bone collagen, which is often well-preserved, even in bones tens of thousands of years old. By incinerating a tiny fragment of preserved bone—usually less than a milligram—researchers can identify the molecules inside it and determine an animal’s diet. 

Scientists have perfected the technique over the years. It was used recently to look at the diet of Neandertals, but this is one of the first studies to use it to look at a nonhuman predator—and the analysis is now sensitive enough to look several steps down the food chain. This enabled Bocherens to determine not only what cave lions ate but also what their prey ate. And that made it possible to tell, for example, whether lions were targeting full-size cave bears or their more vulnerable cubs, because adults and babies eat different diets themselves. “There’s a difference between the [chemical] signal of adults and babies,” Bocherens says. “Babies drink the milk of the mother.” 

As it turned out, this distinction was important. Bocherens’s analysis, reported in the 6 December issue of Quaternary International, revealed that the cave lions occasionally ate bear cubs but not adults. Their favorite food, however, was reindeer, which Bocherens and his team determined consumed massive quantities of lichen, much as their modern descendants did. The cave lion diet, Bocherens says, appears to have been much more finicky than that of today’s lions, which eat just about anything they can catch.
 Grizzly  - Boss of the Woods.
        
  
             
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United States Polar Offline
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#49

From Ursus Arctos on Carnivora, Bengal Tiger vs Smilodon fatalis:


Quote:The study uses a ratio of lower jaw strength for absorbing forces from biting down ("Zx") to lower jaw strength for resisting torsional stresses (i.e. struggling prey) ("Zy"). A high Zx/Zy ratio shows either:

1) The prey was strongly secured by strong forelimbs before the killing bite was made

2) The prey was strongly secured by a group before a killing bite was made

3) Some combination of both

Some Zx/Zy ratios at the canines from this and another study by the same author:

Gray Wolf: 0.90

Lion: 1.25

Tiger: 1.35

Leopard: 1.37

Jaguar: 1.43

Cougar: 1.46

Clouded Leopard: 1.83

American Lion: 1.84

Smilodon Fatalis: 1.89

Smilodon Populator: 2.49

Therrien argues below that the american lion had much stronger forelimbs than modern lions and could likely restain prey securely. He also argues that small groups could have hunted together as well.

The Zx/Zycanine value of Panthera atrox (1.84; Fig. 5)
is much higher than that of other large felids and similar
to Neofelis nebulosa, indicating that dorsoventral stresses
prevailed in the symphyseal region and that labiolingual
and torsional stresses exerted by struggling prey were
relatively lower. While it is generally agreed that the large
body size of Panthera atrox allowed it to tackle very
large herbivores, such as bison, horses, ground sloths,
camels, and proboscideans (e.g. Kurt´en & Anderson,
1980; Harris, 1992; Anyonge, 1993), it seems paradoxical
for this Pleistocene predator to have experienced lower
torsional stresses than by extant lions that hunt smaller
prey (Schaller, 1972). Although extant felids use their
forelimbs and claws to restrain prey (Gonyea&Ashworth,
1975), their jaws must still be able to remain locked on the
neck or muzzle of prey and withstand the unpredictable
stresses induced as prey struggle to escape. If Panthera
atrox were a predator of large herbivores, one would
expect it to have Zx/Zycanine values similar to, or even
lower than, those of extant lions. Claw and tooth marks
left on a Pleistocene steppe bison mummy (Bison priscus;
M. L. Guthrie, 1988; R. D. Guthrie, 1990) suggest that
Panthera atrox adopted killing techniques similar to
those used by modern lions. However, Anyonge (1996)
has shown that the cross-sectional geometric properties
(i.e. bending strength) of the limbs of Panthera atrox,
particularly of the humerus, were much greater than
those of the extant lion, being closer to those of the
brown bear, Ursus arctos. In other words, the extinct
lion had much stronger forelimbs than an extant lion
of similar body size. 
Therefore, large prey could have
been primarily subdued and restrained by the extremely
powerful forelimbs of Panthera atrox, which would have
greatly reduced stresses on the mandible during the
canine bite. Furthermore, because the Zx/Zycanine values
of Panthera atrox are so high, it is possible that cooperative
hunting may have been common practice in that species,
where one or a few individuals would have restrained
a large herbivore while another delivered the canine
killing bite. Indeed, the high degree of cephalization
observed in Panthera atrox (Kurt´en & Anderson, 1980),
the claw and tooth marks left on a bison mummy, and
native American cave paintings (M. L. Guthrie, 1988;
R. D. Guthrie, 1990) suggest that the extinct lion
may have hunted in small groups of two or three
individuals, rather than in a pride. This possibility is
further supported by the size distribution of Panthera
atrox individuals in the Rancho La Brea deposits, which
indicates that the extinct lion did not form prides as
modern lions do but may have hunted in pairs or alone
(Jefferson, 1992).
"Be the reason someone smiles. Be the reason someone feels loved and believes in the goodness in people."

