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tigerluver.animalvsanimal

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Dec 26 13 3:17 AM

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As my first significant contribution to the forum, I would like to initiate a discussion topic on P. spelaea.

To start off, I would like to discuss the body mass of the Early/Middle Eurasion Cave lion, Panthera spelaea fossilis.

There are three fossils on record of P. s. fossilis which exhibit its massive size, specifially:
1. A skull (Greatest length: 484.7 mm). Mentioned by Marciszak et al. (2013). 
2. An ulna (Total length 465 mm). Mentioned by Reichenau (1906).
3. A femur (Total length 470 mm). Mentioned by Teschler-Nicola (2006).

To set the stage, here is a short analysis of theoretical body masses associated with each bone. The following estimates are produced by equations formulated by Christiansen and Harris (2005) and Mazak et al. (2011).
- Skull: Mazak et al. only utilizes condylobasal lengths to derive its formula. Such a measurement is not available for the 484.7 mm skull. Though, assuming this skull holds the same proportions as other P. s. fossilis specimens on record, this skull's condylobasal length would be about 433 mm, resulting in a theoretical mass of 445 kg. Such a value is probably an overestimate, as the equations more often than not overestimates specimens, and significantly overestimates lions. 
- Ulna: This is from probably the largest individual on record. A 465 mm ulna is absolutely massive. The theoretical mass for this specimen in accord to Christiansen and Harris (2005) is 347 kg (the equation used here disregards Lynx lynx). The equation applied here estimates lion masses well, but underestimates tiger and jaguar masses significantly.
- Femur: In according with the equation by Christiansen and Harris (2005), the theoretical mass of this specimen would be 285 kg. This equation seems to result in underestimates across all pantherine species.

Obviously such estimation are not the final word on the mass of P. s. fossilis for the reasons already stated. In time, I will discuss other formulas that could better estimate mass.

Regards and happy holidays to all.

Last Edited By: GuateGojira Jan 22 14 4:07 AM. Edited 1 time

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#1 [url]

Dec 26 13 10:41 PM

Excellent new topic. This will be a very good new source of information for this great cat and any other lion-related specimens.

I will post my data here to about this cat. I am curious, which is the document of Reichenau (1906)? It is the only document of the three that you post here, that I don't have yet.

How are you draw skills? You can make your own images to. It will help to enrich the topic.

Greetings.

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#2 [url]

Dec 29 13 2:40 AM

The Skull

This post will shortly discuss the theoretical mass tied with the large skull (GSL = 484.7 mm, theoretical CBL = 433 mm). 


The Mazak et al. (2011) gave an estimate of 445 kg, and also seems to overestimate mass. The reasoning behind this is explained in Christiansen and Harris (2005), as follows:
"A data sample with many small species would introduce a size-related bias, producing unreliably high body mass estimates for large species."

Mazak et al. used the average body mass and condylobasal lengths of each specie as the database to derive the equation. Thus, from the sample size of 6 data points (n=6), 4 were representative of relatively smaller species (P. pardus, N. nebulosa, P. onca, and P. uncia) while 2 were representative of the large species (P. leo and P. tigris). Graphically, there was an uneven distribution of data points, with the smaller species being represent on one extreme and the large on another. Therefore, the data sample had too many small species relative to the amount of large species represented, and thus there was, "a size-related bias, producing unreliably high body mass estimates for large species" (Christiansen and Harris, 2005). 

Mazak et al. (2011) used a species averaged database to prevent confusion between intra- and inter-specific allometry. Though, in reducing the sample size, the distribution of data became uneven, causing the size-related bias mention above. 

I constructed a logarithmically scaled graph using the same database of specimens from Mazak et al. (2011), but had each individual specimen to represent a data point rather than a specie average representing a data point. This produced a plot with an even distribution of data points. The resulting equation:
log(body mass in kg) = 2.6725*log(condylobasal length in mm) - 4.4587

An implication of this equation is that skull size grows more rapidly than body mass. Furthermore, the data sample used can be more safely applied to P. spelaea as P. spelaea is a distinct species, rather than a subspecie of anomalous species in terms of relative proportions and body mass (e.g. P. t. soloensis to P. tigris), and thus one can assume P. spelaea follows the growth trend of Panthera in general. I realize the wording in this paragraph may be a bit confusing, so just ask if any further clarification is needed on the point I am making.

Finally, the equation discussed yields a theoretical body mass for the 484.7 mm skull of approximately 387 kg.

Guate: The Reichenau (1906) document is one I received working with a professor. The title is: Beitrage zur naheren kenntnis der Carnivoren aus den Sanden von Mauer und Mosbach II.

