Just to wash our mouths out after all the theropod-related unpleasantness yesterday:

What we’re seeing here, in glorious 3D, is the 7th cervical vertebrae of BYU 1252-18531. This is an apatosaurine at the Brigham Young University Museum of Paleontology which the museum has catalogued as “Apatosaurus excelsus” (i.e. Brontosaurus excelsus), and which Tschopp et al. (2015) tentatively referred to Brontosaurus parvus, but which I suspect is most likely good old Apatosaurus louisae.

It’s in the rarely seen ventral view, which really emphasizes the ludicrously over-engineered cervical ribs. Get your 3D glasses on and marvel at how they come lunging out of the screen at you, like giant insects in a 1950s B-movie.

So beautiful.

What if I told you that when Matt was in BYU collections a while ago, he stumbled across a cervical vertebra — one labelled DM/90 CVR 3+4, say — that looked like this in anterior view?

I think you would say something like “That looks like a Camarasaurus cervical, resembling as it does those illustrated in the beautiful plates of Osborn and Mook (1921)”. And then you might show me, for example, the left half of Plate LXII:

And then you might think to yourself that, within its fleshy envelope, this vertebra might have looked a bit like this, in a roughly circular neck:

Reasonable enough, right?

But when what if I then told you that in fact the vertebra was twice this wide relative to its height, and looked like this?

I’m guessing you might say “I don’t believe this is real. You must have produced it by stretching the real photo”. To which I would reply “No no, hypothetical interlocutor, the opposite is the case! I squashed the real photo — this one — to produce the more credible-seeming one at the top of the post”.

You would then demand to see proper photographic evidence, and I would respond by posting these three images (which Matt supplied from his 2019 BYU visit):

BYU specimen DM/90 CVR 3+4, cervical vertebra of ?Camarasaurus in anterior view. This is the photo from which the illustration above was extracted.

The same specimen in anteroventral view.

The same specimen in something approaching ventral view.

So what’s going on here? My first thought was that this speicmen has to have been dorsoventrally crushed — that this can’t be the true shape.

And yet … counterpoint: the processes don’t look crushed: check out the really nice 3d preservation of the neural spine metapophyses, the prezygs, the transverse processes, the nice, rounded parapophyseal rami, and even the ventral aspect of the centrum. This vertebra is actually in pretty good condition.

So is this real? Is this the vertebra more or less as it was in life? And if so, does that mean that the flesh envelope looked like this?

Look, I’m not saying it isn’t ridiculous; I’m just saying this seems to be more or less where the evidence is pointing. We’ve made a big deal about how the necks of apatosaurines were more or less triangular in cross-section, rather than round as has often been assumed; perhaps we need to start thinking about whether some camarasaur necks were squashed ovals in cross section?

Part of what’s crazy here is that this makes no mechanical sense. A cantilevered structure, such as a sauropod neck, needs to be tall rather than wide in order to attain good mechanical advantage that can take the stress imposed by the neck’s weight. A broad neck is silly: it adds mass that needs to be carried without providing high anchors for the tension members. Yet this is what we see. Evolution doesn’t always do what we would expect it to do — and it goes off the rails when sexual selection comes into play. Maybe female camarasaurus were just really into wide-necked males?

Final note: I have been playing fast and loose with the genus name Camarasaurus and the broader, vaguer term camarasaur. Matt and I have long felt (without having made any real attempt to justify this feeling) that Camarasaurus is way over-lumped, and probably contains multiple rather different animals. Maybe there is a flat-necked species in among them?

(Or maybe it’s just crushing.)

In the last post, we looked at some sauropod vertebrae exposed in cross-section at our field sites in the Salt Wash member of the Morrison Formation. This time, we’re going to do it again! Let’s look at another of my faves from the field, with Thuat Tran’s hand for scale. And, er, a scale bar for scale:

And let’s pull the interesting bits out of the background:

Now, confession time. When I first saw this specimen, I interpeted it as-is, right-side up. The round thing in the middle with the honeycomb of internal spaces is obviously the condyle of a vertebra, and the bits sticking out above and below on the sides frame a cervical rib loop. I figured the rounded bit at the upper right was the ramus of bone heading for the prezyg, curved over as I’ve seen it in some taxa, including the YPM Barosaurus. And the two bits below the centrum would then be the cervical ribs. And with such big cervical rib loops and massive, low-hanging cervical ribs, it had to an apatosaurine, either Apatosaurus or Brontosaurus.

