Ghost of Christmas Presents

My first love in science, the first thing that really interested me intellectually, was astronomy. It was early 1986, around the time I turned five. (I was a weird kid.) Anyway, I was really into Star Wars, and wanted to go to space and explore different planets. I had (and still have) a book, "Astronomy Today", with lots of great illustrations and diagrams; sure, I wasn't *reading it* reading it yet, but I got a lot out of those illustrations. Then I discovered that, contrary to the movies, we couldn't exceed the speed of light, so we were basically stuck here. It's a nice enough solar system, but, well, y'know... the realization was terribly disappointing, and space went from an obsession to a pleasant side interest (I still love going to the planetary science sessions when attending GSA annual meetings).

At the time, we had just moved to Tucson. There's a lot of difference geologically between southern Arizona and south-central Minnesota; for starters, a lot more of the geology is right there at the surface in Arizona, and a lot more of it is Mesozoic. Still, Arizona is not Dinosaur Central; it's more like a nice suburb. After nearly 40 years I can't say what it was about dinosaurs that caught my eye. Maybe it was the British Museum toys at the Arizona-Sonora Desert Museum. Whatever it was, the feedback loop started: kid shows interest, people around start feeding the interest, interest intensifies...

The step that sealed the interest in paleontology came at Christmas 1986. There was one rectangular gift with unusual heft. Under the wrapping paper was the Normanpedia, "The Illustrated Encyclopedia of Dinosaurs". Could I read it? Well, okay, sure, I was a great reader for five-and-a-half, but it's safe to assume it took some time and practice before I was conversant (and the book probably accelerated the process, so on top of his scientific accolades, David Norman can take some credit for improving at least one kid's  literacy). But don't forget the second word of the title! The restorations, the photos, the skeletal diagrams! I went through reams of paper tracing skeletons, then drawing outlines around them on the other side of the sheets. That way, there would be a skeleton on one side, a drawing on the other, and if I held the paper to the light, both of them would be there, skeleton revealed inside the body. I loved that book to death; it's in a box of mementos somewhere at home, dust jacket long gone, spine held together by heavy-duty tape, a slip of paper taped over the "dinosauroid" drawing because it creeped me out. I had to get a second copy in the Nineties because the first was falling apart. The second copy also started falling apart after about ten years, so now I'm on my third. Anyway, no particular moral or commentary or insight this time, just wanted to share some holiday nostalgia. See you in 2026!

What do Delaware, Hawaii, and Rhode Island have in common?

Answer: They are the only three U.S. states for which I can't find a record of a mastodon or mammoth.

Hawaii is not exactly a surprise. Certainly proboscideans can swim, but if you want a large mammal that can swim halfway across the Pacific Ocean without dying, it's going to look and function suspiciously like a cetacean instead.

Delaware and Rhode Island, on the other hand, doubtless had mammoths and mastodons tromping over them at some point, because they're surrounded by states (and submerged coastal plains) that have their fossils. They're just both small states not blessed with the most helpful geology, and mantled by urbanization. Rhode Island gets an additional strike from being subjected to glaciers. Someday the fossils will turn up, though. Maybe there's already something; maybe somebody has a funny-looking doorstop they found in 1973 and never paid much attention to, or there's a notice in a local paper published in 1989 about a couple of campers who found a tooth and turned it over to a local historical society, or somebody will hop into the comments of this post and say, "Hey, ya numbskull, ya missed this!"

Now, the nitty-gritty: every once in a while I'd wondered how many states have records of mammoths or mastodons, and occasionally puttered in a source or two. This time I decided to be more thorough. I checked the Hay inventories, Neotoma (which has the old FaunMap contents), and the Paleobiology Database for starters. These three together narrowed down the list of absences to Delaware, Hawaii, New Hampshire, and Rhode Island. If you're curious: Hay didn't have mammoths or mastodons in those four plus Maine and basically omitted Alaska, but had all the rest plus the District of Columbia; Neotoma was missing the four plus Alabama, Mississippi, North Dakota, Vermont, and D.C.; and the PBDB was missing the four plus Connecticut, Louisiana, Massachusetts, Maine, New Jersey, North Dakota, Vermont, and D.C. This is why it's important to check multiple sources; no one source has everything.

Putting aside Hawaii, that left Delaware, New Hampshire, and Rhode Island to check out. Now was the time to drill into the resources of various search engines and museum databases/iDigBio. For the record, I leaned on Google Scholar, Google Books, and the news tab of regular Google (not Google News), with various permutations of "mammoth", "mastodon", "Mammuthus", and "Mammut" plus a state, with additional terms as necessary to knock out particularly obnoxious false positives. This eliminated New Hampshire, which now has a record from a find on land after a couple of teeth found nearby offshore. For a short time I'd thought I'd found one for Delaware, thanks to a short piece in the Winter 1994 issue of "First State Geology" (p. 3). The article mentioned that Jeremy Cloutier of Milford had donated a mastodon tooth to the Delaware Geological Survey, but the tooth derived from offshore clam dredging and therefore doesn't count for our purposes. So, for now, we're still at Delaware, Hawaii, and Rhode Island. (Still a few field days 'til Christmas, if you're feeling like pulling off a Christmas miracle!)

"Archie", the world's largest mounted Columbian mammoth skeleton, on display in Elephant Hall, University of Nebraska State Museum.

A Visual Paleontological Inventory of Utah’s National Park Service Areas

Allow me to indulge in a bit of puffery and congratulations...

Let's go back a couple of years ago. We in the NPS Paleontology Program knew that the new edition of the Utah Geological Association's "Geology of Utah's Parks and Monuments" was not going to have an update of the overall park paleontology chapter. (Which is fair enough, since it was 35 pages in the previous update and was not getting shorter. The new volume is still well worth seeing, though, especially if you haven't seen the earlier editions.) We also were working with Tut Tran, then putting the finishing touches on the paleontological inventory report for Bryce Canyon National Park. These touches included some clever biostratigraphic figures of a kind we'd never used before. Thus was an idea born: Tut would prepare a standalone article featuring similar figures for the rest of Utah, with contributions from various luminaries in NPS and Utah paleontology. This article, "A Visual Paleontological Inventory of Utah’s National Park Service Areas", is now available for your edification and reading pleasure. Congrats, everyone!

