The Cave Lion (Panthera spelaea and Panthera fossilis)

[Ghari Sher]

(12-08-2018, 06:51 AM)Ghari Sher Wrote:

(12-06-2018, 05:53 AM)tigerluver Wrote:

(12-06-2018, 03:49 AM)Ghari Sher Wrote: Just a little query here:

According to some written correspondence by Mauricio Antón and Manuel Salesa, the cave lion had very short distal limb elements and metapodials compared to the African lion. They cite Ballesio (1980), a text which is not accessible to me, to support this claim.

Quote:Here we need to take into account the differences in limb proportions between Pleistocene lions and modern ones: the former are known to have relatively shorter distal segments in their limbs, especially the notoriously short and robust metapodials (Ballesio, 1980).

http://www.academia.edu/16968107/Walking...oichnology

I haven't been able to find any other mention of the cave lion having shorter distal limbs and metapodials than the African lion (though it wouldn't be surprising seeing as the same is seen in the cave hyena as compared to its extant African counterparts), Sabol 2018 doesn't state that the Medvedia lion's metapodials were particulrly short either. Is there any evidence of this being the case?
I know @tigerluver has done some calculations regarding the limb proportions of the European cave lion, maybe that could shed some light on this issue?
From reading that paper you recently linked, it doesn't seem to have had especially distinctive limb proportions, relatively speaking.

I don't think lost in translation would be exactly the right phrase to use, but I believe the issue is caused by the loss of context when describing the bones. Cave lions have "short and stubby" metapodials in the sense that they are very thick for their length so they look "short". The work seems to have missed this contextual point and took the description in a bone to bone context. In proportion to the length of the other long bones, their metapodials are not short in terms of metapodial to long bone length ratios.

Why they have thick metapodials is up in the air. It may be partially to account for their increased robusticity in the rest of their body. Having wide feet would serve as good snow shoes as well.

Hmmm... I see what you mean.
Speaking of wide paws - Protopopov 2016 describe the same condition existing in the cub remainsm their paws are proportionally larger than in African lion cubs. I'm not sure what interpretation can be derived from there.
https://www.researchgate.net/publication...ct_Species

But the robustness of the metapodials really does fit in with what I have kept reading about the cave lion - its robusticity compared to the African lion. Previous publications on P. atrox's morphology have pointed out that its forelimbs were considerably more robust than in an extant lion today, but little is said about the cave lion, which is just subsumed under the "lion" category due to it similar limb proportions proportions far too long.

I recently saw Wheeler & Jefferson (2009)'s paper on P. atrox, and alongside bone measurements he included what he called the "Index of Robustness" or IR, essentially a measure of the robusticity of a longbone which is a function of its length relative to its shaft width. A lower IR=more robust. I decided to calculate that for limb bones in the cave and African lion and see how they (alongside the American one) measured up.
I used some of the cave lion material in Sabol (2018)'s supplementary document that you kindly shared, as well as that of the African lion which, alongside those of the cave lion, I collected from various other works too, including some rather old ones.
Humeri and femora were the available bones to me, and while Wheeler and Jefferson (2009) used the longitudinal width of the femoral shaft of P. atrox in his IR calculations, that measurement was not widely available for the other two cats (particularly the cave lion), and so I re-calculated the IRs of the P. atrox femora using the transverse width (which was also provided by Wheeler & Jefferson (2009) for the femora and was the only shaft width recorded for the humeri).
I didn't separate the groups out to account for sexual dimorphism, though I'm quite certain that there would be some bimodality in the robusticity distributions in each cat, but the cave lion material wasn't at all sexed, and only the P. atrox femora were sexed. But we can talk about that later.

What I found surprised me.

*This image is copyright of its original author

*This image is copyright of its original author

(Made using MS excel)
For both the humeri and the femora, P. spelaea and P. atrox surpassed the extant lion in regards to robusticity, but P. spelaea seems to have surpassed even P. atrox, at least in regards to the mean IR (crossed), but overall their mean IRs seem fairly similar. The material available really shows the overlooked morphological distinctions between the cave lion and the African lion, as well as between it and the American cat.
The cave lion seems to have had very thick longbone shafts, at least in regards to the humeri and femora.

