The Largest Beasts to Walk the Earth

It seems that at least once a year, news of the “largest dinosaur ever found” makes headlines. That’s because almost all the contenders for the biggest saurian are so fragmentary that settling on an uncontested victor is almost impossible, not to mention that new discoveries are being made all the time. There hasn’t been as much of a debate about fossil mammals. For over a century, Paraceratheriuma 26-foot-long, 15 ton, hornless rhino – has been cited as the biggest of the big beasts. But, according to a new paper by Asier Larramendi, ancient elephants are in close competition for the title of the largest mammals to ever walk the Earth.

Such contests rely on how they’re measured, of course. Part of what makes Paraceratherium seem so large is its elongated neck. In terms of tonnage, however, Felisa Smith and colleagues pointed out that the largest prehistoric proboscideans – such as the anchor-tusked Deinotherium – were likely heavier. (It’s similar with sauropods – does Supersaurus hold the title of Biggest Dinosaur because of it was a bit longer, or does Argentinosaurus because it weighed more?) Giants are made by how you measure them.  In this case, Larramendi drew from photos, measurements, and reconstructions of fossil and living elephants to come up with estimates of body mass and shoulder height.

Starting with the bones, Larramendi created virtual restorations of 24 different elephant species. The point, aside from trying to correct errors in articulation that are sometimes made when these animals are mounted in museums, was to restore the elephants as accurately as possible in order to estimate their volume and, from that, their body mass. Some researchers use a different methodology for these sorts of estimates – focused on estimating how body shape changes with size – but Larramendi prefers the volumetric method.

Reconstructions and size estimates for extinct elephants. From Larramendi, 2015.
Reconstructions and size estimates for extinct elephants. From Larramendi, 2015.

Larramendi selected elephants from various times in their 60 million year record. The very early Eritherium azzouzorum, for example, may have weighed 11-13 pounds and been a little more than half a foot high at the shoulder. From such a small start, elephants eventually attained truly enormous sizes. Larramendi calculates the largest Deinotherium weighed 13 tons and stood over 13 feet high at the shoulder. Others were bigger still. The mastodon Mammut borsoni, Larramendi calculates, would have weighed over 15 tons and been almost 13 and a half feet tall at the shoulder, and, based on some scrappy material, Larramendi estimates that the elephant Palaeoloxodon namadicus would have reached 24 tons and an imposing 16 feet at the shoulder.

From Larramendi, 2015.
From Larramendi, 2015.

Part of a femur isn’t much to rest a title on, but, if Larramendi is correct, the largest Palaeoloxodon would have been nearly a foot taller and over 5 tons heavier than the biggest known Paraceratherium. But even sticking to more complete skeletons, Larramendi’s estimates suggest that ancient elephants could get as large as, if not a bit larger than, the previous record holder.

Calling this competition doesn’t interest me, though. Whether a rhino or elephant was the largest land mammal of all time isn’t especially important. What’s curious is that there isn’t a clear winner between the two lineages. (There can’t be until better skeletal material gives us more precise dimensions for these giants.) Just as with the largest dinosaurs, the most superlative mammals of all time seem to cluster around a similar body size. This, as I believe the SV-POW!sketeers have pointed out, might tell us something about the upper limits of body size in these groups. The next step is to see if we can parse out what these limits were and how they altered the shape of life for millions of years, and what those constraints might mean for the future of evolution.

Reference:

Larramendi, A. 2015. Shoulder height, body mass, and shape of proboscideans. Acta Palaeontologica Polonica. doi: 10.4202/app.00136.2014

5 thoughts on “The Largest Beasts to Walk the Earth

  1. Very interesting blogpost, as usual!
    But as many of your readers are not from the USA, please use the metric system, or at least provide both systems!
    Especially when discussing an article about body sizes, that makes it difficult to follow.