Utah has 13 National Park Service units: five parks (Arches, Bryce Canyon, Canyonlands, Capitol Reef, and Zion), six monuments (Cedar Breaks, Dinosaur, Hovenweep, Natural Bridges, Rainbow Bridge, and Timpanogos Cave), one national recreation area (Glen Canyon), and one national historical park (Golden Spike). (There are a few other units in the state designated as "National", such as Grand Staircase-Escalante National Monument and Flaming Gorge National Recreation Area, but they are under different agencies.) This is omitting a few NPS trails (California, Mormon Pioneer, Old Spanish, and Pony Express) and the Virgin Wild and Scenic River, which are complicated to deal with because they don't have formal boundaries the same way the other types of units do; one day I'd like to go over rivers and trails thoroughly, but that's for another day. Anyway, the 13 units give Utah one of the best state-wide paleontological records in the NPS, up there with Alaska and California. Each one of them has something, although certainly some have more than others. The five national parks, Dinosaur NM, and Glen Canyon NRA are the most abundantly supplied and feature the longest records.

Figure 2, comparing the stratigraphic records of Utah's NPS units. Click to embiggen. CC BY 3.0 US.

These seven units are fairly similar stratigraphically and in fossil content, with the exceptions of Bryce Canyon (getting started when most of the others are tapering off) and to a certain extent Dinosaur (longer record and some different formations due to distance from the other units). The classic assortment of rocks generally includes the Permian, Triassic (Moenkopi and Chinle), Jurassic (Glen Canyon Group, San Rafael Group, and Morrison Formation), and part of the Cretaceous (Cedar Mountain Formation and Naturita, the old Dakota Sandstone). Arches, Canyonlands, Capitol Reef, and Glen Canyon in particular are geological and paleontological siblings. There are some older and some younger formations, but generally the sweet spot is between about 300 and 100 million years ago.

You might think, based on Dinosaur's reputation, that the Morrison Formation is the big producer, but as it turns out that's not the case. The Lower and Middle Jurassic rocks that make up so much of the scenic vistas, although not big bone producers, are perhaps the most iconic fossil producers with their fossil tracks. (Which is fitting for a park, because just like you can't put a vista in a box and take it somewhere else, fossil tracks are best appreciated where they are found.) The Chinle and Moenkopi are also good but a bit overlooked (Utah's Chinle having the excuse of being in the shadow of Arizona and New Mexico). Another interval that is very productive is the late Quaternary: several units have cave/rock shelter assemblages, usually thanks to packrats, with the assistance of other animals. Glen Canyon in particular is noted for dung caves, including Bechan Cave ("big feces" in Navajo) and its supply of mammoth dung. The Cretaceous is coming into its own with work at Bryce Canyon, Dinosaur, and Glen Canyon; in fact, work on the Cretaceous made our article outdated almost the day it was published, thanks to Pahasapasaurus gillettei from the Tropic Shale of Glen Canyon (Schmeisser McKean 2025). We were able to get October's Athenar bermani in there in the proof stage, but the plesiosaur managed to wait just long enough. (Darn it.)

This article is not a rewrite of the NPS paleontology chapters in the old UGA volumes. It sets out to do different things and is much more visual in focus, rather than textual. (It even has its own nifty cover image, done by artist Benji Paysnoe in the spirit of the great "Vertebrate Paleontology of Utah" volume.) The main things you will find in this that you won't find in the older chapters are Tut's outstanding diagrams and a grand 62-page appendix listing fossil taxa from each park unit by formation, with citations. Between the nine authors, I think we were able to put together a pretty darn comprehensive appendix, although doubtless we missed something. (If you spot something, let us know, so it can be included if we get the opportunity to do an update.) Me, of course, I'm hoping for more Paleozoic invertebrates, but I'm not turning up my nose at anything!

Figure 39, a sample diagram (Rainbow Bridge NM). You should go to the article to see what's been found at the big parks! CC BY 3.0 US.

We're all excited for this to be out, and we hope you find it useful as well, as a unique look at the paleontological riches of Utah's NPS units.

References

Schmeisser McKean, R. L. 2025. A new species of Pahasapasaurus (Plesiosauria: Polycotylidae) from the Upper Cretaceous Tropic Shale (lower Turonian) of southern Utah, U.S.A. Cretaceous Research 106269. doi: https://doi.org/10.1016/j.cretres.2025.106269.

Tran, T., A. R. C. Milner, J. S. Tweet, D. D. DeBlieux, R. Hunt-Foster, A. B. Shaffer, J. I. Kirkland, E. Warner-Cowgill, and V. L. Santucci. 2025. A visual paleontological inventory of Utah’s National Park Service areas. Geology of the Intermountain West 12: 221–292. doi: https://doi.org/10.31711/giw.v12.pp221-292.

Speleotherium logani

It would be easy to think that we've got the large Late Pleistocene mammals of North America pretty well locked down. After all, every self-respecting animal of that stripe has a long list of synonyms and pseudonyms, sometimes going well back into the 19th century. (All right, to be fair, horses are a mess if you look closely.) Even here, though, there can be surprises. Recently we had the Pacific mastodon and the re-establishment of the dire wolf in its own genus, and now we have the recognition of an entirely new genus and species of scrubox, Speleotherium logani (White et al. 2025).

If you know a bit of Greek, you'll get the idea that the name indicates we're dealing with a beast ("therium") associated in some way with caves ("speleo"). The holotype and best specimen of Speleotherium logani did indeed come from a cave, and in fact was partially encrusted with cave deposits (which certainly give it an unusual look but haven't made it easy to prepare or interpret). A few years ago the skull was photographed for a photogrammetric model, and you can take it for a spin and see the deposits, particularly on the right side of the face. This specimen and others were discovered in 1976 in what is now called Muskox Cave in its honor, within Carlsbad Caverns National Park, New Mexico. The species name honors the discover, Lloyd Logan. Although the name literally translates to "Logan's cave beast", the authors suggest the common name "Logan's austral scrubox" instead (White et al. 2025).

Figure 31 from Kottkamp et al. (2020), used in several other places as well (including White et al. 2025). You're looking at the back of the skull, going off to the right. Other bones are also visible, as well as the coating of cave mineralization.

