It’s time to revisit everyone’s favourite trio of apocryphal super-sized sauropods! (Yes, we’ve talked about this before, but only very briefly, and that was nearly eleven years ago. Things have moved on since then.)

John Sibbick’s classic artwork showing three giant sauropods, including two of Jensen’s three. On the left is Seismosaurus Gillette 1991, which is not directly relevant to today’s post. In the middle is the brachiosaur Ultrasaurus, and on the right the diplodocid Supersaurus. Poor, unloved Dystylosaurus doesn’t get a look-in — perhaps because this was drawn before that name had been announced?

Here’s the story so far …

1. Jensen’s discoveries

In a series of expeditions beginning in April 1972, following a tip from uranium prospectors Eddie and Vivian Jones, Jim Jensen found numerous massive sauropod fossils in the Dry Mesa quarry, southwest Colorado. The Supersaurus pelvis at least was still in the ground as late as August 1972 (George 1973b:51–52) and the excavations continued into 1982 (Jensen 1985:697).

Eschewing such pedestrian venues as Science, Nature or indeed the Journal of Vertebrate Paleontology, Jensen first told the world about these finds in the popular press. The oldest widely circulated work that mentions them is Jean George’s (1973b) piece in Reader’s Digest, condensed from the same author’s piece in the Denver Post’s Empire Magazine earlier that year (George 1973a).

“‘Supersaurus,’ as we shall call him, now awaits an official name and taxonomic classification”, wrote George (1973b:53) — but the piece does not mention the names “Ultrasaurus” or “Dystylosaurus” and I’ve not been able to determine when those informal names became known to the world. (Can anyone help?) We do know that Jensen was informally using the name “Ultrasaurus” as early as 1979 (Curtice et al. 1996:87).

Anyway, for reasons that have never been very clear, Jensen concluded that the remains represented not one, not two, but three gigantic new genera: a diplodocid, which he named “Supersaurus”; a brachiosaurid, which he named “Ultrasaurus”; and an unidentifiable which he named “Dystylosaurus”. All these names were informal at this point, like “Angloposeidon” and “The Archbishop”.

2. Kim’s accidental Ultrasaurus

After Jensen had been using these names informally for some years, Kim (1983) named an indeterminate Korean sauropod as Ultrasaurus tabriensis. Based on the abstract (the only part of the paper in English, apart from the figure captions), Kim was aware of Jensen’s dinosaurs: “Judging by the large size of the ulna the animal may belong to the sauropod dinosaur, which is much bigger than Supersaurus. A new name Ultrasaurus tabriensis is proposed for the convenience of the further study.” While this does not quite go so far as to say that Kim considered the ulna to belong to the same genus as Jensen’s brachiosaur, it seems unlikely that he was aware of Supersaurus but not of Ultrasaurus, and landed independently on the latter name by coincidence. In fact, the “ulna” is a humerus, as shown by Lee et al. (1997).

Either way, in naming his species, Kim inadvertently preoccupied Jensen’s chosen genus name, with conseqences that we shall see below. By all accounts, the material the Kim described is in any case indeterminate, and the genus is generally considered a nomen nudum (e.g. Olshevsky 1991:139, Glut 1997:1001).

Kim 1983, plate 1, parts 1-3, illustrating the proximal portion of the huge “ulna” that the name Ultrasaurus tabriensis was founded on. As is apparent, this is actually the proximal end of a humerus, meaning that the animal is rather less large than Kim supposed — although the 42 cm width across the proximal end is still nothing to be sniffed at. It is about 71% the width of the 59 cm-wide humerus of the Giraffatitan brancai paralectotype MB.R.2181 (previously HMN SII).

Two years after this, and presumably unaware of Kim’s paper or incorrectly assuming his informal use of the name “Ultrasaurus” gave him priority, Jensen published a formal account of his finds, naming them (Jensen 1985). Unfortunately, while the paper does contain formal nomenclatural acts that are valid according to the rules of the ICZN, Jensen did not explain his reasoning for the creation of the new genera, and his selection of type material was problematic, as we shall see below. Also, the specimen numbers that he used have been superseded — I do not know why, but my guess would be that he re-used numbers that were already in use for other specimens, so his own material had to be given new numbers.

