Detour into Choristodera

This week a paper on unusual prehistoric aquatic reptiles was published. I am speaking, of course, of Brownstein (2022) on choristoderes. (If you thought it was going to be spinosaurs, you must be new around here!)

We have a soft spot for choristoderes here at Equatorial Minnesota Towers, even if they rarely rate so much as an "and also featuring Champsosaurus" credit in sci-pop culture. Brownstein (2022) includes the classic Champ but is more focused on the short(er)-faced choristodere Simoedosaurus, although under a fresh coat of taxonomic paint: the North American species S. dakotensis is moved to the new genus Kosmodraco on the grounds of anatomical differences and differences in time and place from the type species, European S. lemoinei. It also gets a new friend, K. magnicornis. Which said, sure... but the species of Kosmodraco still clade more closely to Simoedosaurus than to anything else, so if your genericometer was so tuned, you could still include them in Simoedosaurus with a clear phylogenetic conscience. (There are a lot of anatomical differences between the two genera. My only quibble with Kosmodraco is that it's an awfully pretty name for a choristodere; it sounds more like an an extravagantly crested pterosaur. Meanwhile, Champsosaurus itself is still secretly a taxonomic booby trap waiting to be sprung. Sooner or later someone is going to do something with the various species that is entirely legal by the rules of taxonomy and yet manages to displease everyone else.)

The type skull of Kosmodraco magnicornis. It'll take a moment to orient yourself: the bitey part is surprisingly short (see the "r. lacrimal"? The eye was behind that). What you're looking at is a modest snout attached to a greatly flaring right "cheek" (which comes with a scalloped fringe). Figure 1 in Brownstein (2022). CC BY 4.0.

Either way, the skull is worth a look. There's garden-variety "weird" and then there's "why is this all practically all post-orbital?" (It certainly wasn't for intellect, most of the posterior of choristodere skulls being a series of struts and bars. Note that the maxillary teeth of Kosmodraco only go back as far as the eyes.) Champsosaurus, as we've already seen, had a long, narrow toothy muzzle, with the rest of the skull being broad and low. Kosmodraco had a skull that was still low but much more wedge-shape in dorsal view, as if someone smooshed or amputated a Champsosaurus-like snout. The business end of a Kosmodraco skull is an interesting analog for the modern alligator gar. It's the back of the skull where things get different, as Brownstein (2022) notes. Kosmodraco has a lot more skull going on behind the eyes, which themselves are elevated on skull like those of an alligator. The posterior margin is also ornamented with a series of knobs, and the skull is quite low (Brownstein 2022).

A comparison of the palatal regions of choristoderes Kosmodraco and Champsosaurus to an alligator and an alligator gar also serves as a comparison of basic facial shapes for the four. Similarities between Kosmodraco and the gar aren't as great if you continue through the rest of the skull. Figure 13 in Brownstein (2022). CC BY 4.0.

I'd like to take the opportunity to note that there's another extinct group of aquatic tetrapods with flat, blunt, broad skulls and eyes relatively far forward on the skull, which has so far avoided comparisons to Simoedosaurus/Kosmodraco: the metoposaurid amphibians of the Triassic. Again, it's not a perfect comparison (metoposaur snouts are blunter), but it would seem to point to the long-term existence of a niche for freshwater predators with certain cranial adaptations.

One of the other points noted by Brownstein is there is more diversity in choristoderes than you might suspect by simply looking at the number of genera. For example, as of this writing Champsosaurus is composed of several species over about 20 million years. North American specimens have tended to be lumped with either the short-faced Simoedosaurus or the long-faced Champsosaurus.

References

Brownstein, C. D. 2022. High morphological disparity in a bizarre Paleocene fauna of predatory freshwater reptiles. BMC Ecology and Evolution 22: article number 34. doi:10.1186/s12862-022-01985-z.

Fossil Crocodylomorphs of the National Park Service

For this year's National Park Service fossil group inventory, I've chosen crocodylomorphs, which for convenience I'm going to refer to as "crocs". Crocodylomorpha encompasses the true crocodilians and their closest extinct relatives, which over the years has been defined to exclude major groups of allied Triassic archosaurs (rauisuchids, poposaurs, prestosuchids, etc.). (Technically speaking, traditional Crocodilia is closer to the clade Crocodyliformes, but I have a soft spot for "sphenosuchians" and it's my blog.) Non-crocodilian crocodylomorphs were big players throughout the Mesozoic but came to peter out in the Cenozoic, with holdouts into the Miocene (Sebecosuchia). Some of these non-crocodilian crocodylomorphs looked basically like modern crocodilians and presumably filled very similar niches, but by definition weren't crocodilians*. Others were quite a bit different; for example, small, long-legged terrestrial crocs had a wide distribution from the Late Triassic through the Jurassic, and there were multiple groups of marine forms.

*I have certain misgivings about crown groups, particularly that future stability of usage relies on groups not going extinct (or there would have to be backdating, like radiocarbon dates are pegged to 1950), although at this point I might as well complain about the decline in use of Etruscan.

