Sunday, August 21, 2016

Champsosaurus: Adventures in 19th century taxonomy

It all started innocently enough with a couple of paragraphs in the 1870 Proceedings of the Academy of Natural Sciences, submitted on behalf of Joseph Leidy. The item notes that Leidy had received a reptilian vertebral centrum (the body of the vertebra, minus all the processes and such) collected by Ferdinand Vandeveer Hayden from the Moreau River, probably from Cretaceous rocks. It reminded Leidy of the vertebrae of marine reptiles, specifically Nothosaurus, so he proposed to name it Nothosaurops, type species N. occiduus. So far, so good. He got around to a longer description in 1873, in which he mostly repeated the 1870 information, but also included some figures of the fossil. Unfortunately, somebody goofed, because the text refers to something called Nothosaurus occiduus, but the plates and captions are using Nothosaurops occiduus. This would not be of much modern interest except for the fact that there is a good chance the specimen is the first described fossil of Champsosaurus.

The holotype of Nothosaurops/Nothosaurus occiduus, from Plate 15 in Leidy (1873), with a couple of other fossils erased.

In the Nothosaurops corner we have Hay (1902), Williston (1903, 1908), and Spamer et al. (1995). In the Nothosaurus corner we have Cope (1875), Erickson (1972), and Gao and Fox (1998). To cut to the chase, if it's Nothosaurus occiduus, it's a lot easier to ignore than if it's Nothosaurops occiduus. In the first case, we've already got the genus Nothosaurus, which is a Triassic aquatic reptile much more closely related to plesiosaurs, so we can nudge "N. occiduus" into the pile of marginal nineteenth-century species. In the second case, we've got a unique genus which needs to be dealt with. Williston (1908) flat-out stated that Nothosaurops should be given precedence, which obviously has not been done. Erickson (1972) and Gao and Fox (1998) used Leidy (1873) as the publication of record. They considered Nothosaurops a typographical error and dismissed "N. occiduus" as either a nomen vanum (an "empty name", a term no longer used much because nomen dubium has taken over its territory; Erickson 1972) or a nomen oblitum (a name which can be considered lapsed because of disuse; Gao and Fox 1998). Although I believe that Champsosaurus would practically be a lock for conservation in the face of Leidy's nearly forgotten saurian if the case were put before the International Commission on Zoological Nomenclature, I disagree with Nothosaurops being a typo. Based on Leidy's 1873 description, I suspect that Leidy originally intended a separate genus (1870), but after a few years passed he changed his mind and designated a species within Nothosaurus (1873), and appearances of Nothosaurops are cases where he forgot to revise the captions. (This is not difficult to do, especially in the days before computerized word processors with replace functions.) Furthermore, I don't think that the 1870 publication can be dismissed, especially when most of the text is reused for the 1873 description. Edward Drinker Cope, who named Champsosaurus, had no particular concern about Leidy's saurian; he classified the specimen as Plesiosaurus occiduus (Cope 1875). This wasn't a case of conflict of interest, either, given he wouldn't name Champsosaurus until 1876.

(Curiously, the Paleobiology Database has separate entries for Nothosaurus occiduus and Nothosaurops occiduus. Nothosaurus occiduus is classified as a champsosaurid using Erickson (1972), and Nothosaurops occiduus is classified as a plesiosaur using Williston (1903), apparently having missed his 1908 statement.)

This is not where the fun ends. In a twist that should surprise no one who is familiar with Edward Drinker Cope's practice of nomenclature and taxonomy, Champsosaurus itself hides a bundle of very marginal species. As noted above, Cope named Champsosaurus in 1876. As was his custom, he gave the world not only one species but several, in this case four species based on a grab-bag of Judith River vertebrae differentiated by proportions. For the record, they are C. annectens (the type species), C. brevicollis, C. profundus, and C. vaccinsulensis. The first three are probably all variants on the same thing (Erickson 1972; Gao and Fox 1998), and C. vaccinsulensis is not a champsosaur (Brown 1905; Erickson 1972). For good measure, he later pulled the same trick with New Mexican fossils, coining C. australis (Cope 1881), C. puercoensis, and C. saponensis (Cope 1882). Also in the 1881 article, he accidentally created the completely undescribed and undefined name "C. laticollis" when he noted that the dimensions of the C. australis cervical vertebrae "are about those of the C. laticollis". The usefulness of this comparison is somewhat lessened by his complete omission of the dimensions of cervical vertebrae belonging to "C. laticollis". The one other hint we have to the identity of "C. laticollis" is buried in an anonymous author's "Scientific Intelligence" note in the November 1881 issue of the American Journal of Science. This short piece summarizing Cope (1881) reported that "C. laticollis" is from the Laramie beds (uppermost Cretaceous), but then also states that Cope based the genus on it, so maybe it's just all wet.