3. Jensen’s three sauropods

The following three genera (with their type species) were named, in this order:

1. Supersaurus vivianae, based on the holotype BYU 9025 (BYU 5500 of his usage), a scapulocoracoid measuring 2.44 m in length. To this, he referred an even larger scapulocoracoid whose length he gives as 2.70 m (though Curtice and Stadtman 2001:39 found that this length to be due to optimistic reconstruction); an ischium; either one or two mid-caudal vertebrae (his paper contradicts itself on this); and a sequence of 12 articulated caudal vertebrae. Unfortunately, Jensen’s use of specimen numbers for most of these referred elements is inconsistent, but he is at least consistent in referring to the second scapulocoracoid as BYU 5501.

Supersaurus vivianae referred scapulocoracoid BYU 12962, photographed at the North American Museum of Natural Life. The exhibit text reads: “Supersaurus scapula and coracoid. This is the actual Supersaurus bone that the world saw when the announcement was made of the new animal’s discovery in 1972. The scapula lay in the ground for five more years, waiting for the collection of other fossils that lay in the path of excavation. The flatness of the bone presented a challenge to “Dinosaur Jim” Jensen, who had to figure out a way to get the bone safely out of the ground. He finally accomplished this by cutting the scapula into three pieces. In 1988, Cliff Miles, Brian Versey and Clark Miles prepared the bone for study. It is still one of the largest dinosaur bones known in the world. Specimen on loan from Brigham Young University’s Earth Science Museum. Late Jurassic/Early Cretaceous (about 144 million years ago)

2. Ultrasaurus macintoshi, based on the holotype BYU 9044 (BYU 5000 of his usage), a dorsal vertebra measuring 1.33 m in height. To this, he referred BYU 9462 (BYU 5001 of his usage), a scapulocoracoid measuring 2.7 m in length; BYU 9024 (BYU 5003 of his usage), a huge cervical vertebra; and an anterior caudal vertebra.

Ultrasaurus macintoshi holotype dorsal vertebra BYU 9044, photographed at the North American Museum of Natural Life. (It’s incredibly hard to photograph well because it’s behind reflective glass.)

3. Dystylosaurus edwini, based on the holotype BYU 4503 (BYU 5750 of his usage), a dorsal vertebra. He did not refer any other material to this taxon, and considered it “Family indeterminate” commenting that it “no doubt represents a new sauropod family”. Poor Dystylosaurus has always been the unloved member of this group, and pretty much ignored in the literature aside from the Curtice & Stadtman (2001) synonymisation paper discussed below.

Dystylosaurus edwini holotype BYU 4503, a diplodocoid anterior dorsal vertebra.

In a subsequent paper, Jensen (1987:600–602) removed the big cervical BYU 9024 (BYU 5003 of his usage) from Ultrasauros and reassigned it to Diplodocidae. The text of this paper never refers it to Supersaurus vivianae in particular, but it is illustrated and captioned as belonging to that taxon (Jensen 1987:figures 7A-B, 8C), and this assignment is generally assumed to have been meant.

When Jensen became aware of Kim’s (1983) preoccupation of the name Ultrasaurus, he recognised that his own genus needed a new name. At his suggestion, Olshevsky (1991) erected the replacement name Ultrasauros (with a single-letter spelling difference) for Jensen’s taxon based on the dorsal vertebra BYU 9044. We will use this revised spelling hereon, and the taxon Ultrasaurus Kim 1983 is of no further interest to this story.

The relevant extract from Olshevsky (1991:139).

