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.

Prosauropod rehab

I once wrote that "When you get interested in something, what's already there when you arrive will seem like it has always been there, and everything that comes along later will always seem a little new to you, even if it has been decades." Taxonomic fashion is one of those things. When I got into dinosaurs in the mid-1980s, prosauropods had shrunk to something like a half-dozen well-regarded genera plus some dross and a few names that were too new for someone to have gotten around to sinking. One of the survivors was always Ammosaurus, which somehow parlayed a few pelvic differences from Anchisaurus into family-level separation (Ammosaurus the plateosaurid, Anchisaurus the anchisaurid). Meanwhile, all of Africa's prosauropod diversity was being crammed, with varying degrees of success, into Euskelosaurus, Massospondylus, these things called "roccosaurids", and Aristosaurus. (Aristosaurus? You had to be there.) Since I had no personal memory of the work that had gone on in years before, and at five years old wasn't really pouring through the primary literature, it seemed like things had always been that way. This is not the case, though. The vast majority of this synonymization had taken place in the previous 10–15 years and was the work of just a few authors. What had come before that?

Well, obviously, what had come before that were the decades of describing and naming scads of prosauropods; otherwise, there wouldn't have been anything to synonymize. It seems like people loved to name them and then forget about them. The history of their research is littered with misconceptions, some of which seem blindingly obvious in hindsight (big predatory prosauropods? You sure you're not just looking at shed crowns from carnivorous animals?). It's a matter of lack of respect and interest: If dinosaurs were a sideshow to the evolution of important things (i.e., mammals), prosauropods were a sideshow to the sideshow, a couple of card tables featuring a guy with a big unibrow and someone doing "pull my finger" jokes.

Skip ahead a few decades. Some of my readers will have never heard of Aristosaurus (well, okay, nobody living has heard of Aristosaurus except me and maybe 10 other people who have been sworn to secrecy, and now I've lost my membership in the group for a cheap gag) and know only of Euskelosaurus as this thing that nobody talks about except to badmouth in the "Background" section of papers. You wouldn't remember the heady days of the '00s, when that genus collapsed under the weight of its synonyms. Instead, it will have always been a nomen dubium to you, and if it somehow returned (stranger things have happened), it wouldn't be a resumption of your interrupted Euskelosaurus service but something new. (Why, yes, I do have a sense I'm getting older. The only thing that saved me from being older than all MLB players this year was four games and 3 2/3rds innings from Rich Hill.)

Anyway, I've long had a soft spot for prosauropods. They're never been supremely popular and had the poor marketing insight to go extinct before they could be menaced by particularly charismatic theropods, but they have a subtle charm, like small bipedal ornithischians. Over the past year or so, there's been a series of papers providing redescriptions and historical analyses of some of the most venerable genera and species of prosauropods, including comments on some of those 1970s–1980s synonymizations. They include Barrett and Chapelle (2024) on Massospondylus, Barrett and Choiniere (2024) on Melanorosaurus, and Regalado Fernández et al. (2023) and Schaeffer (2024) on Plateosaurus.

Massospondylus

Another instance of perception not matching up with chronology: one of the things that comes out of Barrett and Chapelle (2024) was that before the 1970s, Massospondylus was kind of a dog of a taxon. It achieved its present exalted status mostly by getting there first and being less of a dog than the other choices. If Richard Owen had decided to lead with Leptospondylus or Pachyspondylus in 1854 instead of Massospondylus, it wouldn't have had that first point in its favor, and then where would we be? When a name was needed to absorb the mid-sized prosauropods of the Early Jurassic of southern Africa, M. carinatus was there waiting. Some of the new material was good enough to support a name, and our conception of Massospondylus carinatus drifted from the varied bones named by Owen to that material, which eventually supplied a neotype (a type or name-bearing specimen selected after a species was named, to replace a type that was lost or not specified). The original material was no longer available to query, having been destroyed in World War II, but casts and line drawings of some of it remain, and Barrett and Chapelle (2024) provided redescriptions of as much as they could. In the end, it's just as well that the species is now represented by a neotype, because that original material is not diagnostic beyond the level of Massospondylidae at best. The drift of the conception of Massospondylus, plus the uncritical assignment of a lot of material since the 1970s, leaves the possibility of surprises once "Massospondylus" specimens are thoroughly examined. This also applies to Melanorosaurus and Plateosaurus. (Granted, I'm not convinced that every new genus and species will stand the test of time, but that's for another time, when the pendulum eventually swings back to consolidation.)