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GrizzlyClaws

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#3 [url]

Dec 29 13 3:52 AM

Panthera spelaea fossilis must have a proportionally very long and narrow skull, while i can imagine that Panthera spelaea spelaea having a body proportion closer to that of tiger.

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#4 [url]

Jan 11 14 6:01 AM

Following with the estimation of weight of the giant Cromerian lion (Panthera spelaea fossilis), using my database and the same Excel sheet that I used for the Ngandong tiger (see the Ngandong tiger topic for details), I calculated the body mass for the skull specimen and the femur one.

 

Here are the results, average of all the calculations:

* Skull (CBL) - 433 mm: 510.3 kg

* Femur (GL) - 470 mm: 367.1 kg

 

Interesting, the skull-weight is way too high, just like a bear. With such a large skull, I will estimate a figure of maybe up to 400 kg and at simple sight, but 500 kg? Surely NO, is like you say, it seems that skulls grow in a different way than the body overall, so the body weight estimation of skulls could be over or under estimated depending of the size of the specimen.

 

Now, with the femur, the estimation of C&H (2009) was of 349.3 kg, using all Panthera but only males. The estimation of Sorkin (2008) was of 385 kg (modern lion CBL of 359.7, femur of 401.5 mm and a maximum weight of 240 kg). The final average of 367.1 kg is very high, but with such massive animals it is be expected.

 

Well, this is the data that I have found; I think that this will help you to establish the weight of this giant cat, just like we have done with the Ngandong tiger.

 

I will put a new list of the skulls of the Cromerian and the Eurasian Steppe lion, actualized with the new data from the document of 2013.

 

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GrizzlyClaws

Posts: 120 Member Since:relative

#5 [url]

Jan 11 14 7:03 AM

The longest scientifically recorded femur for Panthera leo is 401.5mm, while the longest scientifically recorded skull is 419mm.

So Panthera leo got its longest recorded skull longer than longest recorded femur, same as Panthera spelaea fossilis.

These lion-clade cats have proportionally longer skull than the tiger-clade cats. 

Last Edited By: GrizzlyClaws Jan 11 14 7:05 AM. Edited 2 times.

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#6 [url]

Jan 11 14 7:37 AM

The Femur

GuateGojira wrote:

Following with the estimation of weight of the giant Cromerian lion (Panthera spelaea fossilis), using my database and the same Excel sheet that I used for the Ngandong tiger (see the Ngandong tiger topic for details), I calculated the body mass for the skull specimen and the femur one.

 

Here are the results, average of all the calculations:

* Skull (CBL) - 433 mm: 510.3 kg

* Femur (GL) - 470 mm: 367.1 kg

 

Interesting, the skull-weight is way too high, just like a bear. With such a large skull, I will estimate a figure of maybe up to 400 kg and at simple sight, but 500 kg? Surely NO, is like you say, it seems that skulls grow in a different way than the body overall, so the body weight estimation of skulls could be over or under estimated depending of the size of the specimen.

 

Now, with the femur, the estimation of C&H (2009) was of 349.3 kg, using all Panthera but only males. The estimation of Sorkin (2008) was of 385 kg (modern lion CBL of 359.7, femur of 401.5 mm and a maximum weight of 240 kg). The final average of 367.1 kg is very high, but with such massive animals it is be expected.

 

Well, this is the data that I have found; I think that this will help you to establish the weight of this giant cat, just like we have done with the Ngandong tiger.

 

I will put a new list of the skulls of the Cromerian and the Eurasian Steppe lion, actualized with the new data from the document of 2013.

 


Exactly why I have left usage of isometry out. Note, Dr. Christiansen also used regression in his latest document to calculate mass. I'm sure you saw how some species gave irrationally high estimates for the skull, such as the jaguar. The size limit for Pantherines seems to be in the low four hundred kilograms. This makes sense, as where one finds cats, one finds bears. Bears rules the 500+ kg size range, and thus rule the niche that comes with such size. Evolutionary, it would not be beneficial for cats to grow to the size of bears and compete for the same niche. S. populator seems to make it to the mid four hundred kilogram range. I have some comparisons of S. populator bones to bears to show their size.

The femur estimate you got is similar to the one I have found with regression. I assumed that P. spelaea had a build midway between tigers and lions and thus based the regression off a database of only tigers and lions. The database for the formula is based off of 6 specimens, the equation:
log(mass) = 3.6775*log(femur length) - 7.2568
The 470 mm femur would have a mass of 371 kg accordingly.

In all these estimates, if P. spelaea was truly lion it would be a bit lighter as lions have long bone dimensions with relatively less mass out of the Pantherines, around 350 kg.