4. Curtice’s synonymies

This was how things stood, with Jensen’s assignment of the material to his three new genera standing unchallenged, until Brian Curtice came on the scene in the mid 1990s. In a series of three publications (two papers, one abstract), he first synonymised Ultrasauros with Supersaurus, then Dystylosaurus also with Supersaurus, and finally (tentatively) Supersaurus itself with Barosarus. If Curtice’s suggestions were all correct, then there were no new sauropods from Jensen’s work in the the Dry Mesa quarry, just a lot of Barosaurus material.

Was he right? We’ll now consider each of the three publications in turn.

First, Ultrasauros. Jensen had always considered this genus to be a brachiosaurid due to the morphology of the scapulocoracoid BYU 9462 — and indeed this element does seem to be brachiosaurid. Unfortunately, he did not found the taxon on this element, but on the dorsal vertebra BYU 9044. Curtice et al. (1996) re-examined this element, and argued convincingly that it was not an anterior dorsal from a brachiosaurid, as Jensen had thought, but a posterior dorsal from a diplodocid. Since its neural spine morphology matches that of the first preserved sacral spine (S2) of the Supersaurus sacrum, and since it was found between the two Supersaurus scapulocoracoids, Curtice et al. (1996:94) considered BYU 9044 to be a vertebra of Supersaurus (belonging to the holotype individual), and therefore concluded that Ultrasauros was a junior subjective synonym of Supersaurus. They inferred that the referred Ultrasauros scapulocoracoid BYU 9462 therefore did not belong to the same species as the type, since it was brachiosaurid, and referred it to Brachiosaurus sp.

We consider all of Curtice et al.’s (1996) arguments well-founded and convincing, and agree with their conclusions. As a result, both spellings of Jensen’s brachiosaurid genus are now discarded: Ultrasaurus as a nomen dubium, and Ultrasauros as a junior synonym.

Curtice et al. (1996:figure 2). “Uncrushed” Supersaurus vivianae caudal dorsal, BYU 9044, right lateral view.

A few years later, Curtice and Stadtman (2001) took aim at Dystylosaurus. Jensen had argued that it was unique because of the paired centroprezygapophyseal laminae that supported each prezygapophysis from below — and it was from this feature than the genus took its name. But Curtice and Stadtman pointed out that this supposedly unique feature is in fact almost ubiquitous in diplodocids. Because it, too, was found between the two Supersaurus scapulae (close to the Ultrasaurus dorsal), Curtice and Stadtman referred it, too, to Supersaurus, thereby collapsing all three of Jensen’s taxa into one. This argument, too, is well supported and has been generally accepted.

Finally, in a sole-authored abstract, Curtice (2003) hedged about whether he considered Supersaurus to be Barosaurus. I will quote directly, as the line of reasoning is vague and difficult to summarise:

The question of is Supersaurus truly a distinct genus from Barosaurus is now testable. The former Dystylosaurus dorsal vertebra provides an autapomorphy for Supersaurus, that being a strongly reduced bifid neural spine on dorsal four. This loss of bifidity is important for in all other diplodocids the neural spine is still deeply bifurcated on dorsal four. Only Barosaurus has a reduction in cleft depth that far forward in the dorsal column. Supersaurus has all but lost the cleft, more closely resembling the sixth dorsal vertebra of Barosaurus than the fourth.

It is disappointing that this abstract never became a more rigorously argued paper, because the conclusion here is highly equivocal. Curtice appears to be saying that Supersaurus is distinct from Barosaurus — but only on the basis of bifidity reducing two vertebrae more anteriorly in Supersaurus. In other words, he seems to be suggesting that the two taxa are indisinguishable aside from this rather minor difference.

At any rate, this speculation in a conference abstract has generally been ignored, and Supersaurus considered a valid and distinct genus.

5. Jimbo the WDC Supersaurus

In 2008, Lovelace et al. (2008, duh) described WDC DMJ-021, a new specimen of Supersaurus vivianae at the Wyoming Dinosaur Center that is known informally as “Jimbo”. (Confusingly, they refer to the Supersaurus holotype scapulocoracoid by yet a third specimen number, BYU 12962; but this is the revised specimen number of the referred scapulocoracoid, not the holotype.)