Then I got my own personal Cope-getting-Elasmosaurus-backwards moment, courtesy of my friend and fellow field adventurer, Brian Engh, who proposed this:

Gotta say, this makes a lot more sense. For one, the cervical ribs would be lateral to the prezygs, just as in, oh, pretty much all sauropods. And the oddly flat inward-tilted surfaces on what are now the more dorsal bones makes sense: they’re either prezyg facets, or the flat parts of the rami right behind the prezyg facets. The missing thing on what is now the right even makes sense: it’s the other cervical rib, still buried in a projecting bit of sandstone. That made no sense with the vert the other way ’round, because prezygs always stick out farther in front than do the cervical ribs. And we know that we’re looking at the vert from the front, otherwise the backwards-projecting cervical rib would be sticking through that lump of sandstone, coming out of the plane of the photo toward us.

Here’s what I now think is going on:

I’m still convinced that the bits of bone on what is now the left side of the image are framing a cervical rib loop. And as we discussed in the last post, the only Morrison sauropods with such widely-set cervical ribs are Camarasaurus and the apatosaurines. So what makes this an apatosaurine rather than a camarasaur? I find several persuasive clues:

  • If we have this thing the right way up, those prezygs are waaay up above the condyle, at a proportional distance I’ve only seen in diplodocids. See, for example, this famous cervical from CM 3018, the holotype of A. louisae (link).
  • The complexity of the pneumatic honeycombing inside the condyle is a much better fit for an apatosaurine than for Camarasaurus–I’ve never seen that level of complexity in a camarasaur vert.
  • The bump on what we’re now interpreting as the cervical rib looks suspiciously like one of the ventrolateral processes that Kent Sanders and I identified in apatosaurine cervicals back in our 2002 paper. I’ve never seen them, or seen them reported, in Camarasaurus–and I’ve been looking.
  • Crucially, the zygs are not set very far forward of the cervical ribs. By some rare chance, this is pretty darned close to a pure transverse cut, and the prezygs, condyle (at its posterior extent, anyway), and the one visible cervical rib are all in roughly the same plane. In Camarasaurus, the zygs strongly overhang the front end of the centrum in the cervicals (see this and this).

But wait–aren’t the cervical ribs awfully high for this to be an apatosaurine? We-ell, not necessarily. This isn’t a very big vert; max centrum width here is 175mm, only about a third the diameter of a mid-cervical from something like CM 3018. So possibly this is from the front of the neck, around the C3 or C4 position, where the cervical ribs are wide but not yet very deep. You can see something similar in this C2-C5 series on display at BYU:

Or, maybe it’s just one of the weird apatosaurine verts that has cervical rib loops that are wide, but not very deep. Check out this lumpen atrocity at Dinosaur Journey–and more importantly, the apatosaur cervical he’s freaking out over:

UPDATE just a few minutes later: Mike reminded me in the comments about the Tokyo apatosaurine, NSMT-PV 20375, which has wide-but-not-deep cervical ribs. In fact, C7 (the vertebra on the right in this figure) is a pretty good match for the Salt Wash specimen:

UpchurchEtAl2005-apatosaurus-plate2-C3-6-7

NSMT-PV 20375, cervical vertebrae 3, 6 and 7 in anterior and posterior views. Modified from Upchurch et al. (2005: plate 2).

UPDATE the 2nd: After looking at it for a few minutes, I decided that C7 of the Tokyo apatosaurine was such a good match for the Salt Wash specimen that I wanted to know what it would look like if it was similarly sectioned by erosion. In the Salt Wash specimen, the prezygs are sticking out a little farther than the condyle and cervical rib sections. The red line in this figure is my best attempt at mimicking that erosional surface on the Tokyo C7, and the black outlines on the right are my best guess as to what would be exposed by such a cut (or pair of cuts). I’ve never seen NSMT-PV 20375 in person, so this is just an estimate, but I don’t think it can be too inaccurate, and it is a pretty good match for the Salt Wash specimen.

Another way to put it: if this is an apatosaurine, everything fits. Even the wide-but-not-low-hanging cervical ribs are reasonable in light of some other apatosaurines. If we think this is Camarasaurus just because the cervical ribs aren’t low-hanging, then the pneumatic complexity, the height of the prezygs, and the ventrolateral process on the cervical rib are all anomalous. The balance of the evidence says that this is an apatosaurine, either a small, anterior vert from a big one, or possibly something farther back from a small one. And that’s pretty satisfying.