The Muskox Cave fossils were long thought to belong to the shrubox Euceratherium (which is the name used for it at the 3D model above). That's how we labeled them in recent inventories of the park's fossils, for example (Kottkamp et al. 2020, 2022). It was not until White et al. began to prepare the specimens that it became apparent that there was something else here. Euceratherium has a narrow "forehead" and complexly twisted horn cores, whereas the Muskox Cave skull has a broad "forehead" and less curvy cores. The metapodials (metacarpals and metatarsals) are also shorter and stockier, which twigged something else; the authors knew of similarly proportioned metapodials of an animal similar to but much smaller than Euceratherium from sites in Mexico and Belize, but had never had a face to go with them. Speleotherium is therefore not a "one-off" but something that had a fairly wide range in North and Central America, and there are probably other examples that are currently listed as Euceratherium (White et al. 2025). The metapodial anatomy is rather similar to the same bones of the takin (White et al. 2025), which is a Himalayan bovid that's kind of in-between things like muskox, sheep, and goats in appearance and habits. That may give an idea of what Speleotherium was doing. The short and stocky metapodials suggest it was active in rugged terrain, but based on fossil distribution it was not limited to higher elevations (White et al. 2025).

The skull in the process of being captured by photogrammetry (producing the 3D model linked above), showing its best side. Figure 36 in Kottkamp et al. (2020).

Given we're dealing with National Park Service fossils, you can imagine it's been on my radar. It showed up in the 2001 inventory of NPS cave fossils as what is now a grainy photo (Santucci et al. 2001), but we've since been able to feature the image in higher resolution glory for Carlsbad-specific inventories (Kottkamp et al. 2020, 2022). There's been a soft spot for it around here; it even got a coloring page, now part of the "Cenozoic Life in the National Parks" coloring book (just mentally substitute Speleotherium for Euceratherium as the label; the coloring page was prepared long before the bones got their new name).

References

Kottkamp, S., V. L. Santucci, J. S. Tweet, R. D. Horrocks, E. Lynch, and G. S. Morgan. 2020. Carlsbad Caverns National Park: Paleontological resource inventory (public version). Natural Resource Report NPS/CAVE/NRR—2020/2148. National Park Service, Fort Collins, Colorado.

Kottkamp, S., V. L. Santucci, J. S. Tweet, R. D. Horrocks, and G. S. Morgan. 2022. Pleistocene vertebrates from Carlsbad Caverns National Park, New Mexico. New Mexico Museum of Natural History and Science Bulletin 88:267–290.

Santucci, V. L., J. Kenworthy, and R. Kerbo. 2001. An inventory of paleontological resources associated with National Park Service caves. NPS Geologic Resources Division, Denver. Technical Report NPS/NRGRD/GRDTR-01/02.

White, R. S., J. I. Mead, and G. S. Morgan. 2025. Logan's austral scrubox, a new ovibovine (Mammalia: Artiodactyla: Bovidae) from Muskox Cave, Eddy County, New Mexico. New Mexico Museum of Natural History and Science Bulletin 101: 473–494.

Hyoliths VII: The New Blood

Have you ever been working on some mundane task when you suddenly wondered about the latest news from the world of hyoliths? Taking a walk, or merging onto a busy highway, or applying shampoo in the shower? All right, probably not, but if so, we're here for you!

Appropriately for this time of year, we have some news of hyoliths meeting or escaping grisly demises. (Or not, but that's taphonomy for you.) Kraft et al. (2023) published on an exceptionally well-preserved specimen of the Middle Ordovician central European trilobite Bohemolichas incola, including gut contents. The hyoliths are only a small part (quite literally!) of the story, which is well worth checking out if you have any interest in trilobites. The small trilobite (on the order of 35 mm or 1.4 inches long) apparently ate every darn thing it could fit in its mouth that wasn't putting up too much of a fight, including tiny hyoliths, ostracodes, stylophoran echinoderms, and chunks of shells.

The trilobite in question (Figure 1 from Kraft et al. 2023; scale bar 10 mm or 0.4 inches). Hyolith bits are in purple, including one recognizable shell under the trilobite's pygidium (tail segment). CC BY 4.0.

Paleozoic examples of the bilobed trace fossil Rusophycus are often attributed to resting trilobites, and one of the things you can do when you're not moving is pick up a snack. Lee et al. (2025), in a description of Cambrian Rusophycus from China, included an example where the trace was associated with hyolith shells. Unlike classic Rusophycus, thought to occur at the seafloor surface, this example was interpreted as a burrow. Also unlike classic trilobite predation trails, in this case the food had a hard shell. The trace-maker is thought to have been scavenging for hyoliths that had been transported from elsewhere.

Returning to the Ordovician of central Europe, we find a hyolith that was not eaten, although not from lack of trying. Fatka et al. (2023) reported a specimen of Elegantilites custos with healed damage in the form of scratches on its operculum. The culprit in this case is thought to have been an echinoderm, possibly an ophiuroid (brittle star) trying to get in.

Perhaps you'd prefer to think of your hyoliths more in terms of a grand and proud lineage, rather than delicious treats for every passing trilobite and brittle star. If so, Liu et al. (2024) have an analysis of Cambrian hyoliths for you. Using all valid Cambrian genera (N=115), they considered a set of 20 morphological characteristics over time and space. Overall hyolith taxonomic diversity peaked in Series 2 of the Cambrian (roughly speaking, the time when trilobites appeared and therefore kind of like the old "Early Cambrian"). They then keeled over sharply and were at lower levels for much of the rest of the Cambrian, locally reviving to a certain extent in the Early Ordovician. Their decline may have been due to an ocean anoxic event (the Sinsk Event) around 513 to 508 million years ago. The two major wings of hyoliths, the hyolithids (the kind with helens and complex opercula; filter feeders?) and orthothecids (the kind without helens and with simple opercula; deposit feeders?), did not follow the same curves: the orthothecids peaked sooner and felt the bite sooner, whereas the hyolithids didn't really get started until Series 2 and actually peaked just after it before suffering their drop. Morphological diversity was greatest in Series 2, although granted hyoliths had a somewhat limited repertoire.

The curve of Cambrian hyoliths. Figure 1 in Liu et al. (2024). CC BY 4.0.

References

Fatka, O., M. Valent, and P. Budil. 2023. The first healed injury in a hyolith operculum. The Science of Nature 110(50). https://doi.org/10.1007/s00114-023-01879-0.

Kraft, P., V. Vaškaninová, M. Mergl, P. Budil, O. Fatka, and P. E. Ahlberg. 2023. Uniquely preserved gut contents illuminate trilobite palaeophysiology. Nature 622: 545–551. https://doi.org/10.1038/s41586-023-06567-7.