  2. This is based on a distal portion of a femur… That the author didn’t even check as he says: ” Of course, it would be of special interest to revise the Indian Museum of Calcutta collection where these fossils might be stored, in order to study and confirm in particular the colossal size of the distal femur part.” This is NOT how we do real science, we go to the museums to check real specimens and even more with the important ones such as this one.
    Also the only referred giant femur is from Prinsep (1834), with 63 inches. But the femur is not complete, and maybe more important: in 1834 an inch was not an unvariable measure unit and it was slightly different across the world. And that’s why we measure the old specimens to check their measurements. That’s why Amphicoelias fragillimus is also a very doubtful animal. Again, taking a measurement from 181 years ago as absolute truth is a huge mistake, and is not how science should be done.
    Also, taking one small fragment of bone and scaling to a complete set of measurements (shoulder height) is not a good method. Mainly due to allometric relationships between organic parts. Just take a look at different species of mammoths, they do not share the same proportions. So Why this elephant is special?
    Also, the distal part of the femur IS NOT the femur that Prinsep (1834) talk about! This distal portion of femur is nowhere to be found in literature.
    In synthesis, this titanic elephant is the Amphicoelias fragillimus of the mammals. More likely, a myth. But a useful myth, thus we can see to gigantic sized mammals, and question their size. Of course that it can be the case of a proboscidean larger than indricotheres, but this particular case is very, very poorly based on evidence.
    Just don’t buy the conclusions so fast.

  3. Reference:
    Mikael Fortelius, “The largest land mammal ever imagined,” Zoological Journal of the Linnean Society vol. 107 (1993), pp. 85-101. (PDF available on the WWW, courtesy of the “Rhino Resource Center”.)
    Take-home message (from second-to-last paragraph): “Based on these data, it would be difficult to argue convincingly that the largest indricotheres were larger than the largest Proboscideans — both seem to have been slightly more than twice as heavy as the largest African elephants, the heaviest living land mammals.”

    Scrupulously noting that whales are aberrant Artiodactyls, it is noteworthy that Perissodactyls and Proboscideans have produced species significantly larger than the largest (known) Artiodactyls. Is this evidence that hind-gut fermenting is more effective, at very large body sizes, than the ruminants’ way of processing food?

  4. Hello,

    I would like to clarify some doubts on the described material by Prinsep (1834). First, when an author or a group of authors, study a large group of animals in order to estimate their body mass, speed, etc… based on bone measurements, it is not possible to check all the material presented in the study. See for example Paul (1997), Christiansen (2004), O’Gorman and Hone (2012) and a large etc…

    These authors largely based on bibliography for their calculations. Of course, it is strongly recommended to check personally as much material as possible, but when is possible (I personally checked some of the largest remains such as Mosbach M. trogontherii humerus, M. borsoni Milia fossils, giant Stegodon…). For example, Christiansen (2004) based many of his calculations on other author published measurements. I personally corrected him on Taubach femur (he estimated 14.5 tons for this animal, based on unreliable measurement). Through my paper I corrected many errors of past authors, but unfortunately I was not able to check Prinsep (1834) personally, but my obligation as a researcher was to report it.

    Many often, fossil remains are not possible to check, but that is not an impediment to perform studies based on these remains (A clear example is Spinosaurus).

    Prinsep (1834) knew very well how much was a inch (2.54 cm), as anyone can check it in his paper when he translate a 1.11 m femur length to 43.7 inches. On the other hand, I checked the measurements and photos on both femora, in a vector graphics editor program. The results matches perfectly with mm as well as the femur proportions, the other illustrated E. maximus femur and the scale bar of the image. So, the measurements are reliable. Moreover, the measurements given by Prinsep are in perfect accordance with other recently described complete similar sized femur by Meshram and Sonakia (2006). I even lowered Prinsep estimation to 155 cm in order to be conservative (His estimation is also well possible). Finally the gigantic part was said to be nearly one quarter by Prinsep (1834), but I was conservative and assumed 20% larger. It is the best approach with the evidences we have in this moment. Please note that I also report other remains of similar size class in my paper, so a elephant of this size was possible.

    If we consider P. namadicus as the same case as A. fragillimus (which I may agree), we also have to do the same with Indricotherium-Paraceratherium. During the last 80 years, the max. size of the giant hornless rhino was inferred from just a proximal part of an MC III (AMNH 26175), and no one apparently has questioned it. Most complete associated remains of these animals suggest a body size of 8-11 tonnes (See Granger and Gregory 1936; Paul 1997; Fortelius and Kappelman 1993; Qiu and Wang 2007). A body size considerably less than a good number of quite complete preserved proboscidean skeletons.