Lastly, the Sorkin method is a bit unreliable because it mixes and matches masses and bone dimensions. For example, Christiansen and Harris (2005) have a lion weighing 203 kg with a femur length of 401 mm. According to Sorkin's method, he would use the 401 mm length with the greatest mass of lion on record, not 203 kg. Here, one can see the problem. Each extant specimen is like a trial. One can't mix and match data from two separate trials. This explains why Sorkin calculated overestimates for the specimens he assessed, as he used a 320 kg mass as the mass of the bone of the cat which did not weight nearly as much. Maybe for the sake of comparison between fossils this method is okay, but in terms is accuracy, not so much. For that reason, I would consider the Sorkin estimation of the femur invalid. A 401.5 mm femur clearly does not represent a 240 kg, so the basis of comparison is wrong. Furthermore, the Sorkin method is redundant with the isometry method. Averaging the Sorkin result based on only one specimen with the result of isometry applied across many specimen is giving the one base specimen used in the Sorkin method unreliable influence on the average mass. My recommendation is to stick to one method, not average different methods. Formulas are independent. New formulas are derived in hopes to gain accuracy, so I avoid mixing up the results of new and old methods, as doing so kills the purpose of finding more accurate methods of mass estimation.

Last Edited By: tigerluver Jan 16 14 10:10 PM. Edited 5 times.

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#7 [url]

Jan 11 14 9:39 AM

The Ulna

Finally, I will go over the ulna in this short post. 


As I stated before, an ulna of 465 mm is certainly from a record breaking specimen. To predict the body mass without encounter false negative allometry, I again used a database of tigers and lions, with six specimens in total. The equation:
log(mass) = 2.8965*log(ulna length) - 5.1318

The R-squared value was .9, weaker than my other equations. This is because the tiger and lions are significantly different in ulna to body mass proportions, with the former being relatively heavier. Again, I assumed P. spelaea fossilis had a built between the tiger and the lion. The resulting estimate, 393 kg. Putting the ulna into perspective with the Ngandong tiger femur, this specimen probably had a femur of 480 mm as well, give or take. Its mass would be slightly less than the Ngandong specimen (as this specimen is classed as a member of the tiger species, c. 409 kg) again assuming it was not built like a tiger, rather midway between lions and tigers. 

I am looking into evidence to help figure the built of P. spelaea. Two things support it being very lion-like in built, if not synonymous, genetic data and robusticity of the bones, which fall into the range of modern lions. Furthermore, it is likely P. spelaea was morphologically lion-like as both species lived in similar, open landscapes, calling for greater cursoriality, explaining the relatively great width of the long bones.

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GrizzlyClaws

Posts: 120 Member Since:relative

#8 [url]

Jan 11 14 11:03 AM

Their (spelaea and atrox) teeth are also quite lion-like.

A big head Amur tiger measured by Dr. Gewalt has a 50cm head (assuming with 43-45cm skull), and its canine teeth from the gumline were 9cm.

But look at that large spelaea specimen with approximately 45cm skull, it got barely over 7cm from the skull, not from the gumline.

image

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GrizzlyClaws

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#10 [url]

Jan 11 14 11:12 AM

Both male specimens of spelaea and atrox got void within their canine teeth.

image

image

image

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#11 [url]

Jan 13 14 9:35 PM

P. spelaea spelaea body mass

Body mass estimates of the skull measurements Guate collected and published. The estimates are based on equation used for the large P. s. fossilis skull. Again, assuming P. s. spelaea had a heavier built than modern lions, midway between tigers and lions. These estimates would be slight overestimates if P. s. spelaea was more or less identical to modern lions.


image

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#12 [url]

Jan 15 14 11:44 PM

The P. spelaea species seems to have similar or the same long bone proportions as modern lions. Here is an example (epicondylar index = Distal Width/Femur length):

Species Femur length Distal Articular Width Epicondylar Index
P. leo 401.5 83.1 0.206973848
P. leo 401 87.1 0.217206983
P. s. spelaea 419 90 0.214797136
P. s. fossilis 470 99.2 0.21106383

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GrizzlyClaws

Posts: 120 Member Since:relative

#13 [url]

Jan 16 14 9:15 PM

Here is new the study paper about the Panthera spelaea species.

- Now Panthera atrox is considered to be directly evolved from the more archaic Panthera fossilis, not from the recent Panthera spelaea.
- Panthera spelaea did not evolved from Panthera fossilis.
- Panthera fossilis did live in Asia, so those large remain of Panthera spelaea found in Russia were likely belong to Panthera fossilis.

http://onlinelibrary.wiley.com/doi/10.1111/1749-4877.12082/abstract

Last Edited By: GrizzlyClaws Jan 17 14 5:56 AM. Edited 3 times.