Lovelace et al. (2008) did not justify in detail their referral of Jimbo to Supersaurus. The closest they come is this brief passage on page 529–530:

While a scapula is not known for WDC DMJ-021, other elements are identical to axial elements referred to the type individual of Supersaurus. Referral of all material is supported by relative position within their respective quarries (Curtice and Stadtman 2001; Lovelace 2006), size of the skeletal elements, and congruence of phylogenetically significant diplodocid characters between the scapula and referred material.

All of this is kind of weaselly. What it amounts to is this: vertebrae are “identical” to those referred to the BYU Supersaurus (but not really, as we’ll see), and the elements are really big, and the Supersaurus holoype scap comes out in about the same place as Jimbo in a phylogenetic analysis if you code them up separately. This is weak sauce, and I would really have liked to see a much more explicit “Jimbo shares synapomorphies X, Y and Z with BYU Supersaurus” section.

Among the ways in which the justification for this assignment disappoints is that the presacrals that are described as “identical” to the BYU elements are not at all well preserved (Lovelace et al. 2008:figures 3D–E, 4A, 5A): in particular C13, presumably the best preserved cervicals as it is the only one illustrated, is missing the condyle, prezygapophyses and neural spine. It’s not possible to be sure in light of the small monochrome illustrations in the paper, but it does not seem likely that these elements can be reliably assessed as identical to the BYU cervical.

Lovelace et al. (2008:figure 3). Lateral views of cervical vertebrae from A, Diplodocus carnegii (Hatcher 1901); B, Barosaurus lentus (Lull 1919); C, Apatosaurus louisae (Gilmore 1936); D and E, Supersaurus vivianae; demonstrating pneumatic modifications of centra. Supersaurus has the least amount of modification with minimal size for pneumatopores. Internal structure is similar to that seen in other diplodocids (Janensch, 1947). Left lateral view of Cv.13 (D, missing the condyle, prezygapophyses and neural spine; length of incomplete centra 94cm). E, cross section through Cv.11, 5cm posterior of the diapophysis.

The big surprise in the Jimbo paper is that in the phylogenetic analysis (Lovelace et al. 2008:figure 14), the compound BYU+WDC Supersaurus is recovered as an apatosaurine, the sister taxon to Apatosaurus, rather than as a diplodocine as had been assumed in previous studies due to its resemblance to the diplodocine Barosaurus.

The huge specimen-level phylogenetic analysis of diplodocoids by Tschopp et al. (2015) corroborated Lovelace et al’s (2008) referral of the WDC specimen to Supersaurus vivianae, as the two species were sister groups in all most parsimonious trees, with quite strong character support (Tschopp et al. 2015:187). But it placed the Supersaurus clade at the base of Diplodocinae, not within Apatosaurinae as Lovelace et al. (2008) had found.

This, then, was the state of play when Matt and I started to work on Supersaurus during the 2016 Sauropocalypse: Ultrasauros and Dystylosaurus had both been sunk into Supersaurus, and the WDC specimen had been referred to the same species.

Next time, we’ll look what Matt and I found in Utah, and what we think it means for Supersaurus and its friends.

 

