All of these creatures were assumed then to belong to sea lions: most of them were large, and relatively robust, and differed markedly in many respects from true seals. Most of the early descriptions of pinnipeds we now know to be walruses are seemingly obsessed with comparing them to sea lions, and no doubt early workers such as Kellogg were frustrated with the alien nature of many of the fossils: they were about the same size as sea lions, but there were just so many little differences. The answer would not come until much later when decidedly modern researchers like Charles Repenning realized the true walrus affinities of many of these critters (like Pliopedia and Dusignathus).
Unlike the rather large and aberrant jaws of Allodesmus kernensis and Dusignathus santacruzensis, the fossil jaws from San Diego looked like a perfect match for a modern sea lion or fur seal: it had a shallow jaw with small triangular cuspate teeth, and the jaw was rectangular (i.e. the dorsal and ventral margins are parallel). The larger jaws of other better known pinnipeds had too many specialized features to be ancestral to sea lions: Dusignathus had widely flaring jaws without incisors and a canine that projected anteriorly, while Allodesmus lacked cusps on its postcanine teeth, which looked instead like little onions or bulbs. Burleson (1948) assigned these specimens to Pithanotaria, despite the much older age of Pithanotaria starri material described by Kellogg (1922) and the lack of actual morphological characters that identified the jaw of Pithanotaria. Burleson (1948) thought this specimen had a morphology intermediate between Pithanotaria and the modern Northern Fur Seal, Callorhinus.
single rooted, cuspate teeth.
In a much later paper by the preeminent paleo-pinnipedologist Charles Repenning and carnivoran researcher Richard Tedford (1977), this specimen was briefly discussed and they concluded that it did not represent Pithanotaria and was likely much closer to Callorhinus ursinus. After observing trends within the dental evolution of walruses like Imagotaria, Repenning and Tedford (1977) had identified the utility of the stage of root fusion as a taxonomic guide. For example, all modern otariids (fur seals and sea lions), the walrus, and some seals have single rooted teeth, while primitive pinnipeds and terrestrial carnivorans retain a number of double and triple-rooted teeth. For whatever reason, these root lobes coalesced through time and resulted in single rooted teeth in a number of taxa.
The dentary of the holotype of Callorhinus gilmorei, from Berta and Demere 1986.
Repenning and Tedford (1977) were surprised that Burleson (1948) had not noticed the interesting configuration of the tooth roots of this specimen: the third and fourth premolars and the molar were still double rooted, while only the first and second premolars were single rooted; in the modern Northern Fur Seal, Callorhinus ursinus, all the lower premolars and molar are single rooted. It was indeed a fur seal, but retained some interesting primitive features.
In 1986, after an extensive excavation of a bonebed in the San Diego Formation that would be christened the Mission Hills Bonebed, additional remains of this fossil pinniped were discovered including several jaws, teeth, skull fragments, and postcranial bones. The discovery of a partial skeleton of an immature female skeleton allowed Annalisa Berta (San Diego State University) and Tom Demere (San Diego Natural History Museum) to describe the San Diego fur seal as a new species - and sure enough, they found that its features placed it as a close relative of the modern Northern Fur Seal, Callorhinus. They named it Callorhinus gilmorei, named after Dr. Raymond Gilmore. In other regards, Callorhinus gilmorei was a relatively small fur seal - substantially smaller than modern skeletal remains, with less strongly developed cusps on postcanine teeth, and a more 'primitive' state of root fusion.
The new specimen of Callorhinus gilmorei from the Rio Dell Formation of Northern California described by Boessenecker (2011)
Subsequently, Kohno and Yanagisawa (1997) reported a tiny partial jaw from the late Pliocene of Japan. This jaw exhibited double rooted cheek teeth (although the anterior premolars were not preserved), and had accessory cusps on the cheek teeth, so they identified it as Callorhinus gilmorei. This extended the range of the Gilmore fur seal to the western Pacific - similar to the range of the modern Callorhinus ursinus.
Needless to say, I had a few ideas to follow once I started looking into Bushell's fur seal specimen. Fossils of C. gilmorei had so far only been found in Middle to Late Pliocene deposits, whereas in the late Miocene and earliest Pliocene of Japan, California, and Mexico the earlier fur seal Thalassoleon occurred (which has all double rooted teeth, and lacks cuspate cheek teeth, among other differences). The new specimen only has one cheek tooth - but it has a well developed accessory cusp, like C. gilmorei, and the first two premolars are both single rooted - also like C. gilmorei. In addition, it is relatively small - many other modern otariids are substantially larger. Furthermore, C. gilmorei appears to be the only middle-late Pliocene otariid in the entire Northeastern Pacific fossil record, which made the identification process somewhat easier.