Massospondylus as a flowchart. Figure 4 of Barrett and Chapelle (2024). CC BY 4.0.

Melanorosaurus

Proposed synonymizations don't always pan out, of course (one reason not to go overboard committing to the conclusions of New Paper Of The Week). The proposed sinking of Lufengosaurus and Yunnanosaurus into Massospondylus at the height of prosauropod consolidation did not attract much support, and the proposed sinking of Melanorosaurus into Euskelosaurus fared only marginally better. Although it escaped that fate, Melanorosaurus has never seemed to get much respect. It's almost a reflex for authors to mention its questionable type material, as if apologizing for trotting it out. Barrett and Choiniere (2024) went back to the original syntype series (a group of specimens used to bear a name) to see what could be salvaged.

M. readi, like many a species, was based on your standard pile o' bones of unclear association. Barrett and Choiniere (2024) regarded a subset, including several partial or complete vertebrae, an ulna, a radius, an ilium, a partial pubis, two tibiae, a fibula, and parts of four metatarsals, as representing one skeleton that they designated a lectotype (a name-bearing specimen selected from a syntype series). Furthermore, they were able to find several diagnostic features. It's not the prettiest type, but it *is* useable. The authors compared the lectotype to several other specimens frequently used as Melanorosaurus exemplars in the literature. Unfortunately, this part doesn't go quite so well for M. readi; NMQR 3314, one of the more important specimens (including a skull), is excluded on both anatomic and stratigraphic grounds. Another specimen, NMQR 1551, is considered consistent with M. readi but an assignment is not confirmed. (I personally hope it is, because I don't want an excuse for someone to try to bring back "Roccosaurus" as a name.) The upshot is M. readi is valid but not particularly helpful at the moment, pending reclamation of more complete material.

Plateosaurus

If Massospondylus is what you get when specimen assignment goes to the last species standing, Plateosaurus is what happens when people care too much. For sheer mind-numbing taxonomic complexity in the world of dinosaurs, it's hard to beat the adventures of Upper Triassic European prosauropods. An extensive historical review can be found in Regalado Fernández et al. (2023), along with detailed reports on many of the significant specimens involved in the story. One of the things I find useful is rather than synonymize the many lesser lights with a species of Plateosaurus, the authors treat them as dubious species. Why is this useful? Keeping them separate helps to keep the concept of the host species from drifting. It's not as if there's some prize for having the tidiest faunal list, after all; specimens should only be assigned to the level of confidence.

Stratigraphic distribution of German Plateosaurus and friends. I could quote every figure in this article, but then I might as well reproduce the whole thing. Figure 2 in Regalado Fernández et al. (2023) CC BY 4.0.

Coming at Plateosaurus from a different angle, Schaeffer (2024) redescribed the holotype (singular name-bearing specimen) of P. trossingensis, which became the type species of Plateosaurus a few years back (another long story). This gets right at the heart of the taxonomic drift: you have to establish your basis of comparison before you can compare anything to it. Continuing with this, Schaeffer ran a phylogenetic analysis of sauropodomorphs with Plateosaurus trossingensis based just on the holotype. Fortunately, the species behaved predictably in terms of general location and neighbors.

References

Barrett, P. M., and K. E. J. Chapelle. 2024. A brief history of Massospondylus: its discovery, historical taxonomy and redescription of the original syntype series. Palaeontologia africana 58: 97–131.

