Thursday, December 15, 2011

Eocetus, "Eocetus", and Friends

Update (January 28, 2014): "Eocetus" wardii is now Basilotritus wardii. More on my new post, The Third King.

I was shocked that Uhen (2010) remarked that Basilosaurus drazindai and Basiloterus hussaini "probably represent protocetids... akin to Eocetus". This would place the whales outside Pelagiceti and imply that the now-questionable basilosaurids were potentially capable of walking on land, despite being enormous. Unfortunately, other mentions of this revised placement give no further details (Uhen 2008, Uhen et al. 2011) and Uhen (2010) further states the placement is "difficult to determine with certainty" due to scarce materials. I suspect the hypothesis will not be officially discussed until further material is found and/or described... which won't stop me from wildly speculating.

Lumbar vertebrae in right lateral view. From left to right: "Eocetus" wardii (from Uhen 1999), Basiloterus hussaini, and Basilosaurus drazindai - note that the latter-most may be an anterior caudal (from Gingerich et al. 1997). For comparison: Basilosaurus isis vertebrae.
In the description of Basilosaurus drazindai, Gingerich et al. (1997) note a number of "primitive retentions" which resemble the morphology of "generalized archaeocetes": long neural spine and arch; broad, almost-horizontally placed, anterior-projecting metapophyses which project beyond the anterior edge of the vertebral centrum; and paired, posterolateral processes of the neural arch. Aside from the last trait (which I can't confirm without a dorsal view), all of these traits are present in "Eocetus" (Uhen 1999). Additionally, "Eocetus" has elongated transverse processes, unlike the condition of Basilosaurus (Uhen 1999); however, B. drazindai has processes with a 15.5 cm long base (they broke off) relative to the 30 cm centrum (Gingerich et al. 1997), and so probably had a similar, albeit slightly less extreme, condition. The only criterion for placing B. drazindai in the genus Basilosaurus was the size and shape of the centrum (Gingerich et al. 1997), and while they are uncannily similar in shape, everything else seems to be pointing towards a relationship with "Eocetus".

Lumbar vertebrae in anterior view. Ditto order.
As for awkward middle-child Basiloterus, it appears to have a centrum which is slightly more elongated than that of "Eocetus", however the neural arch and maybe the neural spine appear to be narrower. The metapophyses are upwardly-angled (Gingerich et al. 1997), less broad, less anterior-projecting, but still appear to extend past the centrum. The posterolateral processes are absent (Gingerich et al. 1997). The base of the transverse process is 9.3 cm long relative to a 19.5-20 cm centra (Gingerich et al. 1997), proportionally similar to Basilosaurus drazindai. The placement of Basiloterus is thus not clear, and perhaps it was a basilosaurid or an even more derived protocetid.

Maiacetus inuus, a basal "protocetid" (Uhen 2011). From Wikipedia Commons.

Protocetidae is a blatantly paraphyletic "family" of extinct cetaceans from Eocene coastal marine deposits with hip and femur morphology indicating amphibious capabilities (most of the time) and no evidence of flukes (Uhen 2010). Uhen (1999) appears to think that "Eocetuswardii had weight-bearing hips, however Uhen (2010) refers to them as "moderately reduced" and regarded the species as possibly non-amphibious. This is perhaps not surprising since Eocetus, "Eocetus", and an unnamed Pisco Formation species are the sister group of Pelagiceti (Uhen et al. 2011). This could make them closer relatives of Dorudon than Maiacetus, and raises the question of how many typical protocetid traits they actually exhibited. Perhaps they were entirely aquatic tail-based swimmers which just happened to have fairly large vestigial legs.

Dorudon atrox. From Wikipedia Commons.
The scare quotes around "Eocetus" hint at a taxonomic misadventure. "E." wardii was assigned to its genus by Uhen (1999) based on comparisons of its skull and vertebrae to Eocetus schweinfurthi; the problem is, the holotype of E. schweinfurthi is an isolated skull and it is not possible to determine whether the vertebrae referred to it actually represent the species (Geisler et al. 2005). There is overlapping skull material (Uhen 1999), but Geisler et al. (2005) apparently regarded it as too incomplete to warrant unambiguous placement in the genus. Somehow, "Eocetus" and Eocetus formed a clade in phylogenetic analyses (Geisler et al. 2005, Uhen et al. 2011), making it probable that future discoveries will confirm their close relationship.