Indeed, using these measurements and others, I recently formulated some mass estimates for the European cave lion (the Siberian and Beringian forms seemed to have differed in size so their remains were not included).
I used Christiansen and Harris (2005)'s equations to calculate the weight, and the label of the equation is indicated in the below table.
Now in regards to sex.... some of the longbones were already sexed, mainly those from Diedrich's papers, as well as Sabol's. Others were inferred to be male due to their immense size (e.g. the Mladeč femur), and in regards to sorting out the longbones with measured shaft width (which Diedrich does not measure in his papers), I had to infer male/female identity from their lengths in comparison to longbones of known sex. Perhaps that was a bit iffy of me, you be the judge of that, but the results seem to look OK.

*This image is copyright of its original author

The masses derived from humerus articular width, and the mean masses that include these are highlighted in red, since I'm quite skeptical of the accuracy of those figures, they're far greater than the figures from other measurements, and I'm highly doubtful that any cat bar the very largest Smilodon populator individuals could ever hope to attain a mass of around 453.94kg. Hence, in black at the bottom I include mean masses that do not factor humerus articular width in.  Overall, I got a mean mass of 237.96kg for males and 172.21kg for females.
Without factoring in the width of the femoral and humeral shafts, the mass estimate for both sexes goes down to closer to ~220kg for males and ~165kg for females. The cave lion had some noticeable thick limbs, no doubt.

As for that very last column in the table "Percentage Dimorphism", let me explain.
In the literature and on online forums, there are a number of authors who state that the sexual dimorphism present in the cave lion was greater than that today, namely (and I think firstly) Baryshnikov & Boeskorov (2001), and a number of others after them. They calculated that male cave lions were 21% larger than females, which is apparently higher than in the African lion's size difference of only 15%. This same claim, or something similar, has been repeated by later authors such as Sabol (2018).

There are a number of problems with their claim.

Firstly, Baryshnikov & Boeskorov (2001) worked this "21%" figure from the mass estimates they got for Beringian cave lions based on dental measurements -154kg for females, 194kg for males. Simple math shows that assuming this figure to be true, their 21% figure is refuted, as 194/154=1.2597... essentially the males are 26% larger than females, not 21%. What he means is that females are 21% smaller than males, since 100*(1-(154/194)) = ~21%.
Secondly, Both the 2001 authors and Sabol (2018) cite George Schaller's 1972 book The Serengeti Lion in support of their "male African lions are 15% larger than females" figure. As someone who has a digital copy of that book, I can find no mention whatsoever of this 15% figure. Schaller does, however, mention weights of African lions taken in the field. Using Baryshnikov & Boeskorov (2001)'s definition of this male-female size dimorphism, I get much greater levels of dimorphism than is claimed both for them and their Pleistocene relatives. Also looking at other authors, I see the same pattern.

*This image is copyright of its original author

Except for one. The outlier of 12.2% seems to be attributable to the smaller and thereby more biased sample size of lionesses (n=5), where as a more representative sample of males (n=14) was used. 
Disregarding that, I calculated disparities of 25.8-33.8% (mean 30.55%) for differing samples across Africa using the data from Smuts (1980). Combining Schaller (1972) with Smuts (1980)'s database I get an overall mean of a 29.7% disparity between the sexes. This is greater than the dimorphism calculated by Baryshnikov & Boeskorov (2001) for the Beringian cave lion, particularly when you realise that their mass estimate for male Beringian lions is virtually the same as the mean weight of extant Rhodesian males (tabled), but the corresponding females are much smaller than they would be under the calculated dimorphism of the Beringian lion.

The problem is that African lions are highly dimorphic in terms of mass, much more so than is claimed/realised by the cave lion authors. Females are roughly 0.7 times the mass of males, or, as Baryshnikov & Boeskorov (2001) would unaptly put it, males are 29.7% larger than lionesses.