Lee, D.-C., M.‑K. Oh, Y. Zhang, X.‑L. Zhang, J.‑H. Lee, K. Liang, and W. Li. 2025. Two new probable feeding traces of Rusophycus from the Cambrian of China: tracemaker’s behavior and formation mode. Geosciences Journal 29: 1–17. https://doi.org/10.1007/s12303-025-00007-6.

Liu, F., T. P. Topper, L. C. Strotz, Y. Liang, Y. Hu, C. B. Skovsted, and Z. Zhang. 2024. Morphological disparity and evolutionary patterns of Cambrian hyoliths. Papers in Palaeontology 10(2). https://doi.org/10.1002/spp2.1554.

Your Friends The Titanosaurs: Utetitan zellaguymondeweyae

Back almost four years ago when we first waded into Alamosaurus, I noted that "I was waiting on it because I was concerned that it might suffer a taxonomic detonation at any time, so I thought I'd hold off as long as possible." Titanosaurs work on their own schedule. Although people have danced around the issue for decades by suggesting that A. sanjuanensis is dubious or refraining from including some specimens in the species, nobody had taken the step of formally proposing another name for any of the material until Gregory S. Paul in October 2025 with Utetitan zellaguymondeweyae. This genus and species are based on the North Horn specimen that has long been the "practical purposes" type* of A. sanjuanensis.

*Seems like there ought to be a name for non-type specimens that everyone uses instead of the type. "Apatotype", for "deceptive type"? Or "pseudotype" for "false type"? Think like the "Mantel-piece" of Iguanodon mantelli. Brachiosaurus had a whole apatotype species until it was split off for Giraffatitan.

Compact Thescelosaurus Year Ten

Twelve months have rolled along since our previous check on The Compact Thescelosaurus, now clocking in at a decade of existence. With Triassic and Jurassic pseudosuchians for the previous two years, the obvious addition was Cretaceous pseudosuchians, of which there were more than the previous two additions combined. The Cretaceous section wasn't quite so dominated by one group like the Jurassic was by thalattosuchians, but it was a great time to be a notosuchian.

Like this one, Simosuchus clarki. I took this photo when the traveling exhibit visited the Science Museum 11 years ago; Wikipedia is running a photo of the same mount from another stop on the itinerary.

More odds and ends

At the moment I'm preparing the grand annual update to The Compact Thescelosaurus. If you've been following along over the past couple of years, it doesn't take being Sherlock Holmes to guess what it will be. Actually, it doesn't even take being 1940s Radio Program Watson to guess, but do act surprised. In completely unrelated news, there's a taxon published this week I recommend you see: the little croc Thikarisuchus xenodentes. (What, you were expecting the theropod?) T. xenodentes, from the Cenomanian-age Blackleaf Formation of Montana, had a sharply triangular skull in top view and strongly differentiated teeth, among them the expected bitey teeth at the very front and long, low, narrow teeth at the back. This small croc may have had a taste for plants, or perhaps sliced up insects.

Something we love around here at Equatorial Minnesota is historical content about Minnesota geology. You may be familiar with the Minnesota Geological Survey's publication archive. It turns out that there are a bunch of MGS field notebooks scanned and available via the University of Minnesota libraries. You can find notebooks there from paleontologists such as Frederick Sardeson, Robert Sloan, and Clinton R. Stauffer, all of whom curiously have last names starting with "S". (Also interesting: Sardeson's field notebooks all have the left-hand pages filled out, but right-hand pages are much less frequently used. Was Sardeson a lefty?) There are also materials from the Department of Earth and Environmental Science.

Do you prefer to see your geology in the field? I came across a nice section of the St. Peter through Platteville under the I-94 bridge, on the east bank of the Mississippi. I'd been there years before but don't remember being so impressed with it. Maybe it wasn't as well-exposed then, or maybe I just wasn't experienced enough to appreciate it. The Platteville interval at the top was definitely exposed, but maybe this lower exposure is the result of more recent erosion or something. It definitely bears further photography and investigation, as it has a great view of the Glenwood.

The important part looks like this.

And here's what it looks like if I annotate all over it. I'm not entirely confident with the thickness of the Hidden Falls Member; for some reason the Mifflin is really grungy here. The Glenwood bits are open to interpretation (the Nokomis gets "Glenwood/St. Peter" because it's technically in the Glenwood but can't be distinguished from the St. Peter in well logs or gamma logs). The Tonti Member makes up most of the St. Peter Sandstone, if you're curious. There may be some Carimona at the very top. 

Oh yeah, also saw Bridal Veil Falls nearby, which has, y'know, seen better days, but is still running, after a fashion.

Or maybe you'd like some historical trivia? One of the things I wanted to find more about for the Mammoth Cave National Park paleo inventory was whatever became of the type specimen of Lithodrumus veryi, a Mississippian coral possibly collected from the park. It was described in 1904 by George Greene and the specimen has been lost to science since at least 1944. Greene's collection went first to the American Museum of Natural History and then to the National Museum of Natural History, but Lithodrumus veryi apparently went missing, as Easton (1944) couldn't find it at the AMNH. Just a few days ago I was looking up images of tabulate corals when I came upon a post at Louisville Fossils and Beyond that stated one cabinet had not been sold, and its contents were eventually going to the Indiana State Museum. Hope springs eternal! I've sent a message to the Indiana State Museum to see if perhaps L. veryi's type is there.

It's supposed to look like this (Greene 1904: Plate 49). Have you seen it, by any chance?

So that's what's going on around here. (Oh, that and some unusually intense bot "readership", or the whole of Hong Kong has suddenly discovered a passionate interest in the Ordovician fossils of Minnesota and Elliot Formation prosauropods. Maybe I'm cynical.) Tune in for the next post!

References

Allen, H. J., E. W. Wilberg, A. H. Turner, and D. J. Varricchio. 2025. A new, diminutive, heterodont neosuchian from the Vaughn Member of the Blackleaf Formation (Cenomanian), southwest Montana, and implications for the paleoecology of heterodont neosuchians. Journal of Vertebrate Paleontology e2542185. doi: https://doi.org/10.1080/02724634.2025.2542185

Easton, W. M. 1944. Revision of Campophyllum in North America. Journal of Paleontology 18(2): 119–132.

Greene, G. K. 1904. Contribution to Indiana palæontology 1(17): 168–175.

Whatever Happened to Euskelosaurus?

