Body size of the Ngandong tiger (Panthera tigris soloensis)
From GuateGojira:
Data about the great Ngandong tiger (Panthera tigris soloensis) is fully accounted by Von Koenigswald (1933), however, at this time, I have been unable to found the original paper, which by the way, is in German. This is the paper:
Koenigswald, G. H. R. Von, 1933. Beitrag zur Kenntnis der fossilen Wirbeltiere Javas. Wet. Meded. Dienst Mijnb. Ned. Ind., no. 23, 184 pp., 28 pis.
Maybe Peter could be able to found it, but he was not here anymore, sadly.
Now, the next two documents quote the measurements of only two parts of the fossils, the skull and the humerus. So, here we go.
1. Skull:
Brongersman (1935) made an excellent account of fossils of Panthera tigris trinilensis and Panthera tigris oxignata, however he made a little quote about the skull of a specimen of P. t. soloensis, regarding the ratio existing between the greatest length of the skull and the zygomatic width. Check that the name Feliopsis palaeojavanica is invalid and is know now as P. t. soloensis (Mazák, 2010). Here is the full account:
“Feliopsis palaeojavanica was described by Stremme (1911a, p. 56; 1911 b, p. 86, pl. 16 figs. 3, 4, pl. 17 fig. 1) from rather scanty remains. More complete specimens were studied by Von Koenigswald (1933, p. 6) who arrived at the conclusion that the genus Feliopsis could not be separated from the genus Felis in its wider sense. Von Koenigswald compared the fossil specimens to different species and subspecies: Panthera tigris sondaica (Temm.), Panthera tigris tigris (L.), Panthera leo (L.) and Panthera leo spelaea (Goldfuss). In the first place this author compared the ratio existing between the greatest length of the skull and the zygomatic width. This ratio is calculated by him to two points of decimals, but I believe this to be unnecessary as the greatest length of the fossil skull examined by Von Koenigswald is not exactly known. The greatest length actually measured is 373 mm, but according to Von Koenigswald 15 or 20 mm must be added for the part of the occiput which was broken off; in his calculations, however, he used 380 mm for the greatest length, thus adding only 7 mm. The ratio calculated from this measurement (380 mm) is 1.58 (in Von Koenigswald's paper it is given as 1.52; small differences are also found between some of the other calculations made by this author and those made by me) ; if we do add 15 mm to the greatest length actually measured, the ratio becomes 1.61, and if 20 mm is added it becomes 1.63.”
Source: http://www.repository.naturalis.nl/document/149407
A direct approach shows that the only “more or less” full skull of the Ngandong tiger is incomplete, and is estimated a greatest length of 388 – 393 mm. The zygomatic wide is probably of c.250 mm, if we use the ratio of 1.52 which is the original figure given by Koenigswald, which work with the original fossils. In this fashion, I believe that this skull had a greatest length of 390 mm and a zygomatic wide of 250 cm. It is slightly larger than the largest Amur tiger skull reported by Mazák (1981; GL 383 mm, ZW 268 mm) and among the largest tiger skulls in record. However, we must remember that other fossils are even larger (although there are more fragmentary) and until I can found the original paper of Koenigswald, I can’t give more details.
2. Humerus:
The next came from Hooijer (1947), which work with Chinese tigers from the Pleistocene, including the now famous Panthera tigris acutidense. Here is the account:
“The measurements given by von Koenigswald (1933, p. 12) indicate the presence of still larger specimens in the Pleistocene of Java. One humerus is even stated to have a greatest length of 381 mm., others 353 mm., which is about the maximum I found in the lion (No. 5: 352 mm.) and that in a fossil humerus from Chou Kou Tien (Teilhard de Chardin, 1936, p. 15: 355 mm.). In P. tigris altaica (Temminck) the maximum length is 328 mm., and in the other recent tiger humeri it is 311 mm. at the most.”