One more thing: can we take a moment to stand in awe of this freaking thumb-sized cobble that presumably got inside the vertebra through one its pneumatic foramina and rattled around until it got up inside the condyle? Where, I’ll note, the internal structure looks pretty intact despite being filled with just, like, gravel. As someone who spends an inordinate amount of time thinking about how pneumatic vertebrae get buried and fossilized, I am blown away by this. Gaze upon its majesty, people!

This is another “Road to Jurassic Reimagined, Part 2″ post. As before, Part 1 is here, Part 2 will be going up here in the near future. As always, stay tuned.

References

 

I had an interesting opportunity when I was in Utah and Colorado a couple of weeks ago. At Dinosaur Journey in Fruita, Colorado, I went looking for a cast of the Potter Creek Brachiosaurus humerus. I found it — more on that another time — and I also found a cast of BYU 4503, the holotype dorsal vertebra of Dystylosaurus (now almost universally regarded as Supersaurus [but then…]), lurking with it in a corner of the collections room.

Dystylosaurus cast, posterior view.

Somehow I had overlooked the Dystylosaurus cast on all of my previous visits to DJ, which is a shame, because the cast is easy to pick up, flip over, and manipulate. Very much unlike the actual fossil, which combines the charming attributes, shared with many other sauropod vertebrae, of weighing hundreds of pounds but still being awfully fragile.

Dystylosaurus cast, anterior view.

So, hey ya, I had a chance to photograph and measure both sides of the vertebra. You’re not supposed to take measurements from casts, but I figured what the heck, no-one was going to lock me up for it, and I could compare the measurements from the cast to the measurements of the real thing when I visited BYU later in the trip. And that’s exactly what I did. It was easy to make sure I took the second set of measurements the same way I had done the first set, because I took them just a few days apart.

The real deal at BYU.

Here’s what I got. For each measurement, the actual value measured from the real fossil at BYU comes first, followed by the same measurement from the cast at Dinosaur Journey, followed by the difference as a percentage of the first (true) measurement.

  • Total Height (as preserved): 1050mm / 1022mm / -2.6%
  • Max Width (as preserved): 905mm / 889mm / -1.8%
  • Anterior Centrum Height: 400mm / 394mm / -1.5%
  • Anterior Centrum Width: 470mm / 454mm / -3.4%
  • Posterior Centrum Height: 365mm / 352mm / -3.5%
  • Posterior Centrum Width: 480mm / 473mm / -1.5%

They’re not the same! On average, the measurements of the cast are 2.4% smaller than the same measurements taken from the actual bone. (Incidentally, you may be wondering how I measured the posterior centrum faces of the BYU vertebra without flipping it. I used a couple of wooden blocks as orthogonators and measured between them, and I did it at several points to make sure they were truly parallel. In essence, I made giant redneck calipers, a method that Mike and I have had to employ many times when measuring huge, weirdly-shaped fossils. Remind me to show you John Foster’s giant caliper setup sometime.)

Dinosaur Journey cast in right lateral view, big doofus for scale.

Anyway, the discrepancy in the measurements should not be surprising. It is a known phenomenon that when an object is molded and cast, there is a little bit of shrinkage. You can see it bedevil Adam Savage in his quest for the ultimate Maltese Falcon replica in this charming video:

So, on one hand, no outright disasters here; all of the cast measurements are within a few percent of the real measurements, so if all you had was a cast, you could get a pretty good sense of the size of the real thing. But precision counts, even among giant sauropods. In a world where the largest vertebra of Argentinosaurus is only 1cm bigger in diameter than the largest vertebra of Patagotitan, differences like I got with Dystylosaurus would be enough to scramble the order of giant vertebrae. So if you’re ever stuck measuring something from a cast, be forthright and say as much, so that no-one mistakes the cast measurements for the real thing.

Here are some more measurements from BYU 4503, the real thing, for you completists. Note that the vertebra is sheared a bit from right postero-ventral to left antero-dorsal, so figuring out how to take the centrum length is not straightforward. I ended up doing it twice, once orthogonal to the posterior centrum face, and once following the slant of the centrum, both at the mid-height of the centrum, as shown in the little diagram from my notebook (above).

