Tiny frogs of the Chinle

I am embarrassed to realize that after five full years of doing this, with a header mentioning "National Park Service paleontology" and "the Mesozoic", I had not done anything with the Chinle Formation or Petrified Forest National Park. This week offers a fine opportunity to correct this oversight, with the publication of the first frog fossils from the Chinle Formation.

The Chinle Formation is a terrestrial unit, with its colorful rocks deposited in various floodplain, river, lake, and other settings during the Late Triassic. The mosaic of settings led to a variety of rock types, from conglomerates to mudstones, with a healthy supply of volcanic ash from eruption centers to the west. Because we're talking about the Late Triassic, before the end-Triassic extinctions gave dinosaurs the opportunity to fill most of the empty terrestrial niches, the Chinle is full of fossils from all sorts of unusual and obscure animals. (And also petrified wood and freshwater mussels. Lots and lots of petrified wood and mussels!) It and its correlatives are usually good for at least one surprise every few years. Lurking among all of the phytosaurs and aetosaurs and so forth were representatives of lineages that would blossom later. Some of them we know from bones (dinosaurs, early croc relatives, pterosaurs, etc.), others can be suspected based on time and place but haven't yet been found. With this week's announcement, frogs move from the potential to the confirmed (Stocker et al. 2019).

The Chinle Formation is divisible into a number of members, depending on where you are. In ascending order, the units in the Petrified Forest NP area are the Mesa Redondo, Blue Mesa, Sonsela, Petrified Forest, and Owl Rock members. For the frogs, we're dealing with the Blue Mesa and Sonsela members, representing approximately 223 to 213 million years ago. (Following the stratigraphy of the Chinle Formation requires a certain amount of effort and dedication. Not only are different members found in different places, but usage has varied over time.) The three frog-producing localities are spread from the park to the famous Placerias Quarry near St. Johns, Arizona (Stocker et al. 2019).

An outcrop of the Sonsela Member, from the park website (NPS/Andrew V. Kearns).

Frogs, being small and delicate, are not exactly heavily represented in the fossil record. The frog lineage, Salientia, is known back to the Early Triassic, but if you were doing a Compact Thescelosaurus-like project on Mesozoic salientians you'd be finished pretty quickly. Salientia includes early stem-frogs plus the crown group Anura, which is the group consisting of all living frogs, their most recent common ancestor, and everything else that falls within that group. There are to date two Early Triassic stem-frogs, the well-known Triadobatrachus of Madagascar and the somewhat less famous Czatkobatrachus of Poland, with the next named salientian being a respectable frog's jump all the way into the Early Jurassic of Arizona, Prosalirus of the Kayenta Formation. The Kayenta has historically proven stubborn about this whole "absolute dating" thing, but an age in the first half of the Early Jurassic is the consensus, so that's a gap of something approaching 60 million years (approximately 250 to 190 million years). The unnamed Chinle salientian practically splits the difference (Stocker et al. 2019).

The Chinle salientian is known from five tiny fossils. Fortunately, four of them are ilia, which are very distinctive bones in frogs: there is a cup-like socket for the femur and a long thin blade directed anteriorly, lengthened by cartilage, part of the mechanism that gives frogs their spring. In the case of the Chinle form, they are also well into the microvert range: approximately 6.2 mm long, or about a quarter of an inch, the kind of stuff you find when you run sediment through screens as opposed to spotting while hiking around. The whole animal would have been less than about 25 mm long, or about an inch, in the realm of modern miniature frogs (Stocker et al. 2019). So, if you were thinking that ur-frogs might have been prehistoric giants, well... not so much in the Chinle. There *were* enormous amphibians in the Chinle, specifically the metoposaur Koskinonodon and its equally enormous taxonomic history, but it isn't closely related to anything living and it didn't look much like a frog.

The form of the hip joint and shaft suggest that the Chinle form was well on the way to modern frog jumping mechanics (Stocker et al. 2019). It may have been a true anuran, but only time and more fossils will tell. The presence of a stem-frog or true anuran in the Chinle also has some implications for frog distribution and paleoecology. The two Early Triassic stem-frogs lived outside of the tropics, whereas the Chinle form was more or less at the equator in western Pangea, showing that the frog lineage had spread across the supercontinent within the Triassic. Also of interest is the persistence of salientians from the Chinle into the Kayenta; over the length of the Chinle, the local climate became more and more arid (Stocker et al. 2019).

A Triassic frog clings to the snout of a phytosaur, used with permission by Andrey Atuchin (supplied by Adam Marsh/PEFO).

References

Stocker, M. R., S. J. Nesbitt, B. T. Kligman, D. J. Paluh, A. D. Marsh, D. C. Blackburn, and W. G. Parker. 2019. The earliest equatorial record of frogs from the Late Triassic of Arizona. Biology Letters 15:20180922. doi:10.1098/rsbl.2018.0922.

The many moods of Rauffella

Earlier in February, I attended a Geological Society of Minnesota fossil lab hosted by Jeff Thole and Macalester College. At some point someone brought out a fossil that they weren't familiar with; it looked a lot like the final photo in this post, a light-colored object that resembled loops of cord. What this person had was one of the most characteristic but least scientifically appreciated fossils of the Decorah Shale: the trace fossil Rauffella (specifically R. palmipes, as we'll get to later).