Source: http://digitallibrary.amn...nov/N1346.pdf?sequence=1
To understand the size of this humerus, I quote the first table in the first post of this topic:
http://animalsversesanima...ta---weight--tige?page=1
See that of Christiansen& Harris in 2005. The largest tiger (CN5697) had a humerus length of 372.5 mm. Check that this is also the largest humerus in the list. In the second table, of 2007, this same tiger have a shoulder height of 120 cm, so is possible to make a rough estimation and state that a tiger with an humerus of 381 mm would be slightly taller, maybe c.125 cm, which is about the same shoulder height of P. atrox.
3. Analysis:
Using these two fossils, from two distinct specimens, is possible to achieve an idea of how large was P. t. soloensis. The skull was just slightly larger, but even this specimen could be relative larger than modern Amur tigers, which have a maximum head-body length of 208 cm in straight line (Kerley et al., 2005). If can be assumed that the head-body length of a great cat is about five times its greatest skull length (based in a single specimen of Panthera atrox), the estimated head-body length for this specimen of P. t. soloensis could be of 195 cm (390 * 5 = 1950 mm) in the skeleton. However, Christiansen & Harris (2007) state that the average difference of the head-body length measured in the skeleton and the flesh, in tigers, is of 37.1 cm (n=3), here is the calculation:
HB-flesh HB-skeleton Diference
CN5698: 2040 mm 1653.3 mm 386.7 mm
CN5697: 2060 mm 1636.5 mm 423.5 mm
CN6049: 1950 mm 1647.0 mm 303.0 mm
Average: 371.1 mm or 37.1 cm.
Then, based in this result, the head-body length, in the flesh, of this particular P. t. soloensis would be of 232 cm. This is a bit higher, and most be taken in count that the relation “skull length-body length” is based in a single specimen. A large male Bengal tiger with a greatest skull length of 381 mm, had a head-body length of 213 cm “between pegs” (Ward, 1914; Brakefield, 1993); this gives a ratio “skull length-body length” of 5.6, flesh included. Based in this, the head-body length of this specimen of P. t. soloensis could be of 218.4 cm, a more reliable figure as is based in a real specimen, not reconstructed skeletons. It is necessary to make more comparisons, but even then, it is show that this particular specimen was relative longer than modern tigers.
About the humerus of 381 mm, it was obtained a simple relation between “humerus length-shoulder height”. Based in both tables of Christiansen & Harris of 2005 and 2007, it can be calculate that the average ratio for tigers is of 3.12, here is the data:
H-length S-height Ratio
CN5697: 372.5 mm / 1200 mm = 3.22
CN6049: 360.5 mm / 1180 mm = 3.27
CN5698: 350.0 mm / 1000 mm = 2.86
Average: 3.12
So, it can be estimate a shoulder height of c.119 cm. for the humerus of 381 mm, based in an average ratio of 3.12 (with the maximum ratio, the height will be of c.125 cm).
4. Conclusion:
Although the comparative data is based in only to pieces, is possible to achieve a relative good idea of how large was the Ngandong tiger:
* Skull: 390 mm GL and 250 mm ZW.
* Humerus length: 381 mm.
* Estimated head-body length: 218 – 232 cm (depending of the method).
* Estimated shoulder height: 119 cm.
About the weight, I can’t make any direct estimation as I don’t have the diameter of the humerus or the condylobasal length of the skull, however, it is known that the largest tigers (that reached this body sizes) weighed from 260 to 320 kg, so is plausible to estimate an average weight of 290 – 300 kg for these specimens of P. t. soloensis (remember that larger, although fragmentary fossils, exist).
I will appreciate any educated comment or even better, if someone can found the original document of Koenigswald.