  • Centrum Length, left side, orthogonal: 295mm
  • Centrum Length, left side, on the slant: 310mm
  • Centrum Length, right side, orthogonal: 280mm
  • Centrum Length, right side, on the slant: 305mm
  • Max Width across prezygs: 305mm
  • Min gap between prezygs: 19mm
  • Max Width across parapophyses: 620mm
  • Max antero-posterior length of prezyg articular surfaces: 55mm
  • Max antero-posterior depth of hypantrum: 95mm
  • Max antero-posterior depth of fossa between spino-prezyg laminae (SPRLs): 80mm
  • Neural spine cavity, max antero-posterior extent: 40mm
  • Neural spine cavity, max medio-lateral extent: 70mm

Finally, a huge thanks to Julia McHugh at Dinosaur Journey and Brooks Britt and Rod Scheetz at BYU for letting me come play with their huge toys er, hugely important scientific specimens. Rod was particularly helpful, shifting giant things about with a forklift, helping me measure bones that are longer than I am tall, and boxing up loan specimens for me. Mike and I have had really good luck with pro-science curators and collections managers, but the folks at DJ and BYU have always been standouts, and I can’t thank them enough.

Back into the Corner of Shame, artificially tiny Dystylosaurus!

Nothing too serious here, just a fun shot I got while in the collections at BYU this past week. The Brachiosaurus element is metacarpal 1 (thumb column) from BYU 4744, the Potter Creek material. I highlighted my own metacarpal 3. There is a metacarpal 3 from this specimen, but I didn’t see it on the shelf. According to D’Emic and Carrano (2019), the MC3 is 60cm long, vs 57cm for this MC1. So this photo could have been 3cm more impressive!

Oh, ignore the tag on the left that says “radius”. You could be forgiven for thinking that the bone I have my hand on is a radius, but the radius from this individual is 1.34 meters long, or about two-and-a-third times the length of this metacarpal.

Reference

D’Emic, M.D. and Carrano, M.T., 2019. Redescription of Brachiosaurid Sauropod Dinosaur Material From the Upper Jurassic Morrison Formation, Colorado, USA. The Anatomical Record.

I keep wishing there was a single place out there where I could look up Jensen’s old BYU specimen numbers for Supersaurus, Ultrasaurus and Dystylosaurus elements, and find the modern equivalents, or vice versa. Then I realised there’s no reason not to just make one. So here goes! The first column shows the specimen numbers as used in Jensen (1985), and last column contains Jensen’s own assignments except where noted.


Jensen Element New Notes
5000 posterior dorsal vertebra 9044 holotype of Ultrasauros
5001 scapulocoracoid 9462 referred to Ultrasauros
5002 anterior caudal vertebra 9045 referred initially to Ultrasauros, then Supersaurus [1]
5003 mid-cervical vertebra 9024 referred initially to Ultrasauros, then Supersaurus [2]
5500 left scapulocoracoid [3] 9025 holotype of Supersaurus
5501 right scapulocoracoid [3] 12962 referred to Supersaurus, although found first
5502 sequence of 12 caudals [4] 9084 referred to Supersaurus
5503 right ischium [4] 12946 referred to Supersaurus
5504 two mid-caudal vertebrae [4] ?9077[5] referred to Supersaurus
5750 anterior dorsal vertebra 4503 holotype of Dystylosaurus

By the way, does anyone know why the numbers were changed?

 

Notes

[1] This diplodocid caudal, which is obviously diplodocid based on Jensen’s own illustrations (Jensen 1985:figures 2D,E, 3E), was reassigned to Supersaurus by Curtice (1995).

[2] Jensen (1987:602-603) recognised his own error in referring this cervical to the brachiosaurid taxon Ultrasaurus, based on its bifurcated neural spine. He “provisionally refer[red] it to the Diplodocidae” in the text, but without specifying a genus or species. However in caption to illustrations in the same paper (Jensen 1987:figures 7A, B, 8C) he names the element as Supersaurus vivianae without comment.

[3] Jensen’s (1985) original description describes BYU 5500 (=BYU 9025) as a right scapulocoracoid, implying that BYU 5501 is the left; but this is incorrect.

[4] Jensen’s original Supersaurus/Ultrasaurus/Dystylosaurus description is confusing and contradictory in his assignment of specimen numbers. In his systematic palaeontology section, Jensen (1985:701) says that BYU 5502 is the ischium, BYU 5503 is the pair of mid-caudals and BYU 5504 is the sequence of 12 caudals. But the description on the same page contradicts this, giving the assignments shown here. The casting vote goes to the caption of Jensen (1985:figure 7), in which part A illustrates BYU 5503, the ischium; and parts C, D and D1 illustrate caudals that do not appear to be part of sequence of twelve.