References

  • Curtice, Brian D. 2003. Two genera down, one to go? The potential synonomy [sic] of Supersaurus with Barosaurus. Southwest Paleontological Symposium 2003, Guide to Presentations. Mesa Southwest Museum, January 25 2003. Unpaginated.
  • Curtice, Brian D. and Kenneth L. Stadtman. 2001. The demise of Dystylosaurus edwini and a revision of Supersaurus vivianae. Western Association of Vertebrate Paleontologists and Mesa Southwest Museum and Southwest Paleontologists Symposium, Bulletin 8:33-40.
  • Curtice, Brian D., Kenneth L. Stadtman and Linda J. Curtice. 1996. A reassessment of Ultrasauros macintoshi (Jensen, 1985). M. Morales (ed.), “The continental Jurassic”. Museum of Northern Arizona Bulletin 60:87–95.
  • George, Jean. 1973a. Supersaurus, giant of the giants. Denver Post,
    Empire Magazine. May 13, 1973, pp 14ff.
  • George, Jean. 1973b. Supersaurus, the biggest brute ever. Reader’s Digest (June 1973):51–56.
  • Glut, Donald F. 1997. Dinosaurs: the Encyclopedia. McFarland & Company Inc., Jefferson. 1076 pp.
  • Jensen, James A. 1985. Three new sauropod dinosaurs from the Upper Jurassic of Colorado. Great Basin Naturalist 45(4):697–709.
  • Jensen, James A. 1987. New brachiosaur material from the Late Jurassic of Utah and Colorado. Great Basin Naturalist 47(4):592–608.
  • Kim, Haang Mook. 1983. Cretaceous dinosaurs from South Korea. Journal of the Geological Society of Korea 19(3):115–126.
  • Lee, Yuong-Nam., S. Y. Yang and E. J. Park. 1997. Sauropod dinosaur remains from the Gyeongsang Supergroup, Korea; pp. 103–114 in S. Y. Yang, M. Huh, Y.-N. Lee and M. G. Lockley (eds.), International Dinosaur Symposium for Uhangri Dinosaur Center and Theme Park in Korea. Journal of Paleontological Society of Korea, Special Publication 2.
  • Lovelace, David M., Scott A. Hartman and William R. Wahl. 2008. Morphology of a specimen of Supersaurus (Dinosauria, Sauropoda) from the Morrison Formation of Wyoming, and a re-evaluation of diplodocid phylogeny. Arquivos do Museu Nacional, Rio de Janeiro 65(4):527–544.
  • Olshevsky, George. 1991. A revision of the parainfraclass Archosauria Cope, 1869, excluding the advanced Crocodylia. Mesozoic Meanderings 2:1–196.
  • Tschopp, Emanuel, Octávio Mateus and Roger B. J. Benson. 2015. A specimen-level phylogenetic analysis and taxonomic revision of Diplodocidae (Dinosauria, Sauropoda). PeerJ 2:e857. doi:10.7717/peerj.857

 

Illustration talk slide 19

Illustration talk slide 20

Illustration talk slide 21

Illustration talk slide 22

This whole section, including the title, is mostly swiped from Mike’s Tutorial 17.

Other posts in this series are here.

Papers referenced in these slides:

Caudal pneumaticity in saltasaurines. Cerda et al. (2012: fig. 1).

Earlier this month I was amazed to see the new paper by Cerda et al. (2012), “Extreme postcranial pneumaticity in sauropod dinosaurs from South America.” The title is dramatic, but the paper delivers the promised extremeness in spades. Almost every figure in the paper is a gobsmacker, starting with Figure 1, which shows pneumatic foramina and cavities in the middle and even distal caudals of Rocasaurus, Neuquensaurus, and Saltasaurus. This is most welcome. Since the 1990s there have been reports of saltasaurs with “spongy bone” in their tail vertebrae, but it hasn’t been clear until now whether that “spongy bone” meant pneumatic air cells or just normal marrow-filled trabecular bone. The answer is air cells, loads of ’em, way farther down the tail than I expected.

Caudal pneumaticity in diplodocines. Top, transverse cross-section through an anterior caudal of Tornieria, from Janensch (1947: fig. 9). Bottom, caudals of Diplodocus, from Osborn (1899: fig. 13).

Here’s why this is awesome. Lateral fossae occur in the proximal caudals of lots of neosauropods, maybe most, but only a few taxa go in for really invasive caudal pneumaticity with big internal chambers. In fact, the only other sauropod clade with such extensive pneumaticity so far down the tail are the diplodocines, including Diplodocus, Barosaurus, and Tornieria. But they do things differently, with BIG, “pleurocoel”-type foramina on the lateral surfaces of the centra, leading to BIG–but simple–camerae inside, and vertebral cross-sections that look like I-beams. In contrast, the saltasaurines have numerous small foramina on the centrum and neural arch that lead to complexes of small pneumatic camellae, giving their vertebrae honeycomb cross-sections. So caudal pneumaticity in diplodocines and saltsaurines is convergent in its presence and extent but clade-specific in its development. Pneumaticity doesn’t get much cooler than that.