Well, based in facts, the Smilodon populator is the real champion of the cats in the weight issue. I have found this document (check table 1):
http://www.app.pan.pl/archive/published/app51/app51-407.pdf
The data on it is based in the weight estimation of Christiansen & Harris (2005), so the average is reliable. According with this, the average weight for Smilodon populator was 304.45 kg (range 220-400 kg), the largest on record. However, check that this average includes male and female specimens together. For comparison, the average weight for modern Bengal tiger would be of 162.9 kg (males of 200.4 and females of 125.4 kg; according with my data). Modern lions would be of 150 kg (males of 175 and females of 125 kg; the figure for males is from my data, the figure from females is from Yamaguchi).
About Macairodus kabir, yes, its weight was inflated as it was used the formula of Anyonge (1993). However its fossils are large and it was possible that it were of the same weight that the largest Pantherines, but surely lighter than the S. populator.
Finally, about the size of the fragmented fossils of Panthera tigris soloensis, I have no idea of the size, just the descriptions of Brongersman (1935), Hooijer (1947) and now Hooijer (1949), which I have found here:
http://www.rhinoresourcecenter.com/pdf_files/117/1178933675.pdf
Check that he states that in the Pleistocene Java AND Sumatra have been found fossils of the same size that the modern Indian tiger. He also states that the Wanshien tiger (Panthera tigris acutidense) was as large as the recent Indian subspecies, BUT its metapodials were more massive than those of the recent tiger; “a character which the Plesitocene Chinese tiger has in common with the fossil tiger from Siberia described by Tscherski (1892).”
To understand what this means, we must remember that the metapodials are the long bones of the hand (metacarpals) and feet (metatarsals) which connect the digits to the centers. In humans, five are present in each hand and foot. This bones support much of the weight of the animal.
With this evidence we can establish that if this bones are more massive in P. t. acutidense than in the modern Indian tiger (weight from 170-260 kg), it seems that, even when they were of the same size (like state Kitchener (1999), they were obviously heaver, possible reaching up to 300 kg or more, like the 1900-1940 Manchurian tigers.
The document says more: “The metapodials of the Javan Pleistocene tiger are more slender than those of the Pleistocene continental tiger”. This “Pleistocene tiger” was surely the “Punung tiger”, which was much smaller than P. t. soloensis and was possibly the direct antecessor of the modern Javan tiger (Panthera sondaica sondaica; according with Mazák and Groves (2006)), which according with Hertler & Volmer (2008), weighed between 140 and 189 kg. Is important to mention than these weights were estimated in base of Van Valkenburgh (1990), which use dentition and skull measurements, instead of limb dimensions (for details see Volmer (2005)).
This paragraph from Mazák and Groves (2006) is very interesting:
“The evolution of mainland and Javanese tigers was summarized in a diagram by Groves (1992), who argued that the Late Pleistocene ancestor of the modern Javan tiger, P. t. soloensis, was completely different from its Early/Middle Pleistocene predecessors (which it presumably replaced), and already had the modern Javan form’s high M1 index and narrow occiput. The evidence indicates that the now extinct Javan tiger must also be assigned to a distinct species, Panthera sondaica, and to which the Ngandong tiger also belongs.”
In other words, it seems that the tiger present in Java (Panthera tigris oxygnatha) was replaced by the new arrival of the continental tiger at the Sonda shelf (possible descendents of Panthera tigris acutidense). The result of this replacement (or mix) was P. t. soloensis (or better, Panthera sondaica soloensis), with the large size but with some characteristics of the already present Pleistocene Javan tigers (described by Mazák & Groves (2006)). With time, the large size disappeared, given place to a new form and more specialized species of cat, the modern Sonda tiger (Panthera sondaica).
Is interesting to mention that Groves state that P. t. oxignata was more primitive than P. t. trinilensis, but the strata and the fossil’s age shows otherwise (Aziz et al. 1995; Meijaard, 2004).
So, we can estimate the size of Panthera tigris soloensis and Panthera tigris acutidense more or less accurately, but until we can found the original paper of Koenigswald (1933), I can only use the two fossils described in this topic (the skull and humerus).