Out of Cope's work, we are left with six formally described species still considered to be Champsosaurus, one species outside of Champsosaurus, and one "species" that could be anything from a faulty memory to a typo for C. brevicollis to a species he wanted to name but never got around to. The six actual Champsosaurus species are all based on various vertebrae, all useful in illustrating the presence of Champsosaurus, but not so helpful for doing taxonomic comparisons. More recent authorities such as Erickson (1972) and Gao and Fox (1998) have made the case that these early species can be regarded as belonging to the genus Champsosaurus, but none of them are diagnostic within Champsosaurus. This includes C. annectens, the type species. To get to a species that can be diagnosed, you have to wait until Brown (1905), with C. ambulator and C. laramiensis.

References:

Brown, B. 1905. The osteology of Champsosaurus Cope. Memoirs of the American Museum of Natural History 9(1).

Cope, E. D. 1875. The Vertebrata of the Cretaceous formations of the West. Report of the United States Geological Survey of the territories. F. V. Hayden, United States Geologist in charge, vol. 2.

Cope, E. D. 1876. On some extinct reptiles and Batrachia from the Judith River and Fox Hills beds of Montana. Proceedings of the Academy of Natural Sciences of Philadelphia 28:340-359.

Cope, E. D. 1881. A Laramie saurian in the Eocene. The American Naturalist 15:669–670.

Cope, E. D. 1882. Contributions to the history of the vertebrata of the lower Eocene of Wyoming and New Mexico made during 1881. Proceedings of the American Philosophical Society 20:139–197.

Erickson, B. R. 1972. The lepidosaurian reptile Champsosaurus in North America. Science Museum of Minnesota Monograph 1: Paleontology.

Gao, K.-Q., and R. C. Fox. 1998. New choristoderes (Reptilia: Diapsida) from the Upper Cretaceous and Palaeocene, Alberta and Saskatchewan, Canada, and phylogenetic relationships of Choristodera. Zoological Journal of the Linnean Society 124:303–353.

Hay, O. P. 1902. Bibliography and catalogue of the vertebrata of North America. U.S. Geological Survey, Washington, D.C. Bulletin 179.

Leidy, J. 1870. [On Nothosaurops occiduus.] Proceedings of the Academy of Natural Sciences of Philadelphia 22:74.

Leidy, J. 1873. Contributions to the extinct vertebrate fauna of the Western Territories. Report of the U. S. Geological Survey of the Territories, F. V. Hayden; U. S. geologist in charge, 1:14-358.

Spamer, E. E., E. Daeschler, and L. G. Vostreys-Shapiro. 1995. A study of fossil vertebrate types in the Academy of Natural Sciences of Philadelphia: taxonomic, systematic, and historical perspectives. The Academy of Natural Sciences of Philadelphia Special Publication 16.

Williston, S. W. 1903. North American plesiosaurs, part 1. Geological Series Field Museum of Natural History 2(1).

Williston, S. W. 1908. North American plesiosaurs: Trinacromerum. The Journal of Geology 16(8):715–736.

2 comments:

  1. Nothosaurops cannot be a nomen oblitum, since it was used as valid after 1899 (ICZN Article 23.9.1.1). Neither Erickson's nor Gao and Fox's opinions are valid since they were apparently unaware of Leidy's original 1870 description of Nothosaurops. Indeed, Gao and Fox missed the obvious statement by Leidy (1873) that he intended to same Nothosaurus occiduus as a new species of the genus, not to name a new genus Nothosaurus- "In form and construction it resembles the vertebral centra of Nothosaurus, an extinct reptile of the Triassic formation of Europe, and probably it belongs to an animal of the same order if not the same genus." I agree with your conclusion that Leidy originally thought occiduus was a distinct genus, then decided it was congeneric with Nothosaurus, but forgot to change the plate caption. Thus I would say Nothosaurops occiduus is a valid name, with Nothosaurus occiduus being an objective junior synonym.

    I don't think we have to worry about Nothosaurops being a senior synonym of Champsosaurus though, because it's near certainly indeterminate within some subclade of Choristodera. Similarly, given our vastly improved knowledge since 1905, I doubt Brown's opinion that the C. annectens' type axial centrum is generically diagnostic. Champsosaurus is in dire need of a detailed restudy of its original types and probably a subsequent ICZN petition establishing C. laramiensis or ambulator as the neotype.

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    1. I agree that Nothosaurops is probably not diagnostic enough to pin onto Champsosaurus in any kind of formal way, although I do think it's kind of a Thespesius/Edmontosaurus situation.

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