Your Friends The Titanosaurs, part 9: "Campylodon", Clasmodosaurus, Choconsaurus, and Daxiatitan

I've come up against some judgement calls at this point. There are two varieties for this exercise. The first includes a group of names based on poor material that have had little study but which are sometimes/often stuck into Titanosauria, usually from historical inertia or the "provenance argument". The provenance argument goes something like this: "the only described sauropods from this time and place are titanosaurs, ergo this is a titanosaur". The provenance argument is a seductive bit of taxonomic stereotyping, but I'm wary of it. For one thing, the implicit flip side is "therefore there cannot be anything else here than titanosaurs". When the provenance argument is correct, it's not because you put in any effort or had any kind of insight, and when it isn't, you look stupid for being lazy. When confronted with this dilemma and, say, a handful of teeth which have no particular characteristics that would put them in one group or another, I don't see what's wrong with "indeterminate sauropod". The other type of judgement call includes the frustrating taxa that sometimes end up in a clade and sometimes end up outside of it, generally because they are in that fuzzy transitional zone.

For this particular entry, I had cause to consider examples of both types: in the historical orphan bin are "Campylodon" ameghinoi (=Campylodoniscus) and Clasmodosaurus spatula, and in the fuzzy clade bin is Chubutisaurus insignis. I decided that the position of Chubutisaurus is too unstable to be confident it's a titanosaur at this point, and so left it out. We can always cover it later on, with some of its similarly ambivalent cronies. As of this writing, others in a similar equivocal position which I currently have outside of Titanosauria include Angolatitan adamastor, the two Huanghetitan species, Ligabuesaurus leanzai, Mongolosaurus haplodon, Phuwiangosaurus sirindhornae, Tastavinsaurus sanzi, Triunfosaurus leonardii, and Wintonotitan wattsi. Two things they almost all have in common: late Early Cretaceous age, and when they do show up in Titanosauria they're just barely over the line. The other two were more problematic because there's not much material and they've never attracted much interest. I decided to cover them here together as examples.

Bajadasaurus pronuspinax

Like their cousins the rebbachisaurids, the dicraeosaurids have been on a roll over the past year. I featured the new oldest dicraeosaurid, Lingwulong shenqi, last year. This week saw the publication of another new dicraeosaurid, from a much more typical time and place (the Early Cretaceous of South America), but with a unique style all its own: Bajadasaurus pronuspinax.

Concerning an Isotelus and its acquaintances

Here we have a big ol' rock.

New Pages: Geologic Time Scale and Classification Diagrams

So far the year has been quiet (well, up until Friday), so I thought I'd address a couple of nagging things, which I've done by adding two pages. The first page is a geologic time scale figure, from one I worked up for National Park Service reports. It's the kind of thing I thought would be handy to have on hand here for reference, instead of having a link to an external site. It'll be updated from time to time as dates are refined. The Quaternary is rather cramped, although I don't feel particularly apologetic for shortchanging Homo sapiens.

The second page is a sort of "map" to the classifications used in the various sheets over at The Compact Thescelosaurus, made with classic ASCII cladograms. (I contemplated drafting them in other ways, but none of them were as amenable to updating.) I thought this would be useful for visualizing the mess of classification columns. The process also forced me to look at the positions of a few clades, as you may have noticed from the updates sheet.

It's not connected to either topic, but I've also added a paragraph of new information to the post on the "Kweichow sauropod" after coming across a mention of it in Averianov and Sues (2017).

Finally, so as not to leave the post without an image, here's one attached to a quick story:

Back in spring 2001 I was on a field trip to the Badlands/Black Hills area of southwestern South Dakota. We were stopped along a road near Deadwood for lunch. One of the professors said something to the effect that "there are fossils in this formation." I looked down at the chunks of rock at my feet and said "You mean like this?"

Yeah, like this.

From my notes the source is the Whitewood Formation (or Dolomite, or Limestone), fittingly enough an (surprise, surprise) Upper Ordovician unit. More on its cephalopods can be found in Miller and Furnish (1937).

References

Averianov, A., and H.-D. Sues. 2017. Review of Cretaceous sauropod dinosaurs from central Asia. Cretaceous Research 69:184–197. doi:10.1016/j.cretres.2016.09.006.

Miller, A. K., and W. M. Furnish. 1937. Ordovician cephalopods from the Black Hills, South Dakota. Journal of Paleontology 11(7):535–551.

Your Friends The Titanosaurs, part 8: Bonitasaura, Borealosaurus, and Brasilotitan

And so we reach Part 8, otherwise to be known as "Bonitasaura and these other two." Bonitasaura is one of the best-known sauropods, even if it isn't quite a household word, and also has some unusual jaw features going for it. Brasilotitan has some of the same jaw features, but is not known from nearly as much material and came along several years later. Borealosaurus has an uncertain grip on Titanosauria, and gripping things takes some creativity when you have either no fingers or just nubs.