Pneumatic ilia in saltasaurines. Cerda et al. (2012: fig. 3).

But it does get a little cooler. Because the stuff in the rest of the paper is even more mind-blowing. Cerda et al. (2012) go on to describe and illustrate–compellingly, with photos–pneumatic cavities in the ilia, scapulae, and coracoids of saltasaurines. And, crucially, these cavities are connected to the outside by pneumatic foramina. This is important. Chambers have been reported in the ilia of several sauropods, mostly somphospondyls but also in the diplodocoid Amazonsaurus. But it hasn’t been clear until now whether those chambers connected to the outside. No patent foramen, no pneumaticity. It seemed unlikely that these sauropods had big marrow-filled vacuities in their ilia–as far as I know, all of the non-pneumatic ilia out there in Tetrapoda are filled with trabecular bone, and big open marrow spaces only occur in the long bones of the limbs. And, as I noted in my 2009 paper, the phylogenetic distribution of iliac chambers is consistent with pneumaticity, in that the chambers are only found in those sauropods that already have sacral pneumaticity (showing that pneumatic diverticula were already loose in their rear ends). But it’s nice to have confirmation.

So, the pneumatic ilia in Rocasaurus, Neuquensaurus, and Saltasaurus are cool because they suggest that all the other big chambers in sauropod ilia were pneumatic as well. And for those of you keeping score at home, that’s another parallel acquisition in Diplodocoidea and Somphospondyli (given the apparent absence of iliac chambers in Camarasaurus and the brachiosaurids, although maybe we should bust open a few brachiosaur ilia just to be sure*).

* I kid, I kid.**

** Seriously, though, if you “drop” one and find some chambers, call me!

Pectoral pneumaticity in saltasaurines. Cerda et al. (2012: fig. 2).

But that’s not all. The possibility of pneumatic ilia has been floating around for a while now, and most of us who were aware of the iliac chambers in sauropods probably assumed that eventually someone would find the specimens that would show that they were pneumatic. At least, that was my assumption, and as far as I know no-one ever floated an alternative hypothesis to explain the chambers. But I certainly did not expect pneumaticity in the shoulder girdle. And yet there they are: chambers with associated foramina in the scap and coracoid of Saltasaurus and in the coracoid of Neuquensaurus. Wacky. And extremely important, because this is the first evidence that sauropods had clavicular air sacs like those of theropods and pterosaurs. So either all three clades evolved a shedload of air sacs independently, or the basic layout of the avian respiratory system was already present in the ancestral ornithodiran. I know where I’d put my money.

There’s loads more interesting stuff to talk about, like the fact that the ultra-pneumatic saltasaurines are among the smallest sauropods, or the way that fossae and camerae are evolutionary antecedent to camellae in the vertebrae of sauropods, so maybe we should start looking for fossae and camerae in the girdle bones of other sauropods, or further macroevolutionary parallels in the evolution of pneumaticity in pterosaurs, sauropods, and theropods. Each one of those things could be a blog post or maybe a whole dissertation. But my mind is already thoroughly blown. I’m going to go lie down for a while. Congratulations to Cerda et al. on what is probably the most important paper ever written on sauropod pneumaticity.

References

  • Cerda, I.A., Salgado, L., and Powell, J.E. 2012. Extreme postcranial pneumaticity in sauropod dinosaurs from South America. Palaeontologische Zeitschrift. DOI 10.1007/s12542-012-0140-6
  • Janensch, W. 1947. Pneumatizitat bei Wirbeln von Sauropoden und anderen Saurischien. Palaeontographica, Supplement 7, 3:1–25.
  • Osborn, H. F. 1899. A skeleton of Diplodocus. Memoirs of the American Museum of Natural History 1:191–214.