In principle, any dinosaur name considered dubious or a synonym can be brought back into use unless it is an objective junior synonym (based on the same specimen as a previous name) or suppressed by the International Commission on Zoological Nomenclature. In practice, some names are much more likely than others to rise out of the mire. For example, when was the last time you thought about Polyonax? (Is this the first time you've ever had occasion to think about Polyonax?) Then there are historical names based on questionable material that once were widely used but have now fallen completely out of favor. Think Monoclonius, Palaeoscincus, or Trachodon, long-time favorites that have staggered into obsolescence and cheap dinosaur toys.

Pulaosaurus qinglong

We're admittedly getting to Pulaosaurus qinglong a bit after announcement and description, but what's a few weeks after 160 million years? "Hypsil"-type dinosaurs are always going to be welcome here, and this one also has potential gut contents, albeit left for a later date, so let's extend a welcome and have a look after the jump break.

Minnesota Spring Inventory

While working on something else recently, I stumbled on the Minnesota Spring Inventory. More importantly, especially in terms of time consumption, the inventory also has a great interactive map.

See any patterns?

You can zoom and select any point, or if you are interested in a particularly named spring you can type the name in the search bar to find it. Clicking on a point gives a pop-up box with information about the spring, and sometimes there are one or more links to photos.

For example, this is Big Spring at Beaver Creek State Park.

Incidentally, it looks like this. Highly recommended!

If you're curious about what the various data fields are measuring, the rules and guidelines used for documenting Minnesota springs are in this document.

If you're curious about natural or human features, there's a lot of interest to be had playing around with the map. Why human features? Springs that flow reliably and with a significant volume are great places to live, especially in the days before municipal waterworks, and many of these are of archeological and historical interest. For example, fooling around in the Minnesota River Valley area, I stumbled on one called Sacred Heart Geyser with this note: "Water was bottled and shipped to Chicago about 1900."

Obviously springs are significant natural features as well, but not only as just springs. At a glance at the map, for example, it's very easy to pick out the karst terrain of Minnesota. They can also be used as proxies for geological horizons. Springs form where groundwater isn't moving down, but laterally out of a surface. One of the biggest reasons why groundwater isn't moving down is because the flow is interrupted by a change in lithology. Maybe there's porous sandstone over non-porous shale, or limestone with fractures over shale, or pretty much anything over shale. Groundwater hits the non-permeable surface and spreads laterally, and if it runs into open air before it finds a path downward, a spring may form. (Or just a seep, which is more diffuse and less powerful.) As noted in the document, there are several geological contacts that are prone to this in Minnesota, such as glacial drift over the Decorah Shale and the Magnolia Member of the Platteville over the more shaley Hidden Falls Member. Many of the springs in Minneapolis and St. Paul are probably at one of these two contacts. (Less scientifically, springs in some settings will be found with exposed bedrock, and anywhere you've got exposed bedrock and flowing springs has a good chance of interesting geological and scenic viewing...*)

(*as always, land ownership restrictions and common sense about terrain awareness apply) 

Going up (and down!) in section, southeastern Minnesota

I was recently on a trip to southeastern Minnesota for several days. While there, inevitably I ended up with some geological photos. I must confess that I've never spent any time down here outside of passing through, and this was actually the first occasion I'd spent any time above the Cummingsville, so it was nice to see the overlying units I'd only read about before. If you'd like to know what we're up against, I recommend the Minnesota Geological Survey's geologic atlas of Fillmore County (the trip wasn't entirely in Fillmore County, but you'll get the idea).

Fossil Ursids of the National Park Service

Here we are, a little later than usual for the annual "Fossil [Group] of the National Park Service" but present nonetheless. I decided to get in another group of carnivorans, this time the bears (Ursidae). If you've been here for the cats and dogs, you can probably guess the basic shape of things: a few records earlier in the Cenozoic, then a big slug in the Pleistocene into the Holocene, with many of the records representing living species. That is indeed how it goes with the bears as well, with a couple of quirks: the bear record goes back as far as the dog record, into the Chadronian (Late Eocene), but is never as diverse as the records for either cats or dogs. Essentially you get a couple of genera or species in a formation, and that's about it. Bears, it seems, have never felt the need to be profuse about the business of being bears. It's not quite "one-bear-fits-all", but bears favor generalism, and there's only room for so many species of generalists in one place and time.

Odds and Ends

Several items worth a brief comment...

Minnesota state fossil

By Minnesota law, Castoroides ohioensis, the "giant beaver", became Minnesota's state fossil on July 1, 2025. C. ohioensis is a rare case of a write-in winning (the original slate of candidates did not include it). Call me a lot of things (crazy, no fun, pessimistic about human nature), but I have a hard time believing the giant beaver won based on its merits as a fossil found in Minnesota, as opposed to a bunch of people on the Internet thinking it would be funny for Minnesota's state fossil to be a giant beaver. Which, I suppose, is a reasonable response to the idea of state symbols in general and state fossils in particular.

Enigmacursor

Speaking of pessimism, no sooner are all previous names for Morrison Formation "hypsils" declared dubious than a paper by the same authors comes out naming... a new Morrison Formation "hypsil". (Not the first time this kind of thing has taken place, but usually the declaration of invalidity and the new name come in the same paper.) Anyway, the new one is Enigmacursor mollyborthwickae, based on a partial skeleton from Colorado featuring vertebrae, ribs, and most of the girdles and limbs. It's not one of the more famous (well, "famous", but bear with me) unnamed Morrison specimens like BYU ESM 163R or "Barbara" but instead was recovered a few years ago. (Some days it feels like famous unnamed specimens *never* get described, but I digress.) Given the matter of chimeric specimens discussed in the Nanosaurus paper, a quarry map would have been a nice inclusion. E. mollyborthwickae looks and phylogenetically acts about the way you'd expect out of a Morrison "hypsil". I wouldn't expect that it actually *is* the closest relative to Yandusaurus hongheensis among all dinosaurs known to date (for one thing, that's putting a lot of faith in Yandusaurus, one of those dinosaurs that seems more complete and better known than it is), but that's about where Morrison "hypsils" always turn up.

Tyrannosaurus

Meanwhile, on the reincarnation of the Dinosaur Mailing List, the Dinosaur Mailing Group, the hot topic has been Tyrannosaurus and nanotyrants. Some things never change, after all. It's not a spontaneous irruption, but driven by Gregory Paul's new review of latest Cretaceous western North American tyrannosaurs. This follows on the paper a couple of years ago proposing to split T. rex into three species. Paul further revises Tyrannosaurus in this paper, supporting not only the removal of Nanotyrannus lancensis, but also Stygivenator molnari. (If you remember using the word "aublysodont" seriously, you know what's up, and there's an excellent chance you're also at least 40.) These two species are interpreted as having crossed over from Appalachia (the eastern half of North America) with the dwindling of the Western Interior Seaway, which is a fascinating idea worth further consideration.