- Roy T. Bennett
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Brazil Matias Offline
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#50

I want to congratulate you for the high level of information in this Post, based on scientific evidence, with very interesting conclusions and questions.


We always try to understand the concrete facts with concrete, objective analyzes using a practical, conclusive directive. I believe that, like the other Pleistocene mammals, the felines Panthera Spelaea, Panthera Fossilis and Panthera Atrox have met a higher growth pattern due to a diversity of concrete factors and others entirely unmeasurable that will have no practical means of proving themselves. Most of the answers on the whys? Of its bigger size, it is in the other mammals contemporary to all these felines. I believe that the body volume of a predator meets a mathematical principle that determines its adaptive success. In simplistic terms we have: speed to reach prey, force to shoot prey, need to become hegemonic, or better adapted in its environment to face other top predators and organic / environmental ability to better synthesize foods. The great card, so to speak, is to understand the evolutionary process by which all the mammals of that time passed and to frame the variants of the "Panthera" family in this context. Nature did great evolutionary experiments and in the Panthera branch it was no different.


I have a more global, procedural view, where determinisms are never fully determinant. Understanding the multiplicities of variants in measures of skulls, teeth, bone structure, body mass etc is an unpleasant task. The more we try to understand these organic processes through which these "archaic lions" have passed, the more we realize that we still lack countless paleontological discoveries that can finally elucidate such questions. Let's wait for the analytical results on the two puppies of Panthera Spelaea.
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United States tigerluver Offline
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#51

I've updated the library. There are over 20 articles to read if interested. Those not linked I can probably get you if you PM me. The anatomical bank also gained 3 images.
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United States tigerluver Offline
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#52

With a few words, I made an indication that I found P. fossilis to have proportionately longer tarsal and carpals than other pantherines. Here's my basis. 

First I noticed that in Marciszak's figures, P. fossilis, both the early and middle form, consistently had longer metacarpal and metatarsals than P. spelaea. 

*This image is copyright of its original author


Thus, I would expect P. fossilis handful of long bone specimens to show the same trend. This was not the case. Something could have been different, namely the proportions of distal to proximal bones of the species. However, with a small sample size for P. fossilis, analysis is difficult. Nevertheless, I consulted the realms of biostatistics, utilizing the confidence interval. I tested two proportions, namely the ratio of femur length to tibia length and the ratio of femur length to metatarsal III (3) length.:


*This image is copyright of its original author

What you see above are the different ways I used the true average, the minimum 95% interval average, and the maximum 95% interval average for each bone measurement. By mixing and matching different intervals, I calculated different ratios. The then averaged these ratios to come up with a statistically most likely value for each ratio. This way, I attempted to make something more concrete out of a few pieces.

The results show that in all likelihood, P. spelaea and P. fossilis did significantly different in proportion. To account for the small sample size, I derived the p-value at a confidence level of 99% (alpha = 0.01), meaning there is only 1% chance these tests produced a false result, at least by the rules of biostatistics. As you can see above, both tests' p-values were comfortably below the cutoff for significance in different of 0.01, meaning nothing here is a borderline conclusions. 

Therefore, tentatively it seems P. fossilis was a longer legged cat than P. speleaa by quite a bit. Thus, it would have been a very, very tall cat. The fact that distal bones (tibia and metatarsal 3) are proportionately longer than proximal bones (femur) also indicates that P. fossilis was more cursorial than P. spelaea. By extension, as P. leo shares its ratios with P. spelaea, P. fossilis would have been quite proportionately taller and more cursorial than its closest living relative in P. leo.  


What is interesting to note is that despite this seemingly higher rate of cursoriality, P. fossilis does not seem to have given up robusticity. The giant 465 mm ulna I found to be about the same in robusticity as the American lion (P. atrox), a species with a femur to tibia almost the same as P. spelaea. Moreover, looking the humerus fragment, Thus, P. fossilis could be looked as a paradigm for the peak of evolutionary efficiency. It was a long limbed cat, well adapted for covering distance perhaps more effectively than the modern lion. Yet at the same time, it also had bones wider than the modern lion, indicating a powerful beast capable of bringing down the Pleistocene's big game. This combination is hard to find in any modern carnivore. All in all, it seems P. fossilis is unique. It had wide bones, part of which was used for increased muscle mass and another part which aided its long legs as they strode during its cursorial life. 