[5] Curtice et al. (2001:36) say “An additional caudal vertebra (BYU 9077) is referred to (and figured as) Supersaurus in the text of Jensen (1985)”. This probably refers to Jensen 1985:figure 7:C, D, D1, which are captioned as follows: “C, BYU 5033, Supersaurus vivianae, referred specimen, ischium [sic]. D, D1, BYU 5504, Supersaurus vivianae, referred specimen, caudal vertebra.” Since part C of the figure is clearly a caudal vertebra, and since BYU 5503 is also illustrated as an ischium in part A of the same figure(!), it seems most likely that the caudals in part C and parts D and D1 of this figure are the pair described as BYU 5044 on pages 701-704.

 

Commentary (i.e. pointless whining)

For all his innovations in skeletal mounting and his amazing discoveries in the field, Jensen was evidently a markedly careless palaeontologist in many respects, and his contempt for specimen numbers in particular has created enormous problems. Even within a single page — even within a single figure caption — he was capable of contradicting himself on the numbers assigned to specimens. Most illustrations don’t give specimen numbers at all. And while in many respects the later work of Curtice et al. (1996) and Curtice and Stadtman (2001) is much better, they did the world no favours by simply switching to the new specimen numbers without providing a definitive key like the one I am trying to build here. It’s pretty silly that, 23 years on, we are reduced to guesswork like note 5.

 

References

  • Curtice, Brian D. 1995. A description of the anterior caudal vertebrae of Supersaurus vivianae. Journal of Vertebrate Paleontology 15(3):25A.
  • 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.
  • 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.

One of the strange things about Jensen’s 1985 paper is that the abstract implies that he informally considered the Ultrasauros scapulocoracoid to be the type specimen.

Cast of BYU 9462, scapulocoracoid referred to Ultrasaurus macintoshi (possibly intended to the be the holotype), at Brigham Young Museum. This photo is one of a series in which I turned the cast in place to obtain photos for a photogrammetric model.

Here’s what Jensen (1985:697) says:

From 1972 to 1982 three exceptionally large sauropod scapulocoracoids […] were collected from the base of the Brushy Basin Member of the Upper Jurassic, Morrison Formation, in western Colorado. Two of the scapulae are conspecific, but the third represents a second genus and possibly a new family. The two conspecific specimens are described here as Supersaurus vivianae; the second genus is described as Ultrasaurus mcintoshi.

But on page 704, he formally and unambiguously nominated the dorsal vertebra as the holotype:

Family Brachiosauridae
Ultrasaurus macintoshi, n. gen., n. sp.
[…]
Holotype.—BYU 5000, posterior dorsal vertebra.
Referred material.—BYU 5001, scapulocoracoid.

Stranger still, two years after this, Jensen (1987:603) straight up claimed – quite incorectly — that the scap was the Ultrasaurus holotype:

In 1979 a scapulocoracoid, 2.70 m (8’10”) long (Figs. 6A-B, 9I) was collected in the Dry Mesa Quarry. This scapula, BYU 5000 [sic; he meant BYU 5001], is readily referrable to the Brachiosauridae (Fig. 9H) and is the holotype of Ultrasaurus macintoshi Jensen, 1985.

But it sayin’ it’s so don’t make it so. The joint evidence of the 1985 abstract and the 1987 extract suggest that Jensen probably intended the scap to be the holotype and somehow accidentally designated the wrong element — or was persuaded to do so against his own judgement. But however it came about, the scap is not the holotype.

BYU 9462, the scapulocoracoid referred by Jensen to Ultrasauros. Mike Taylor for scale, doing a Jensen. Note that the actual specimen is very much a mosaic of bone fragments, rather than the solid, complete bone that the cast might suggest.

Instead, the holotype remains the large posterior dorsal vertebra BYU 9044 (BYU 5000 of Jensen’s usage) which Curtice et al. (1996) convincingly showed to be diplodocid, and referred to Supersaurus, making Ultrasaurus (and its subsequent replacement Ultrasauros) a junior synonym of that name.

Ultrasauros macintoshi holotype dorsal vertebra BYU 9044, in left lateral view, photographed at the North American Museum of Natural Life. Sorry about all the reflections off the glass case.

But wait, wait. We’ve shown that there are probably two big diplodocids in the Dry Mesa quarry: Barosaurus (represented by the big cervical BYU 9024) and something different (represented by the “Dystylosaurus” dorsal, BYU 4503). The Ultrasauros holotype vertebra probably belongs to one of these (unless there are three big diplodocids in there but we’ll ignore that possibility). But we can’t tell whether the Ultrasauros dorsal belongs with the Barosaurus cervical or the Dystylosaurus dorsal.