The NPS record of croc fossils turns out to be sparser than I expected: there are 17 park units with solid records (albeit two of these being reworked or washed up, making them hard to place stratigraphically), and another couple potential records. Here is the requisite map and its accompanying long caption:

Click to embiggen. The sites mentioned in this post are: 1. John Day Fossil Beds National Monument; 2. Bighorn Canyon National Recreation Area; 3. Fossil Butte NM; 4. Dinosaur NM; 5. Colorado NM; 6. Curecanti NRA; 7. Bryce Canyon National Park; 8. Glen Canyon NRA; 9. Petrified Forest NP; 10. Chaco Culture National Historical Park; 11. Theodore Roosevelt NP; 12. Badlands NP; 13. Agate Fossil Beds NM; 14. Niobrara National Scenic River; 15. Big Bend NP; 16. Waco Mammoth NM; 17. Gateway NRA; 18. Fort Washington Park; 19. Cumberland Island National Seashore.

These 19 units are primarily in the Colorado Plateau and northern Great Plain, and these two areas correlate in large part to temporal distribution: the Colorado Plateau records are mostly Jurassic and Cretaceous, and the Great Plains records are Cenozoic. A couple of compact diagrams will show this:

Part 1 shows the Mesozoic, Paleocene, and Eocene records, which make up the bulk of the reports.

Part 2 shows the few younger records; the two that can't be pinned down are added to keep them company.

You can probably guess a lot of the story if you have some familiarity with the stratigraphy of western North America. As so many other groups of terrestrial vertebrates, the place to go in the NPS for Triassic crocs is Petrified Forest National Park, where "sphenosuchians" have been found in the famous Chinle Formation. (Ignore the phytosaurs; they only look like crocs.) After that, possible early croc tracks have been found in the Navajo Sandstone of Glen Canyon National Recreation Area; with all of the Early Jurassic tracks in the Colorado Plateau parks, there are likely other track records. We have no body fossil records in the parks' rocks yet, though (the facies aren't as forgiving as elsewhere). Four parks have records for the Late Jurassic: Bighorn Canyon National Recreation Area has swim traces attributed to crocs in the Sundance Formation, and no points for guessing what's represented at the other three. (It's the Morrison Formation.) NPS Morrison crocs are best known from Dinosaur National Monument, which primarily has the well-represented Amphicotylus (formerly Goniopholis), but also produced the type specimen of the diminutive Hoplosuchus kayi.

The Cretaceous is more sparsely represented, with nothing confirmed from the Early Cretaceous. Ot the Late Cretaceous records, neither Bryce Canyon NP (Straight Cliffs and Wahweap microvertebrate remains) nor Chaco Culture National Historical Park (Menefee isolated material) have much to speak of. Big Bend NP, on the other hand, has the most impressive croc record in the NPS. Granted, that's an easy call when you can point to the type specimen of the suitably Texas-sized Phobosuchus riograndensis (now a species of Deinosuchus), but the park also has by far the longest record of crocs in the NPS. Five formations are represented: the Aguja Formation and Javelina Formation, both Late Cretaceous; the overlying Black Peaks Formation, which straddles the Cretaceous and Paleocene; the Early Eocene Hannold Hill Formation; and the Middle Eocene Canoe Formation. Recently a second croc species has been named from Big Bend NP fossils: Bottosaurus fustidens, from the Paleocene part of the Black Peaks Formation. Other taxa are present, but have not been studied in as much detail.

Looking elsewhere in the Paleocene, there is a single record of a partial bone from the Aquia Formation at Fort Washington Park, and Theodore Roosevelt NP has crocs in the Bullion Creek and Sentinel Butte Formations, comparable to nearby Wannagan Creek (only not quite so concentrated). The Eocene is fairly good for NPS crocs. Apart from Big Bend, we have croc fossils in: the Wasatch Formation at Fossil Butte NM; the Clarno Formation at John Day Fossil Beds NM; and the Chadron Formation at Badlands NP. The type specimen of Caimanoidea visheri (now considered a synonym of Alligator prenasalis) may have come from Badlands NP.

And that's almost the end. Crocs disappeared from the drying interior of North America during the middle Cenozoic. For the Miocene, we have one *very* sketchy potential record from the early Miocene Anderson Ranch Formation of Agate Fossil Beds NM and better records of crocs from the middle Miocene Valentine Formation of Niobrara National Scenic River (including the type specimen of Nordenosaurus magnus, originally described as a big lizard but now identified as a small crocodilian). At Waco Mammoth NM there is late Pleistocene alligator material, but we are otherwise lacking Pleistocene crocodilian records. Two units have material of uncertain provenance: a scute found in dredge material at Cumberland Island National Seashore and various croc fossils that have washed up at Gateway NRA.

What I Did While I Was Out

I made no post last week because I was traveling for work. Generally, I don't have the time to work on a post while traveling, and this was no exception. In order to get the most bang for our buck on work travel, we try to schedule as many projects as possible, and this trip was no exception. I had four separate projects scheduled over eleven days.

Won't somebody please think of the champsosaurs?

Every so often, you may stumble across a crocodile-like thing lurking at the edge of your typical "two-page spread of all kinds of prehistoric critters" found in popular dinosaur books. If the book is of any quality, it includes a legend identifying the denizens, and the crocodile-like thing will be tagged "Champsosaurus" (if there isn't an error in the legend, of course). Unless you're intrigued by the name, or something else tickles your fancy, you'll probably continue along. There are a lot of crocodiles and crocodile-like things to keep track of, after all. What makes this one unusual?