    Also would like to point that the size of the largest individuals of many dinosaurs have been estimated from very fragmentary remains (For example: Spinosaurus, Magnapaulia, Torvosaurus, Argentinosaurus, Puertasaurus, Xenoposeidon, Triceratops, Eotriceratops, Mamenchisaurus, Apatosaurus, Giraffatitan, Alamosaurus, Stegosaurus, Ankylosaurus, Therizinosaurus, Giganotosaurus and a very very long etc…) is the daily bread of the researcher focused on estimating the body size of extinct forms.

    The shoulder height estimated for a 185 cm for a Palaeoloxodon is conservative. Concerning to mammoths as one pointed out, the ratio between skeletal shoulder height to femur length is practically the same in the largest different species (see my paper). The same is for Palaeoloxodon, this ratio for P. antiquus and P. Recki (Koobi Fora), is exactly the same (2.56), and should be noted that in the largest elephant individual, this ratio tend to increase because the allometry (see my paper). So, applying a very conservative ratio of 2.55 to P. namadicus we obtain 472 cm of skeletal SH and 498 cm in the SH in the flesh as a result.

    My estimation for the largest individuals are a rough approximation only (the same was also for Indricotheres for the last century), but based on solid remains, proboscideans were larger than indricotheres.

    References:

    Christiansen, P. 2004. Body size in proboscideans, with notes on elephant metabolism. Zoological Journal of the Linnean Society 140: 523–549.

    Fortelius, M. and J. Kappelman 1993. The largest land mammal ever imagined.
    Zoological Journal of the Linnean Society 108:85-101.

    Granger, W. and Gregory, W. K. 1936. Further notes on the gigantic extinct rhinoceros, Baluchitherium, from the Oligocene of Mongolia. Bull. Amer. Mus. Nat. Hist. 72, 1-73.

    Larramendi, A. 2015. Shoulder height, body mass, and shape of proboscideans. Acta Palaeontologica Polonica. doi: 10.4202/app.00136.2014

    Meshram, S.N. and Sonakia, A. 2006. Fossil femur of a giant elephant (cf. Elephas
    namadicus) from middle Pleistocene, Central India. Indian Miner 60 (3–4): 199–202.

    O’Gorman EJ, Hone DWE 2012. Body Size Distribution of the Dinosaurs. PLoS ONE 7(12): e51925. doi:10.1371/journal.pone.0051925

    Paul, G.S. 1997. Dinosaur models: the good, the bad, and using them to estimate the
    mass of dinosaurs. In: D.L. Wolberg, E. Stump, G.D. Rosenberg (eds.), DinoFest
    International Proceedings, 129–154. Academy of Natural Science Philadelphia,
    Philadelphia,

    Prinsep, L. 1834. Note on the fossil bones on the Nerbudda valley discovered by Dr.
    G.G. Spilsbury near Narsinhpur. Journal of the Asiatic Society of Bengal 3: 396–403.

    Qiu, Z.-X. and Wang, B.-Y. 2007. Paracerathere fossils of China. Paleontologica Sinica 193 (29): 1–396.

  5. Hello,

    I would like to clarify some doubts on the described material by Prinsep (1834). First, when an author or a group of authors, study a large group of animals in order to estimate their body mass, speed, etc… based on bone measurements, it is not possible to check all the material presented in the study. See for example Paul (1997), Christiansen (2004), O’Gorman and Hone (2012) and a large etc…

    These authors largely based on bibliography for their calculations. Of course, it is strongly recommended to check personally as much material as possible, but when is possible (I personally checked some of the largest remains such as Mosbach M. trogontherii humerus, M. borsoni Milia fossils, giant Stegodon…). Christiansen (2004) based many of his calculations on other authors published measurements. I personally corrected him on Taubach femur (he estimated 14.5 tons for this animal, based on unreliable measurement). Through my paper I corrected many errors of past authors, but unfortunately I could not check Prinsep (1834) personally, but my obligation as a researcher was to report it (In the future I’ll do my best to check the remains personally).

    Many often, fossil remains are not possible to check, but that is not an impediment to perform studies based on these remains (A clear example is Spinosaurus).