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#14 [url]

Jan 17 14 5:51 AM

Great find GrizzyClaws. If anyone need the full article, please PM me.

This excerpt changes a lot of ideas that we have gone by:
image

I remember GrizzyClaws proposed a splitting Panthera further into the tiger and lion clads. This document brings up an interesting point that lion-like cats originated from Africa. On the other hand, the tiger's origins are in East Asia. The two clads may be more distant than we previously though.

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GrizzlyClaws

Posts: 120 Member Since:relative

#15 [url]

Jan 17 14 6:00 AM

So far there four subspecies for Panthera spelaea.

- Panthera spelaea fossilis
- Panthera spelaea atrox
- Panthera spelaea spelaea
- Panthera spelaea vereshchagini

But the author even suggests to rename Panthera youngi as Panthera spelaea youngi, which i strongly disagree. Since this great cat indeed bears some similarities with the Panthera spelaea species, but the recent discover shows this cat has tiger-like short facial profile. I think it is more likely an extinct tiger-clade member.

Last Edited By: GrizzlyClaws Jan 17 14 6:03 AM. Edited 3 times.

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GrizzlyClaws

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#16 [url]

Jan 17 14 6:05 AM

Also, Panthera spelaea spelaea and Panthera spelaea vereshchagini formed a same inner branch within the Cave lion species.

While Panthera spelaea fossilis shows more affiliation with Panthera spelaea atrox.

So there are two inner branch within the Cave lion species; the spelaea-vereshchagini branch and the fossilis-atrox branch.

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#17 [url]

Jan 17 14 7:35 AM

I may have missed something, but I think that the author is actually classifying the Cromerian lion and the Upper Cave lion as distinct species. They also cite another work (Hanko and Korsos, 2007) as evidence. Though, they still encompass both these species into the lion-clad, as well as Panthera youngi. So the classification according to this author would be:
Lion-clad:
-P. spelaea
-P. fossilis
-P. atrox
-P. youngi

The author is classifying P. youngi as part of the lion-clad due to it mandible features. But I also have some doubt about this. Some of the recent P.t. sondaica mandibles have slight lion-like features, but at the same time look a lot like the P. youngi mandible. Maybe the tiger's concave mandible is a more recent trait. Then again, P. fossilis and P. spelaea both expanded to the end of east Asia, so it would be plausible for another lion-like specie to make it to China.

But how plausible is the question? P. youngi coming from Africa would have to compete the daughter species of P. zdanskyi, which had been around eastern Asia for much longer, and therefore, probably much better adapted to out-compete an alien species from a different region. I wouldn't expect a new cat specie of similar size having much success here, and its time in the vicinity of P. zdanskyi's daughter species would be short lived. It would make more sense for P. youngi to one of P. zdanskyi daughter species than an one whose roots are not native to the area.

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GrizzlyClaws

Posts: 120 Member Since:relative

#18 [url]

Jan 17 14 7:55 AM

Fully agreed, East Asia (especially China) seems to be fully dominated by the tiger-clade members.

Also, the squarish mandible is not an exclusive feature for lion-like cats, since all Pleistocene tigers did have this archaic feature, which even extends into some modern tigers.

Moreover, the recent discovered skull of Panthera youngi shows the tiger-like facial profile.

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GrizzlyClaws

Posts: 120 Member Since:relative

#19 [url]

Jan 17 14 8:01 AM

Panthera spelaea (Beringian lions) seemed to have stopped right after the northern shore of the Amur River, they didn't cross the river thus penetrating into the Manchurian woodland.

Also, the Panthera spelaea specimens near Manchuria didn't evolve with impressive size, so i believe they might not be very successful to live near tiger's domain.

Same for tiger, those lived in Beringia and Alaska also didn't become dominant species in the region.

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#20 [url]

Jan 17 14 8:21 AM

For two pantherine species to coexist, they would have to have different niches, which would be governed by their relative sizes. That's why tiger and leopards coexist and the same can be said for the African great cats. P. spelaea and P. tigris are of comparable size, whichever species got to a specific region first would probably inhibit the other from expanding into the same region.

Nevertheless, maybe tiger and lion clad species never overlapped because of their evolutionary adaptations rather than niche clashes. The lion-clad started out in the open savanna of Africa, and such species would evolve to be long legged and cursorial, therefore limiting their expansion to areas of only open terrain. On the other hand, the tiger lineage evolved in more wooded areas, calling for stouter bodies, not a beneficial feature for regions which would require greater cursoriality.

These new studies give more support to notion that P. fossilis and P. spelaea were lion-like in built, so soon I'll give some estimates of body mass with lion proportions.

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