Will it settle anything? I don't expect it to. At this point it feels like everyone is well beyond being burned out on the topic and is firmly set in their ways. It doesn't help that Nanotyrannus, along with spinosaurs and exaggerated dinosaur sizes, is one of the most obnoxious long-running issues among dinosaur enthusiasts, providing further incentive to stay the heck away. It's a pity, though, because there's something something weirdly Alioramus-y going on with some of these specimens, and all we ever do is circle around the same couple of points we always have.

Probably 25 years ago or more I posted something on the DML about a taxonomic discussion, and another member suggested we'd be better off not using the names at all and just using specimen numbers. I thought it was crazy at the time since they're a lot harder to keep track of, but with time I'm definitely coming around to the idea.

Lithodendron

The second issue of Lithodendron, Petrified Forest National Park's journal, has just come out. Contributions to this issue include a report of a large silesaurid (and coelophysoid) from the park, and the printing of a previously unpublished stratigraphic study of the park from 1940 (always good to see formerly overlooked research get published!).

Cenozoic Life and Mesozoic Life in the National Parks coloring books

Finally, some uncomplicated fun: We've recently produced coloring books of Cenozoic and Mesozoic life in the National Parks to go with the Prehistoric Life in the National Parks coloring book from a few years back. They feature a mix of artwork from the original coloring book plus newer pages. The links above will take you to the data store pages for the two books, which can be downloaded freely as pdfs. Parks with notable Cenozoic or Mesozoic fossils may also have physical copies available.

References

Maidment, S. C. R., and P. M. Barrett. 2025. Enigmacursor mollyborthwickae, a neornithischian dinosaur from the Upper Jurassic Morrison Formation of the western USA. Royal Society Open Science 12(6):242195. doi: https://doi.org/10.1098/rsos.242195

Paul, G. S. 2025. A presentation of the current data on the exceptionally diverse non-tyrannosaurid eutyrannosaur and tyrannosaurini genera and species of western North America during the End Cretaceous North American Interchange. Mesozoic 2(2): 85–138. https://doi.org/10.11646/mesozoic.2.2.1

Feeding Diamantinasaurus

It's no secret that we like titanosaurs here at Equatorial Minnesota, and another favorite topic is potential gut contents in herbivorous dinosaurs (albeit we don't see a lot of papers on it). When Poropat et al. (2025)'s paper on the potential gut contents of a specimen of Diamantinasaurus matildae came out a couple of weeks ago, it was an obvious paper to cover here. True, diamantinasaurus may not be titanosaurs by phylogeny, but they're close, and they're still titanosaurs in our hearts.

The Find

A weekend or so back I traveled down to Whitewater State Park to participate in their Fossil Field Trip event. As part of this, visitors go to a site off of the park where they can hunt for fossils (participants can keep two). It was very well presented and attended! The site is in the Cummingsville Formation, which is the limestone that follows the Decorah Shale. (Actually, it's a bit more complicated than that: much of the Cummingsville Formation in Olmsted County is roughly the offshore equivalent of the more nearshore Decorah of the Twin Cities, so where we were is probably equivalent to a horizon in the upper half of the Decorah quarry walls at the Brickyard.) Unlike the distinctive green-gray Decorah, the Cummingsville presents as a tan unit. It's a shaly limestone at the site, so it doesn't quite have the strength of a pure limestone, but it does better than the Decorah.

Weathering Cummingsville at the site.

The Cummingsville is not as fossiliferous as the Decorah, even accounting for my different search image, and the site is a known fossil site, so the going was slow at the beginning. Then I found this:

Well, I did do a little prep work and applied water before taking this picture, and the rock is about palm-sized all told, but you get the idea.

I'd been hoping to find a receptaculitid, because those are common in the Cummingsville and I haven't found one, but this was something better. Even in an unprepared state it was obviously an echinoderm based on the plates, but what kind? A crinoid calyx? Something else?

If you know what the arrows are pointing to...

The prep work exposed the little vent-like features, meaning it was some kind of rhombiferan cystoid. The cleaning also made it more obvious where the plates met, which can be kind of confusing when there are all these ridges to mislead the eye.

A handy diagram! The dotted lines are where the plate edges become obscure or are lost.

If you don't know what it is, check this out:

Figure 1 from Parsley (1970), with red lines and dots added by me. Original image CC BY-NC-SA 3.0.

Yup, this is a Pleurocystites, or at least a good-sized chunk of one. Either P. squamosus (Parsley 1970) or P. strimplei (Brower 1999) would be appropriate for the Cummingsville; there's not really enough there to tell between them, but I'm not complaining! It's fun to be in the right place at the right time with the right knowledge and experience.

References

Brower, J. C. 1999. A new pleurocystitid rhombiferan echinoderm from the Middle Ordovician Galena Group of northern Iowa and southern Minnesota. Journal of Paleontology 73(1): 129–153. 

Mossler, J. H. 2008. Paleozoic stratigraphic nomenclature for Minnesota. Minnesota Geological Survey, St. Paul, Minnesota. Report of Investigations 65.

Parsley, R. L. 1970. Revision of the North American Pleurocystitidae (Rhombifera-Cystoidea). Bulletins of American Paleontology 58(260): 135–213. 

The Continuing Story of Nanosaurus agilis

Recently Barrett and Maidment (2025) published a paper on the state of Morrison hypsilophodont-things, which is of great interest here because after two long posts on Nanosaurus agilis, we're solidly invested in its fate. How did it fare? Short answer: not very well. But, on the other hand, neither did anybody else. Well, Drinker was shown more appreciation than probably anyone has given it since 1990, but that's not saying much.

So, what to call these happy fellows at the Denver Museum of Nature and Science?