All of this means that Ultrasauros is a synonym, but we don’t know of what. It might be Barosaurus; it might be Supersaurus, whatever that is, if it’s not a nomen dubium; and it might be Dystylosaurus, if Supersaurus is a nomen dubium. Yikes.

Well, then. Is it Barosaurus? Here are the dorsal vertebrae of the fairly complete AMNH specimen, in a composite that I put together a few years ago from McIntosh’s (2005) illustrations:

Barosaurus lentus AMNH 6341 dorsal vertebrae 1 to 9 in anterior, left lateral and posterior views. Modified from McIntosh (2005:figure 2.5)

We can compare these with the photo above of the Ultrasauros dorsal in left lateral view, and with this one in posterior view:

Ultrasauros macintoshi holotype dorsal vertebra BYU 9044, in posterior view, photographed at the North American Museum of Natural Life. Sorry about all the reflections off the glass case.

I wouldn’t want to hang too much on those poor quality, postage-stamp-sized monochrome photos of the Barosaurus dorsals. And I’m also more than aware of the imperfections in my photos of the “Ultrasauros” dorsal. But to the naked eye, there’s nothing here that immediately screams they couldn’t be the same thing.

Lull’s (1919) monograph on the original Barosaurus specimen YPM 429 also illustrated a posterior dorsal, which he designated D9. Lull helpfully provided both drawings and photographs:

Lull (1919: plate IV: parts 4-6). Barosaurus lentus holoype YPM 429, 9th dorsal vertebra in anterior, right lateral and posterior views (line drawing).

Lull (1919: plate IV: parts 4-6). Barosaurus lentus holoype YPM 429, 9th dorsal vertebra in anterior, right lateral and posterior views (photographs).

With something a bit more substantial to go on, the case for the Ultrasaurus vertebra being Barosarus doesn’t look so good.

Most obviously, its centrum is much longer than that of the Barosaurus dorsal — and indeed, than any posterior dorsal vertebra of any diplodocid. This character is the reason — the only reason — that Jensen (1985:704) initially thought it was brachiosaurid: “Ultrasaurus shares the family characteristic of a long dorsal centrum with Brachiosaurus, but in other features it has no parallel with that genus”. Curtice et al. (1996:90) argued that “extensive transverse and oblique crushing artificially elongate the centrum […]. Without the crushing […] the centrum shrinks considerably in length”. Based on my photos, I can’t really see any justification for this claim, but Curtice spent waaay more time with this specimen than I have done, so I’m going to hold that observation lightly.

But there are other features of BYU 9044 that are not a good match for Lull’s illustrations. These include a less robust looking and more prominently laminated subzygapophyseal neural arch, and a neural spine that is anteroposteriorly broader but transversely narrower than in Lull’s specimen. Also, the apex of the neural spine in anterior or posterior view is convex in BYU 9044 but concave in YPM 429.

None of these characters can be considered to definitely separate BYU 9044 from Barosaurus, especially in light of that element’s crushing, the imperfect preservation of Lull’s specimen, the possibility of serial variation, and the fact that I am working only from photographs and drawings of both. But when you put all the differences together, they combine to at least suggest that Ultrasaurus is not Barosaurus — and that it is therefore most likely Supersaurus/Dystylosaurus.

So what about the scapulocoracoid?

It looks brachiosaurid, as Jensen observed. Curtice et al. (1996) concurred, and referred it to Brachiosaurus sp. In fact, when compared with the best-preserved scapula of a known brachiosaurid Giraffatitan HMN Sa 9), it’s not all that similar:

Brachiosaurid scapulocoracoids. Left: cast of BYU 9462, right scapulocoracoid referred to Ultrasauros macintoshi, at Brigham Young Museum, with Mike Taylor for scale. Right: HMN Sa 9, left scapula only (coracoid is not co-ossified) of Giraffatitan brancai, scaled to same blade length as BYU 9462, photo by FunkMonk (Michael B. H.), CC By-SA.

It’s apparent, when looking at the two scaps together, that there are significant differences: BYU 9462 is in every respect less robust, having a less expanded distal blade, a more constricted midshaft, a less promiment and narrower acromial ridge and a much less robust ventral ridge. In addition, the acromion process is hooked in Sa 9, so that its tip projects laterally, whereas it is rounded in BYU 9462. Finally, the shapes of the distal blades differ, having a gently rounded profile in BYU 9462 but a distinct kink in Sa 9 where the dorsal part of the margin inclines anterodorsally.