Next up: Other fossil otariids from California and Oregon, and the Pleistocene Callorhinus
specimen.
References:
Berta, A., and T. A. Demere. 1986. Callorhinus gilmorei n. sp., (Carnivora: Otariidae) from the San Diego Formation (Blancan) and its implications for otariid phylogeny. Transactions of the San Diego Society of Natural History 21:111–126.
Boessenecker, R.W. 2011. New records of the fur seal Callorhinus (Carnivora: Otariidae) from the Plio-Pleistocene Rio Dell Formation of Northern California and comments on otariid dental evolution. Journal of Vertebrate Paleontology 31:2:454-467.
Burleson, G. L. 1948. A Pliocene pinniped from the San Diego Formation of southern California. University of California Publications in Zoology 47:247-254.
Kellogg, R. 1922. Pinnipeds from Miocene and Pleistocene deposits of California. University of California Publications, Bulletin of the Department of Geological Sciences 13:23–123.
Kohno, N., and Y. Yanagisawa. 1997. The first record of the Pliocene Gilmore fur seal in the Western North Pacific Ocean. Bulletin of the National Science Museum, Tokyo 23:119–130.
Repenning, C. A., and R. H. Tedford. 1977. Otarioid seals of the Neogene. US Geological Survey Professional Paper 992:1–87.
Needless to say, I had a few ideas to follow once I started looking into Bushell's fur seal specimen. Fossils of C. gilmorei had so far only been found in Middle to Late Pliocene deposits, whereas in the late Miocene and earliest Pliocene of Japan, California, and Mexico the earlier fur seal Thalassoleon occurred (which has all double rooted teeth, and lacks cuspate cheek teeth, among other differences). The new specimen only has one cheek tooth - but it has a well developed accessory cusp, like C. gilmorei, and the first two premolars are both single rooted - also like C. gilmorei. In addition, it is relatively small - many other modern otariids are substantially larger. Furthermore, C. gilmorei appears to be the only middle-late Pliocene otariid in the entire Northeastern Pacific fossil record, which made the identification process somewhat easier.
Next up: Other fossil otariids from California and Oregon, and the Pleistocene Callorhinus
specimen.
References:
Berta, A., and T. A. Demere. 1986. Callorhinus gilmorei n. sp., (Carnivora: Otariidae) from the San Diego Formation (Blancan) and its implications for otariid phylogeny. Transactions of the San Diego Society of Natural History 21:111–126.
Boessenecker, R.W. 2011. New records of the fur seal Callorhinus (Carnivora: Otariidae) from the Plio-Pleistocene Rio Dell Formation of Northern California and comments on otariid dental evolution. Journal of Vertebrate Paleontology 31:2:454-467.
Burleson, G. L. 1948. A Pliocene pinniped from the San Diego Formation of southern California. University of California Publications in Zoology 47:247-254.
Kellogg, R. 1922. Pinnipeds from Miocene and Pleistocene deposits of California. University of California Publications, Bulletin of the Department of Geological Sciences 13:23–123.
Kohno, N., and Y. Yanagisawa. 1997. The first record of the Pliocene Gilmore fur seal in the Western North Pacific Ocean. Bulletin of the National Science Museum, Tokyo 23:119–130.
Repenning, C. A., and R. H. Tedford. 1977. Otarioid seals of the Neogene. US Geological Survey Professional Paper 992:1–87.
Bobby, I'm catching up on reading your blog and what can you tell me about Pithanotaria from Santa Barbara? From the Santa Barbara Formation? I didn't know there were any larger vertebrates from the Santa Barbara Formation. I did some work for a mapping project on the Santa Barbara Formation and one of these days hope to publish at least some of that work, but its on the back burner for now. Still I'd like to hear more about what's found there.
ReplyDeleteBest,
Chuck
Hey Chuck! The type and paratype material of Pithanotaria starri is from the Sisquoc Formation in the vicinity of Lompoc. I was just referring to the general area - sorry for the confusion.
ReplyDeleteThe SDNHM has two pinniped bone fragments from the Santa Barbara Formation - I visited the coastal locality in High School, and I suspect that if some careful looking was done, some good vertebrate material could be found - which would be very interesting given the latest Pliocene to Pleistocene age of the unit.