Barrett, P. M., and J. N. Choiniere. 2024. Melanorosaurus readi Haughton, 1924 (Dinosauria, Sauropodomorpha) from the Late Triassic of South Africa: osteology and designation of a lectotype. Journal of Vertebrate Paleontology 44(1): e2337802. doi: https://doi.org/10.1080/02724634.2024.2337802

Regalado Fernández, O. R., H. Stöhr, B. Kästle, and I. Werneburg. 2023. Diversity and taxonomy of the Late Triassic sauropodomorphs (Saurischia, Sauropodomorpha) stored in the Palaeontological Collection of Tübingen, Germany, historically referred to Plateosaurus. European Journal of Taxonomy 913(1): 1–88. doi: https://doi.org/10.5852/ejt.2023.913.2375

Schaeffer, J. 2024. Osteological redescription of the holotype of Plateosaurus trossingensis (Dinosauria: Sauropodomorpha) from the Upper Triassic of SW Germany and its phylogenetic implications. Journal of Systematic Palaeontology 22(1): 2335387. doi: https://doi.org/10.1080/14772019.2024.2335387

Mbiresaurus and Tuebingosaurus

Things have picked up since late June. In the past couple of weeks, two new "prosauropods" have been published that cover both ends of the prosauropod spectrum. One, Mbiresaurus raathi, represents the early part of sauropodomorph evolution, while the other, Tuebingosaurus maierfritzorum, is close to the transition from clear-cut "prosauropods" to clear-cut "sauropods". Long-time readers will be familiar with the author's inexplicable fondness for prosauropods, so let's invite them in.

Issi saaneq and Brighstoneus simmondsi

A couple of new non-theropod names have come down the pipe since the beginning of November, and I thought I'd cover them together. Let's tackle the earlier of the two (both geochronologically and publication-wise) first.

Kholumolumo ellenbergerorum and "Thotobolosaurus"

With the twin unofficial missions of Equatorial Minnesota to cover Elliot Formation "prosauropods" and find out what happened to unpublished dinosaurs of the past, it was imperative that Kholumolumo ellenbergerorum receive a post. Before we get into the actual science of K. ellenbergerorum, though, let's step back through the mists of time. First we'll stop off in the far-off world of 1985.

You've just gotten the Normanpedia (Norman 1985) and are busy committing it to memory. On page 97, in the "Dubious" section of the list of prosauropod genera, is Thotobolosaurus. It doesn't ring a bell, so you pull out "A Field Guide to Dinosaurs" (Lambert 1983). It shows up on page 104, as a "roccosaurid" with Riojasaurus and something called Roccosaurus, which wasn't in the Normanpedia. "Roccosaurids" apparently have "unusually sharp fangs and an especially strong joint linking hips with the backbone", but might actually be melanorosaurids. This last bit just ends up being confusing because (stepping outside of 1985 for a moment) "A Field Guide to Dinosaurs" was caught up in the contemporary fad for lumping all kinds of prosauropods, so Melanorosaurus is sunk into Euskelosaurus. Anyway, Thotobolosaurus is "a big prosauropod that lived in early Late Triassic Lesotho, southern Africa." Okay, sure. Does "The New Dinosaur Dictionary" (Glut 1982) have anything more, and can it explain what exactly is a Roccosaurus?

Ngwevu intloko

For our first "prosauropod" entry since Ledumahadi mafube back in September 2018, we return to the Upper Elliot Formation of South Africa. This time around, it's massospondylid Ngwevu intloko. Or, is it just a distorted, young, or otherwise odd individual of the Upper Elliot classic Massospondylus carinatus?

Ledumahadi mafube

For reasons which are mysterious to me, this blog has become full of sauropodomorphs of one kind or another. It's been a good year for them. The latest is Ledumahadi mafube from the Elliot Formation of South Africa, an early giant near the base of the sauropod tree.