Barrett and Maidment (2025) went over the type specimens of N. agilis, N. rex (Othnielia), Laosaurus celer, L. consors (Othnielosaurus), L. gracilis, and, via illustrations, Drinker nisti. (There is a slight advance on Carpenter and Galton 2018, in that we now get the implication that Bob Bakker has D. nisti as opposed to the whereabouts being unknown.) They find none of the type specimens to be diagnostic. The one that comes off the worst is L. consors. Material cataloged as the type is an assemblage rather than an individual. To be fair, Marsh knew he had multiple individuals at the time, but then he should have been more careful about specifying a type. The parts that had been on display as a panel mount at Yale may be one associated individual, consisting of most of the cervical and dorsal series (just centra), possibly six sacrals, parts of the left shoulder girdle and the pelvis, a partial left femur and complete left foot, and parts of the right femur, tibia, and astragalus. This is a lot of parts/partials plus a lot of plaster, which is not encouraging. The rest of the material is a chimeric mix including at least juvenile dryosaur and hypsil material. The only one of the six that ends up being interesting is D. nisti, which has some dental and jugal features reminiscent of pachycephalosaurs (but is still not diagnostic, although it would be nice to have the type material in hand to be sure).

Where does this leave the Morrison hypsil(s), which Carpenter and Galton (2018) had declared N. agilis? Anonymous, for the time being. Carpenter and Galton (2018) looked upon the pile of Morrison hypsil bits and proposed it was "all" Nanosaurus agilis. Barrett and Maidment (2025) looked upon the same pile and clutch of names and regarded it as a taxonomic dead end, to be set aside to allow a fresh start for more complete and better preserved specimens (with quarry maps and documented associations and such).

At heart, we're seeing two different approaches to taxonomy, and which one you choose depends on how pragmatic you are and how bound you feel by existing names. If you want a species with a holotype featuring robust apomorphies, N. agilis is not for you. We saw that in the comments section of the last post: most of the characters cited by Carpenter and Galton (2018) are widely distributed among hypsil-things, with just a couple that might have some particular use. However: Is there a hypsil-thing in the Morrison that is anatomically consistent across specimens, whether or not said specimens are diagnostic across Ornithischia? If so, is it reasonable to call this hypsil *something*, knowing that it may be revised later? If so, the oldest existing name is Nanosaurus agilis. If you go that route, I'd recommend looking into a neotype, though. ("All would be well, if, if, if, if, if...")

References

Barrett, P. M., and S. C. R. Maidment. 2025. A review of Nanosaurus agilis Marsh and other small-bodied Morrison Formation “ornithopods". Bulletin of the Peabody Museum of Natural History 66(1): 25–50. doi: https://doi.org/10.3374/014.066.0102.

Carpenter, K., and P. M. Galton. 2018. A photo documentation of bipedal ornithischian dinosaurs from the Upper Jurassic Morrison Formation, USA. Geology of the Intermountain West 5:167–207. doi:

The downside of reef building?

I was reviewing text for a website a few weeks back dealing with aspects of the history of life, and a couple of things struck me about biological reefs. First, a quick look at reef-builders through time (a useful overview can be found here if you'd prefer more flesh on the bones, or corallites or shells or whatever may be more appropriate):

The first multicellular reef-builders were the archaeocyathan sponges, who flourished briefly in the Cambrian but did not even make it to the end of the period. Corals, in the form of rugose and tabulate corals, spread in the Ordovician but took a while to make reefs. They were joined by stromatoporoid sponges (layered like stromatolites, spelled like stromatolites, but not stromatolites) and various microbes, with the Devonian being an apex of reef-building. The classic stromatoporoid-tabulate reefs of the Devonian went kaput in the End-Devonian extinction. Permian reefs were a conglomeration of just about everything that couldn't get out of the way: various algae, sponges, bryozoans, and other less obvious things. This assortment bought it at the end of the Permian. False starts with scleractinian corals in the first part of the Mesozoic gave way to the rudist bivalves in the Cretaceous. The rudist reefs went out with non-avian dinosaurs, marine reptiles, pterosaurs, ammonites, and so forth at the end-Cretaceous extinction. Finally we get to the big scleractinian coral reefs in the Cenozoic, with some sponge and oyster reefs and such for variety.

So far, a typical pattern: group of organisms branches into reef building, reefs spread and become ecologically complex, reefs flourish for a while, mass extinction wipes out reefs. Then after a hiatus reefs become fashionable again, with some other group laying the foundation for a new iteration, and the cycle continues. A couple of observations come to mind. First, reefs seem to be an obvious evolutionary path for immobile marine invertebrates. It may take some time, but some group always takes up the baton after another falters.

Then, the other shoe. What happens to the previous reef-builders? Seen any archaeocyaths lately? Any vacation packages advertising stromatoporoid reef visits for their island getaways? Run across any rudists while snorkeling? Could it be that once a group goes all-in on the reef habit, it's stuck with it?

Furthermore, reefs have a habit of getting smacked in mass extinction events. Does a reef inherit a narrowing range of environmental restrictions from its components as its complexity increases? Does it become vulnerable to unpredictable instability, such as some minor constituent going through a bad patch leading to collapse via a Rube Goldbergian-cascade of events? More broadly, does reef building amount to an evolutionary Faustian bargain, in which a group becomes dominant for a while by locking itself into a doomed arrangement? (Granted, we're all doomed in the final analysis, but some of us are more obviously doomed than others.) Or am I just playing the gloomy Minnesotan?

Arcola Bluffs Day Use Area

It's been a good while since our last St. Croix post, hasn't it? I wanted to let you in on a fun place I recently visited for the first time; Arcola Bluffs Day Use Area. This site is not widely known; there's this article (with much more artistic photos than my own), an NPS cultural landscape assessment from 2018 that weighs in at 204 mb (NPS 2018; absolutely worth it if you want a thorough understanding of the site, and also includes a section on Fairy Falls), and then some short pieces here and there, and that's about it. Visiting it, though, you'll discover great geology, views of the river and the historic Soo Line High Bridge, forest and prairie settings, and some evocative ruins.

Titanosaur osteoderms, 2025 update

Occasionally I glance at the site statistics, but since I don't use anything detailed there's not a lot to get from them except that it's fun to see where the obvious spoof hits are coming from (Singapore, lately; sometimes Hong Kong or Russia). Sometimes I can tell that a post has been picked up elsewhere and gotten a few views. Over the last month or so, there have been an unusually large number of visits to "Titanosaur osteoderms: functions and conclusions", and at the same time a similar number of visits have come from a service at the University at Buffalo. Conclusion? Seems like someone at the university is using the post in a course. If that's what's going on, this one is for you!

Since I wrote the osteoderm series back in 2019, there have a few reports of interest on titanosaur osteoderms. I added several overlooked and new records to the distribution post in 2019–2020, and there hasn't been much change there since. One tangential note, also applicable to "Titanosaurs of Yesterday", is a further advance in the study of Agustinia ligabuei, the spiky sauropod that wasn't. Bellardini et al. (2022) published an analysis that found A. ligabuei was not a titanosaurian or even a macronarian, but a rebbachisaurid. (And it's still not armored, either.)