Another interesting aspect to note is that in the Marciszak et al. (2014), earlier P. fossilis consistently had longer metacarpals and metatarsals than the middle P. fossilis. Perhaps this could be used as a line of evidence for the chronological link between P. fossilis and P. spelaea, whereby the intermediate proportions of the tarsals and carpals in the middle P. fossilis is proof a transition period in the lineage. Perhaps these late middle Pleistocene P. fossilis are also the direct ancestors of P. atrox, explaining why P. atrox, while sharing skull characteristics with P. fossilis, did not share its proximal to distal bone proportions. Thus, the next step could be to investigate the climate and habitat changes that could have triggered the body proportion change.
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Canada GrizzlyClaws Offline
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#53

What does explain the unique skull characteristics of Panthera spelaea?

It is a significant morphological shift due the adaption or Panthera spelaea simply belonged a completely different lineage from Panthera fossilis?

Both Panthera spelaea and Panthera fossilis should belong to a lineage that diverged from Panthera leo back in 1.89 mya?
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United States Polar Offline
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#54

@GrizzlyClaws,

Would you say that the skull of the European Cave Lion is more robust than the modern African Lion skull in any way? The former seems to be near-equal with the latter in terms of width-to-length ratio.
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Canada GrizzlyClaws Offline
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#55

(01-14-2017, 10:27 AM)Polar Wrote: @GrizzlyClaws,

Would you say that the skull of the European Cave Lion is more robust than the modern African Lion skull in any way? The former seems to be near-equal with the latter in terms of width-to-length ratio.


The late Pleistocene Eurasian Cave lion did have a very wide muzzle like tiger, while the middle Pleistocene European Cave lion and the late Pleistocene American lion did have more consistent proportion with the modern lion.

Apart from the muzzle, Panthera spealea was not more robust than Panthera fossilis and Panthera atrox.
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United States Polar Offline
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#56

So in conclusion, the American Lion retained the extreme cursoriality as Fossilis but lost some of the bone robusticity (in leg bones)?

What can be said, though, is the loss of bone thickness isn't as significant. The American Lion still has a stronger humerus than a modern lion when considering size parity, and the former has longer, more cursorial bones than the latter.
"Be the reason someone smiles. Be the reason someone feels loved and believes in the goodness in people."

- Roy T. Bennett
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Canada GrizzlyClaws Offline
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#57

I think you just nail it.

And for odd reasons, Panthera spelaea just convergently evolved toward tiger. You can see the striking similarities between the bones of Panthera spealea and the robust late Pleistocene tigers.
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United States tigerluver Offline
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#58
( This post was last modified: 01-14-2017, 11:35 AM by tigerluver )

@Polar , I would say the opposite. The American lion dropped high cursoriality and retained bone robusticity. I say this with scant data, but Reichenau's P. fossilis ulna was about as robust as those found by Merriam and Stock, while P. atrox's MT3/Femur ratio of about 0.33 and femur/tibia ratio of 1.17 indicates that it was not as tall and cursorial as P. fossilis. Thick bones go far into the African Panthera lineages, remember the 111 cm wide distal end of P. shawi. 

@GrizzlyClaws , I'll get back to you on the skull issue. Barnett et al. (2016) came up with the 1.89 mya divergence using just a P. spelaea and then assumed/inferred P. fossilis falls on that lineage. Not a bad assumption as P. shawi is found around that time, but not guaranteed until P. fossilis DNA somehow gets found, extracted, and tested.
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Canada GrizzlyClaws Offline
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#59
( This post was last modified: 01-14-2017, 12:42 PM by GrizzlyClaws )

Well, it would be quite hard to extract the DNA from Panthera fossilis, since this species died too early. However, its daughter species Panthera atrox could be a good option if there is any fresh mummified carcass that is available in the places like Yukon and Alaska.

BTW, the discovery of those two mummified Cave lion cubs was like winning the lottery, and they were also quite recent, just the dawn before the early human civilization. And it did miraculously help the scientific community to solve the mystery of the Cave lion DNA. Not sure if they would be that lucky next time.
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Italy Ngala Offline
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#60
( This post was last modified: 01-18-2017, 02:37 AM by Ngala )

Interesting table with the measurements of the Atlas of various felines.

A Find of Panthera leo spelaea GOLDFUSS (Felidae, Mammalia) from unknown Stratigraphic Position of Potočka zijalka (Slovenia) Krofel, 2004

Abstract:
"The atlas of a large felid from a small unpublished collection of the 1928-1935 excavation in Potočka zijalka has been studied. It belongs to the cave lion (Panthera leo spelaea), most likely to a female. This is the highest elevation fi nd belonging to the said species in Slovenia and the second one in the Slovenian Alpine region."
"Man still bears in his bodily frame the indelible stamp of his lowly origin." C. Darwin
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