Bagualosaurus agudoensis: another sauropodomorph ahead of its time

Our subject today is the newly described Bagualosaurus agudoensis from lower Upper Triassic rocks of southern Brazil. If I'd known back in March 2016 that I'd have the opportunity in a couple of years to write about another "prosauropod" that was "ahead of its time", and that it would include a partial skull justifying a terrible "head" pun, maybe I'd have come up with another title then. Oh, well; spilled milk and all that.

Xingxiulong

Two dinosaurs were published on February 16, 2017. One of them was Isaberrysaura mollensis, which has gotten a lot of press because it's a weird basal ornithischian with gut contents. The other was Xingxiulong chengi, which hasn't gotten as much attention, although the Wikipedia article is pretty extensive. Xingxiulong is among what we used to call "prosauropods", now known as basal sauropodomorphs. It is represented by most of the skeleton, excepting the tips of the jaws, most of the hands, and the coracoids and sternal elements. It also provides me a half-point on my prediction for "prosauropods", which I'll take because it's been kind of a slow year so far.

Meroktenos thabanensis: not Early Jurassic, just ahead of its time

It's been about a month since Meroktenos thabanensis made its (re)appearance, but I'm just now getting around to it. A bunch of other topics muscled ahead of it, which is fitting, I suppose. Prosauropods (by which the reader should understand "sauropodomorphs what ain't sauropods"; if you should want me to write "basal sauropodomorph" over and over again instead, I accept large cash bribes) are among those dinosaurs which never seem to get much respect. They have been fertile subjects for arm-waving, though, one result of which being that a survey of popular-audience dinosaur books will turn up wildly differing depictions over the years. Bipedal or quadrupedal? Herbivorous, omnivorous, predatory, or scavenging? A stem leading into sauropods, a bunch of rungs, or a distinct empire of prosauropods? Gleefully extreme lumping or splitting? What about herrerasaurids? What about Teratosaurus? (if that last one means anything to you, congratulations, you're in your mid-30s or older, or you study rauisuchids.) Prosauropods have also not been helped by a certain vague uniformity of body shape, nor have they been done any favors by geography. North America has an unimpressive record for these dinosaurs, so far including Anchisaurus (let's not kid ourselves about Ammosaurus being distinct), Sarahsaurus, Seitaad, the unnamed Nova Scotia form, and odds and ends. For a group that did not have horns, frills, spikes, bony armor, crests, ridge-backs, spectacular size, or big pointy teeth, and had the poor sense to go extinct before the appearance of tyrannosaurs or dromies to prey upon them, not being well-represented in North America has not helped their exposure. But I digress.

Where are they now: 1980s dinosaur rumors, sauropodomorphs

And now we return to 1982. We have seen the future of theropods, now let us turn to the future of sauropodomorphs. One future was that of Supersaurus and Ultrasaurus, a future of every large sauropod having to out-superlative the sauropod before it. This future petered out before it got any farther, kneecapped by an honest mistake half a world away and a terminal case of bonebed chimerism. Another future was that of Roccosaurus, the fanged prosauropod, and it was a terrible bleak cheesy early-'80s dystopian future, with leather and chains and synthesizers. That future never came to pass. Others involved overlooked fossils from Portugal and central Asia taking their rightful places, or promises of skulls from India, or an unusual elongate cervical vertebra from Wyoming, or a skeleton from Tendaguru.

Anchisaurus is older than you might think

Anchisaurus is kind of a second-tier dinosaur, not up there with Tyrannosaurus, Triceratops, Velociraptor, and their ilk, but it gets a fair amount of exposure, especially if you pull in the old synonym Yaleosaurus. It gets points for having a long history, having a few interesting anecdotes associated with it, being one of the first early dinosaurs known from good remains, and beginning with the letter "A"; this latter bit is important, because it means Anchisaurus ends up near the beginning of children's A-Z dinosaur books. (Is this why so many dinosaur names begin with "A"?)