What does all this mean? We don’t know. I’m certainly not arguing that BYU 9462 is not brachiosaurid, as it does seem to differ less from Giraffatitan scapulae than from those of other sauropods. All I’m saying is that it’s not all that Giraffatitan-like. But then every bone that we know from both Giraffatitan and Brachiosaurus is significantly different between them (Taylor 2009:798), so if a subsequently discovered associated skeleton one day shows us that this is just what the scapulocoracoid of Brachiosaurus altithorax looks like, it would not be a huge shock.

Still, as things stand, I’m not really convinced that the referral to Brachiosaurus sp. — based on a not-particularly-close resemblance to a completely different brachiosaurid — is rock solid. Had the scap been the type specimen, as Jensen probably intended, I would consider that the sound move would be to continue to consider Ultrasauros as a distinct taxon from Brachiosaurus, unless and until an associated specimen demonstrates that synonymy is warranted.

But that’s all in Shoulda-Coulda-Woulda territory. In fact the scapulocoracoid is not the type specimen, and so the name Ultrasauros remains sunk, even though we can’t tell whether it’s a synonym of Barosaurus, Supersaurus or Dystylosaurus. That will remain the case unless someone takes the initiative to raise a new name for the scapulocoracoid — which we can, at least, be confident does not belong the diplodocid Ultrasauros. I think that would be a reasonable move for someone to make, but it’s not one that I feel moved to make myself.

… and with that, I think we have finally reached the end of this series. We may revisit it in the future to say more about Jimbo, or maybe Dinheirosaurus, but this series has been the substance of what we have to say. Hope you’ve enjoyed it!

References

 

 

Poor Dystylosaurus. Always the bridesmaid. No-one seems to care much about it, yet the one and only vertebra that bears that name is the single most diagnostic elements out of all the individual bones that have been assigned to Supersaurus over the years.

A nice drawing of the “Dystylosaurus” dorsal vertebra in anterior and right lateral views. It’s probably Tracey Ford’s work (awaiting confirmation), from the PaleoFile page on Supersaurus.

Unfortunately, we weren’t able to learn a whole ton about this vertebra on the Sauropocalypse visit. We did see it, but it was flat on its back on a shelf not much taller than the anteroposterior length of the bone itself, so we weren’t able to get a good look at it in anything but dorsal and ventral views. If we’d had more time to get things arranged, I’m sure the BYU people would have been happy to get it down from the shelf for us, but we simply had so much to do in their collections that time was never made for it.

BYU 4503, the holotype and only element of Dystylosaurus edwini, an anterior dorsal vertebra. here seen in approximately dorsal view with anterior to the top. Matt Wedel for scale.

Matt actually got some rather better photos a few years ago, though (based on his comment on that post), there are probably no more than the couple in that old blog-post. (By the way, notice how very different the colour of the bone appears in Matt’s old photos from how it appears in my more recent one above.)

Why do I say so confidently that the Dystylosaurus vertebra is diagnosable? Because it has a whole suite of characters that tell us it’s an anterior dorsal vertebra from a diplodocid (dual centroprezygapophyseal laminae, anteroposteriorly compressed spine composed primarily of spinozygapophyseal rather than spinodiapophyseal laminae, drooping transverse processes), yet two features of the spine are never seen in such vertebrae: the spine is wholly unsplit without even a hint of bifurcation, even featuring macronarian-like lateral apices; and it’s hollow inside rather than being constructed from intersecting plates of bone. (You can see the internal hollow in the photo above.)

So what happens to its genus name given the doubts about Supersaurus‘s diagnosability? The general trend of comments on these posts has been that Supersaurus should stand or fall on its holotype, and I am inclined to agree that parachuting in the Dystylosaurus vertebra or Jimbo as a neotype to save the name would be a mistake. For one thing, despite its numerous appearances in kids’ books, the name Supersaurus is not that important in the technical literature: for example, no-one has named a clade Supersaurinae or similar. For another, the holotypic scapulocoracoid BYU 9025 is only questionably undiagnosable. There would always be the possibility that if someone nominated a neotype and wrestled it through the ICZN petition process, someone else would find a good solid way to diagnose the original holotype. That would be embarrassing.

The rare ventral-ish view of the Dystylosaurus dorsal vertebra BYU 4503. Sorry it’s not better. I do have 93 photos of it in this shelf, all of them individually pretty terrible, which I took in the forlorn hope that one day we’ll get photogrammetry software simple enough and clever enough to make some kind of model out of them.

So I think we need to simply accept that the name Dystylosaurus, while perfectly diagnosable based on its holotype and only specimen, is destined to remain a junior synonym for as long as Supersaurus is considered taxonomically valid.