As noted in the Menucocelsior arriagadai post, Rolando et al. (2022) was not just a description of a new titanosaur taxon, but also included reports of material from other titanosaurs. Among these specimens were four isolated osteoderms from the Cerro Matadero site of the Allen Formation. Three represent the "ellipsoidal" form (bulb-and-root) of D'Emic et al. (2009) and the other is a "keeled" osteoderm. Rolando et al. interpreted the keeled osteoderm as perhaps from the tail or back of a saltasaur and the two more complete ellipsoidal osteoderms as perhaps aeolosaur osteoderms from the hip region.

Another report also discussed in another post is Fronimos (2021) on an osteoderm of a Big Bend titanosaur (e.g., Big Bend Alamosaurus). To paraphrase, this is a large oval and unkeeled osteoderm from the upper Javelina. It is tall, symmetrical, 19.9 cm long (7.83 in), not hollow, not differentiated into a bulb and root, and does not have a cingulum. It resembles the North Horn Formation Alamosaurus osteoderm and the osteoderms of Mendozasaurus neguyelap and unnamed South American forms (Fronimos 2021). Fronimos (2021) regarded the most likely functions as mineral storage, local defense, and display.

Concerning the function of titanosaur osteoderms, Silva Junior et al. (2022) published a study using finite element analysis to evaluate titanosaur osteoderms versus likely titanosaur adversaries, specifically the bites of abelisaurs and baurusuchid crocs. They found that bites had less of an effect on solid osteoderms (i.e., those without hollow internal spaces), and interpreted this to indicate that solid osteoderms could do more than provide mineral storage. On the other side of the mineral storage question, Broeckhoven and du Plessis (2022) made an analysis of osteoderms in armadillo lizards. Using micro-computed tomography, they found that the female lizards in their study had denser, more compact osteoderms than males, and observed the presence of TRAP-positive cells (tartrate-resistant acid phosphatase, involved in bone resorption and breakdown). Denser osteoderms may help maintain a minimum level of mineral density for reproduction and provide defensive strength to osteoderms also being used for a mineral storage function. The authors did not find a difference in density between seasons, and suggested this meant the osteoderms were only subject to resorption during particularly stressful conditions, or that it only took place during certain phases of embryo growth. They concluded that mineral storage for reproduction is a plausible function for osteoderms in female reptiles.

Finally, I've saved the most interesting update for last. In an abstract, Filippi et al. (2023) described an articulated tail, MAU-Pv-CO-726, from the Bajo de la Carpa Formation of Cerro Overo–La Invernada, Patagonia, Argentina. (Yes, another Bajo de la Carpa mystery titanosaur!) MAU-Pv-CO-726 includes 25 caudals, 11 chevrons, and two osteoderms in place, with another nearby. The pair of osteoderms is associated with the last anterior caudal, found flanking the side and underside of the tail (about where the chevron articulates with the caudal). They are described as bulbous, oval, a little more than 10 cm (4 in) long, and feature a medial ridge and tapered spine on the lateral half, whereas the solitary osteoderm is described more like a classic bulb-and-root. Filippi et al. found MAU-Pv-CO-726 to be the sister taxon of Rinconsaurus caudamirus, marking the first evidence of an armored rinconsaur (and no doubt causing R. caudamirus's usual best friend Muyelensaurus pecheni great phylogenetic distress). They interpreted the location of the osteoderms as evidence of a defensive function. The osteoderms' placement also has implications for paleoart; titanosaur restorations usually put osteoderms on upper-lateral surfaces. But, then again, if you want my opinion I think titanosaurs were too diverse for a one-size-fits-all approach to osteoderms, in function, anatomy, or placement.

References (note that a couple are different from previous usage, as those were online preprints)

Bellardini, F., R. A. Coria, G. J. Windholz, A. G. Martinelli, and M. A. Baiano. 2022. Revisiting the Early Cretaceous sauropod Agustinia ligabuei (Dinosauria: Diplodocoidea) from southern Neuquén Basin (Patagonia, Argentina), with implications on the early evolution of rebbachisaurids. Historical Biology 35(12): 1–27. doi: https://doi.org/10.1080/08912963.2022.2142911

Broeckhoven, C., and A. du Plessis. 2022. Osteoderms as calcium reservoirs: Insights from the lizard Ouroborus cataphractus. Journal of Anatomy 241(3): 635–640. doi: https://doi.org/10.1111/joa.13683

D'Emic, M. D., J. A. Wilson, and S. Chatterjee. 2009. The titanosaur (Dinosauria: Sauropoda) osteoderm record: review and first definitive specimen from India. Journal of Vertebrate Paleontology 29(1):165–177.

Filippi, L. S., F. Bellardini, A. Paulina-Carabajal, P. Cruzado-Caballero, J. González-Dionis, A. H. Méndez, F. Gianechini, K. Ulloa-Guaiquin, A. Garrido, I. Maniel, Y-N. Lee, and K. Do-Kwon. 2023. Articulated osteoderms on a titanosaur tail from Cerro Overo–La Invernada (Bajo de la Carpa Formation), Upper Cretaceous, Northern Patagonia Argentina: Paleobiological and paleoecological implications. Publicación Electrónica de la Asociación Paleontológica Argentina 24(R3): R67–R68.

Fronimos, J. A. 2021. Morphology and neurovascular anatomy of a titanosaur (Dinosauria, Sauropoda) osteoderm from the Upper Cretaceous of Big Bend National Park, Texas. Cretaceous Research 120: 104670. doi: https://doi.org/10.1016/j.cretres.2020.104670

Rolando, M. A., J. A. Garcia Marsà, F. L. Agnolín, M. J. Motta, S. Rozadilla, and F. E. Novas. 2022. The sauropod record of Salitral Ojo del Agua: An Upper Cretaceous (Allen Formation) fossiliferous locality from northern Patagonia, Argentina. Cretaceous Research 129: 105029. doi: https://doi.org/10.1016/j.cretres.2021.105029

Silva Junior, J. C. G., F. C. Montefeltro, T. S. Marinho, A. G. Martinelli, and M. C. Langer. 2022. Finite elements analysis suggests a defensive role for osteoderms in titanosaur dinosaurs (Sauropoda). Cretaceous Research 129: 105031. doi: https://doi.org/10.1016/j.cretres.2021.105031