Again, here is the name of the document, if someone can found it, will be EXCELENT!!!
* Koenigswald, G. H. R. Von, 1933. Beitrag zur Kenntnis der fossilen Wirbeltiere Javas. Wet. Meded. Dienst Mijnb. Ned. Ind., no. 23, 184 pp., 28 pis.
Incredibly, but this seems right. Check that the best weight estimation (Wheeler & Jeferson, 2009) for the female Panthera atrox estate an average of 178 kg (N= 32; range 142-210 kg). It is larger than any modern female cat, but not as large as those from males. It seems that the sexual dimorphism was stronger in the prehistoric specimens.
For comparison, the skulls of the Eurasian steppe lion (Panthera leo spelaea) were of the same size than modern African lionesses. So, they were no larger, only males were 8-10% larger than modern lions.
The bear dogs (like Anphicyon ingens) were indeed larger and heavier than any modern or prehistoric cat. Check the fossils, they were massive. These animals surpassed the mark of 500 kg and I even think that our best great cats were no match for them in a single combat.
Panthera leo spelaea, or steppe Eurasian lion, like Dr Diedrich says, is regarded as been 8-10% larger than modern lions (Patterson, 2004). The largest skull at this day measured 420 cm in greatest length (Diedrich, 2007) [it came from Siegsdorf, Germany], which is about the same size that the largest modern African lions. Wikipedia, in French, mention a skull of 430 cm from England, but don’t quote ANY sources for it.
Here is a list of all the skulls of P. l. spelaea that I have found:
Specimen | Greatest length (mm) | Condylobasal length (mm) | Zygomathic wide (mm) | Source |
MCSNB 5127 (♀) | 324.2 | 295.6 | 225.1 | Bona, 2005 |
Pocala 25260 | 370 | 354 | 264 |
Pocala 25262 (♀?) | 311 | 311 | 225 |
Pocala 1929, U Bologna | – | 360 | 253 |
AZE.K.13.29 | 417.4 | 368.6 | 283.4 |
Gailenrenth | – | 362 | 276 |
Herm | – | 350 | 245 |
Cajarc | 393 | 350 | 265 |
Vence | 354 | 320 | 241 |
K-1, Kondakovka Kolyma | 345.1 | 326.2 | 249 | Sotnikova & Nikolskiy, 2005 |
IPBPS-1, Duvannyi Yar, Kolyma | 359.4 | – | – |
GIN-1123, Smolensk Russ. Plain | 360 | 328 | 238.2 |
Beroun skull (♀) | 302 | – | – | Diedrich, 2007 |
Srbsko Chlum-Komín (♀) | 305 | – | – |
German Perick Caves (♀) | 302 | – | – |
Arrikrutz | 380 | – | – |
Siegsdorf | 420 | – | – |
Sandford Hill Cave | – | – | 231 | Dawkins et al., 1866 |
Hotton Cave | 336 | – | 234 |
Sundwig Brithis Museum (♀?) | 304 | – | – |
Vence (Alpes-maritimes) | 360 | – | – | Wikipedia (French), 2011 * |
England | 430 | – | – |
* This skulls are quoted with no reference, so they are unreliable.
♀ Specimens estimated to be females.
As you can see, they were just slightly larger than modern lions. Dr Diedrich shows many images and pictures of real complete skeletons of this great cat. I will put all of them here (even when this is not a topic about this cat) to show they sizes, this is thanks that Dr Diedrich put a size bar with all of them.
About the weight, based in skull and bone sizes, I estimate that this “lions” weighed the same than modern Bengal tigers, which is about 170-260 kg.
The Beringian lions (Panthera leo vereschagini) were slightly smaller than it Eurasian relative (Kurtén, 1985), so is possible that they were of the same size than modern African lion, with weighs of 130-240 kg.
This is my appreciation based in real fossils and not popular myths and web-pages.