    Prinsep (1834) knew very well how much was a inch (2.54 cm), as anyone can check it in his paper when he translate a 1.11 m femur length to 43.7 inches. On the other hand, I checked the measurements and images on both femora, in a vector graphics editor program. The results matches perfectly with mm as well as the femur proportions, the other illustrated E. maximus femur and the scale bar of the image. So, the measurements are reliable. Moreover, the measurements given by Prinsep, are in perfect accordance with other recently described complete similar sized femur by Meshram and Sonakia (2006). I even lowered Prinsep estimation to 155 cm in order to be conservative (His estimation is also well possible). Finally the gigantic distal part (reported by Prinsep (1834; see fig.11)) was nearly one quarter larger than the preceding femora (Prinsep 1834), but I was conservative and assumed a 20% larger. It is the best approach with the evidences we have in this moment. Please note that the large described femora belonged to an still growing elephant, and I also report other remains of similar size class in my paper, so a elephant of this size was possible.

    If we consider P. namadicus as the same case as A. fragillimus (which I may agree), we also have to do the same with Indricotherium-Paraceratherium. During the last 80 years, the max. size of the giant hornless rhino was inferred from just a proximal part of an MC III (AMNH 26175), and no one apparently has questioned it. Most complete associated remains of these animals suggest a body size of 8-11 tonnes (See Granger and Gregory 1936; Paul 1997; Fortelius and Kappelman 1993; Qiu and Wang 2007). A body size considerably less than a good number of quite complete preserved proboscidean skeletons.

    Also, I would like to point that the size of the largest individuals of many dinosaurs have been estimated from very fragmentary remains (For example: Spinosaurus, Magnapaulia, Torvosaurus, Argentinosaurus, Puertasaurus, Xenoposeidon, Triceratops, Eotriceratops, Mamenchisaurus, Apatosaurus, Giraffatitan, Alamosaurus, Stegosaurus, Ankylosaurus, Therizinosaurus, Giganotosaurus and a very very long etc…) is the daily bread of the researcher focused on estimating the body size of extinct forms.

    The shoulder height estimated for a Palaeoloxodon with a femur length of 185 cm, is conservative. Concerning to mammoths as one pointed out, the ratio between skeletal shoulder height to femur length is practically the same in the largest different species (see my paper). The same is for Palaeoloxodon, this ratio in P. antiquus and P. Recki (Koobi Fora), is exactly the same (2.56), and should be noted that in the largest elephant individuals, this ratio, tend to increase because the allometry (see my paper). So, applying a very conservative ratio of 2.55 to P. namadicus, we obtain 472 cm of skeletal SH and 498 cm in the SH in the flesh as a result.

    My estimation for the largest individuals are a rough approximation only (the same was also for Indricotheres for the last century), but based on solid remains, proboscideans were larger than indricotheres.

    References:

    Christiansen, P. 2004. Body size in proboscideans, with notes on elephant metabolism. Zoological Journal of the Linnean Society 140: 523–549.

    Fortelius, M. and J. Kappelman 1993. The largest land mammal ever imagined. Zoological Journal of the Linnean Society 108:85-101.

    Granger, W. and Gregory, W. K. 1936. Further notes on the gigantic extinct rhinoceros, Baluchitherium, from the Oligocene of Mongolia. Bull. Amer. Mus. Nat. Hist. 72, 1-73.

    Larramendi, A. 2015. Shoulder height, body mass, and shape of proboscideans. Acta Palaeontologica Polonica. doi: 10.4202/app.00136.2014

    Meshram, S.N. and Sonakia, A. 2006. Fossil femur of a giant elephant (cf. Elephas namadicus) from middle Pleistocene, Central India. Indian Miner 60 (3–4): 199–202.

    O’Gorman EJ, Hone DWE 2012. Body Size Distribution of the Dinosaurs. PLoS ONE 7(12): e51925. doi:10.1371/journal.pone.0051925

    Paul, G.S. 1997. Dinosaur models: the good, the bad, and using them to estimate the mass of dinosaurs. In: D.L. Wolberg, E. Stump, G.D. Rosenberg (eds.), DinoFest International Proceedings, 129–154. Academy of Natural Science Philadelphia,

    Prinsep, L. 1834. Note on the fossil bones on the Nerbudda valley discovered by Dr.
    G.G. Spilsbury near Narsinhpur. Journal of the Asiatic Society of Bengal 3: 396–403.

    Qiu, Z.-X. and Wang, B.-Y. 2007. Paracerathere fossils of China. Paleontologica Sinica 193 (29): 1–396.

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