"Eocetus" wardii is clearly related to unnamed Pisco Formation material which exhibits the same distinctive traits (moderate centrum elongation, elongated neural arches and spines and transverse processes, strange pock-marked texture) with the only difference being that the unnamed material is 35% smaller (Uhen et al. 2011). The Egyptian vertebrae dubiously assigned to Eocetus schweinfurthi (figured in Uhen 1999) also seem quite similar (including the pock-marks), and if it is also a member of this clade, it would indicate a sizable trans-oceanic range. This in turn could be taken as evidence of the whales being largely pelagic... of course this is quite speculative.

There of course remains much to be known about these cetaceans, and perhaps future discoveries will be enlightening as to how similar they were to the pelagic cetaceans, as well as the origins of Pelagiceti. I really hope it turns out that a Basilosaurus-sized animal could walk on land.


Geisler, J. H., Sanders, A. E., and Luo, Z-X. (2005). A New Protocetid Whale (Cetacea: Archaeoceti) from the Late Middle Eocene of South Carolina. American Museum Novitates 3480, 1-65. Available.

Gingerich, P. D., Arif, M., Bhatti, M. A., Anwar, M., & Sanders, W. J. (1997). Basilosaurus drazindai and Basiloterus hussaini, new Archaeoceti (Mammalia, Cetacea) from the middle Eocene Drazinda Formation, with a revised interpretation of ages of whale-bearing strata in the Kirthar Group of the Sulaiman Range, Punjab (Pakistan). Contributions from the Museum of Paleontology, University of Michigan 30 (2), 55-81. Available.

Uhen, M. D., Pyenson, N. D., Devries, T. J., Urbina, M., and Renne, P. R. (2011). New middle Eocene whales from the Pisco Basin of Peru. Journal of Paleontology 85(5), 955-969. doi:

Uhen, M. D. (2010). The Origin(s) of Whales. Annual Review of Earth and Planetary Sciences 38, 189–221. Available.

Uhen, M. D. (2008). Basilosaurids. In: Perrin, W. F., Würsig, B., and Thewissen, J. G. M. (eds.) Encyclopedia of Marine Mammals, Second Edition. Elsevier: Burlington, Massachusetts. Available.

Uhen, M. D. (1999). New Species of Protocetid Archaeocete Whale, Eocetus wardii (Mammalia: Cetacea) from the Middle Eocene of North Carolina. Journal of Paleontology 73(3), 512-528.

Weems, R. E., Edwards, L. E., Osborne, J. E., and Alford, A. A. (2011). An occurrence of the protocetid whale "Eocetus" wardii in the Middle Eocene formation of Virginia. Journal of Paleontology 85(2), 271-278. Available.


Allen Hazen said...

Slightly tangential question: how, in Cetacea derived enough that there is no longer a "Sacro-iliac"connection such as is found in land mammals, do you distinguish lumbar from "anterior caudal" vertebrae? I have looked at mounted skeletons of modern cetacea, and failed to spot any obvious transition in form. ... Eventually you get to centra that have chevrons below them (below and between...), and I take it that these ought to be classified as caudal--but are ONLY the centra with chevrons caudals?
With enough soft anatomy (which I'm not sure we have!), you could determine which vertebrae are closest to the innominate remnants, and they might be fair candidates for classification as sacra's, particularly if (is this the case?) they have some sort of ligamental connection to the vertebrae. ... Mounted skeletons typically have the hind-limb remnants suspended beneath the backbone, and I have wondered whether their exact location (which vertebrae are immediately over them) are known or guess-work!
(B.t.w.: your cetacean blogs are appreciated even when not commented on!)

Cameron McCormick said...

Yep, the vertebrae with chevrons (or "ventral/haemal midline processes") are caudals. While some early workers thought they could distinguish sacrals (I remember Kellogg listed Basilosaurus with them), they're just lumped in with lumbars now. There doesn't seem to be any way to determine where the hindlimbs of fossil whales with unattached innominates are, other than presumably floating somewhere right before the tail.

Whales of the World and Buchholtz and Schur(2004) have pretty good reviews that are freely available.

Cameron McCormick said...

Oh, and it means a lot that somebody out there appreciates these! At some point I'll write about Basilosaurus, but I keep on running into these other interesting whales...

Allen Hazen said...

BAsilosaurus, humbug! I learned about it from Herbert S. Zim's "The Great Whales" when I was a small child in the 1950s, and Zim called it Zeuglodon. So, ICZN and priority be damned, I'll always think of it as Zeuglodon.
And Merry Christmas to you! (Thank you for the references, which I will study after Christmas.)

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