What was the case for the cave lion?
I'm a bit doubtful that the mass estimates by Baryshnikov & Boeskorov (2001) are accurate, since they only used teeth to estimate mass, so the puported dimorphism may be off, a suspicion which is raised by my mass results.
As you can see/ work out from the tables, in P. spelaea (at least the European ones), lionesses are estimated at about 0.73 times the mass of males, or, as Baryshnikov & Boeskorov (2001) would put it, males are approximately 27.42% larger than females. This size disparity is, of all things, smaller than what is seen in extant P. leo, where lionesses are about 0.7 times the mass of males and males are approximately 29.7% larger than females, but not greatly so, and I can't help but attribute this rather minor difference in size dimorphism simply to imprecisions in my own calculations/ smaller sample sizes available. In all honestly, it appears that sexual dimorphism in P. spelaea was similar to that seen in extant P. leo.

In older forums, such as on Tapatalk, I have heard claims to the effect that European late Pleistocene lionesses were no bigger than those of Africa today, and that it was only the males that exceeded extant African males in size by about 10%.
I find this to be false, cave lionesses seem to have exceeded African lionesses in size by a similar extent to the male cave lions dwarfing the African ones. In this instance, male cave lions seem to have been a whopping ~32.2% larger than extant ones, as were the lionesses, give or take.

Indeed comparing them to extant Bengal tigers, where males average around 208kg and females 135, the former species seems to fall short by a margin, though I'm sure the earlier Holocene/Pleistocene tigers would have reached much more sizeable proportions that today with their better access to prey,etc.
https://www.scribd.com/document/55287778...Tiger-2015
(I hope @GuateGojira doesn't mind me posting links to his stuff here, I can't find any other reliable figures for Bengal tiger mass)

In any case, the cave lion was a massive cat indeed.

Thoughts/feedback would be appreciated.

Just a little update to the mass estimation, I realised that I used the equations for olecranon length by substituting in the "maximal width" of the ulna that Diedrich had listed in his tables.
I had assumed they equalled the olecranon length that is weightable by the equations of Christiansen & Harris (2005). If so, then it gave a severe underprediction of 172kg in males and 116kg in females, though given that cave lions probably did not differ this drastically in terms of their ulna proportions when compared with modern cats, I'm going with the conclusion that "maximal width"=/= "olecranon length". In any case, I excluded the Olecranon length and Humerus articular width rows from the table to give (what I think is) a more accurate mass estimate.

*This image is copyright of its original author

I got an average weight of 247kg in European male cave lions and 181kg for an average lioness. This is absolutely huge, no cat today reaches such average sizes.
By comparison, here's what I was able to get for P. atrox, bone measurements from Wheeler & Jefferson (2009), though I did have to sex a few myself, perhaps that was a bit iffy of me, but in any case, here's what I got using the same equations:

*This image is copyright of its original author

I got a mean weight of 264kg for male and 193kg for female P. atrox. Still bigger on average than European P. spelaea, but the size disparity is much less that I had previously figured, with P. atrox averaging about ~7% larger than European P. spelaea.

However, as you can see from the P. atrox table, the 264kg/193kg weight was averaged from a number of different measurements, not all of whom were available for P. spelaea (and vice versa).
I'm inclined to use these weights as being closer to the real-life weights since they take into account more of the animals' proportions/measurements, but for an oranges-to-oranges comparison I'll use the weights derived from the 6 (widely) available measurements in common between the two cats (Humerus length, Humerus diaphysis LM, Femur length, Femur diaphysis LM, Ulna length, Tibia length), then the mean male mass for P. atrox goes down to 257kg and for P. spelaea it marginally climbs to 249kg, with male P. atrox being ~3% bigger than European P. spelaea.

In any case, it seems that P. atrox was the bigger cat, but not by much; European P. spelaea and P. atrox likely did not differ significantly in size. Reply