But it does leave Dystylosaurus in a bit of a quantum superposition. When Supersaurus is considered diagnosable, it ceases to exist, like a cat in a box. When Supersaurus is considered undiagnosable, it pops back into existence, like … well, a cat in a box. It’s an unsatisfactory kind of existence, but I think that’s the way it has to be.

So Dystylosaurus has its day — and it ends up being disappointing. Despite being perfectly diagnosable, it’s dependent for its validity on our assessment of other taxa. Some fossils just can’t catch a break.

Having surveyed what we know from the published literature about Jensen’s Big Three sauropods, and what Matt and I concluded about its big cervical BYU 9024, and having thought a bit more about the size of the BYU 9024 animal, we’re getting to the point where we can consider what all this means for Jensen’s taxa.

The Supersaurus pelvis BYU 13018 in right lateral view, at the North American Museum of Ancient Life (NAMAL). Signage reads: “Supersaurus pelvis. In 1988 the pelvis of Supersaurus was discovered at Dry Mesa Quarry. Brian Versey, Cliff Miles and Ken Stadtman of Brigham Young University’s Earth Science Museum found the pelvis while they were trying to close the quarry for the season. The discovery generated a huge media event, making headlines around the world. This pelvis is the largest dinosaur bone complex ever discovered. It is on display here for the very first time. Specimen on loan from Brigham Young University’s Earth Science Museum. Late Jurassic/Early Cretaceous (about 144 million years ago)

As Curtice and Stadtman (2001:36-39) pointed out, Supersaurus is actually known from quite a lot of material, all assigned to the holotype individual. I’ll quote them at length rather than paraphrasing, but if you want a tabular summary, you can skip the quote and pick up down below.

Supersaurus vivianae roll call

The name “Supersaurus” first appeared in a Reader’s Digest article (George, 1973) describing a pair of 8′ long scapulocoracoids uncovered from Dry Mesa Dinosaur Quarry near Delta, Colorado. When formally described (Jensen, 1985) a number of elements were referred to the holotype including the left scapulocoracoid discovered in 1972 (BYU 9025), a right scapulocoracoid (BYU 12962), a right ischium (BYU 12946), a distal proximal caudal vertebra (BYU 12843) and 12 articulated mid-caudal vertebrae (BYU 9084). An additional caudal vertebra (BYU 9077) is referred to (and figured as) Supersaurus in the text of Jensen (1985). The specimen numbers used in Jensen (1985), no longer valid, have created confusion in the literature (e.g., Paul, 1988) and thus current BYU specimen numbers are used here throughout.

Jensen (1987) later referred a mid-cervical vertebra (BYU 9024) and Curtice and Curtice (1996) a proximal caudal vertebra (BYU 9045), both originally assigned to Ultrasauros, to Supersaurus. Numerous additional elements belong to Supersaurus, including a left ischium (BYU 12555), which is clearly the mate to the referred right ischium (BYU 12946), a right pubis (BYU 12424), a carpal (BYU 12390), a phalanx (BYU 9000), a left ulna (BYU 13744), at least five caudal vertebrae (BYU 4839, 9045, 12639, 12819, 12843) and a pelvis (BYU 13018) consisting of a left ilium and four sacral vertebrae.

Jensen never referred the two Supersaurus scapulocoracoids to the same individual due to a 260 mm discrepancy in length. Stripping away the paint and resin on BYU 9025 revealed the proximal end had been inadvertently lengthened during preservation. Close examination of the actual bone surface nets a total scapulocoracoid length less than 50 mm longer than BYU 12962, an amount easily accounted for by scapular variation and thus here both are referred to the same individual. Concerning the large brachiosaur scapulocoracoid (BYU 9462) Jensen (1985) listed as part of the material belonging to Ultrasauros, it is demonstrably smaller than the largest Tendaguru scapula and has been referred to Brachiosaurus sp. (Curtice and Curtice, 1996; Curtice et al., 1996). As such all exceptionally large sauropod elements from Dry Mesa Dinosaur Quarry can be referred to one of two individuals, one a Supersaurus and one a Brachiosaurus.

A dorsal vertebra (BYU 9044) referred to Supersaurus (Curtice and Curtice, 1996; Curtice et al., 1996) results in Ultrasaurus macintoshi becoming a junior synonym of Supersaurus vivianae, as BYU 9044 was the type specimen of Ultrasauros. A second dorsal vertebra, BYU 12814, is also here referred to Supersaurus based on its similarities to BYU 9044. All of the three large dorsal vertebrae mentioned herein were found within the confines of the paired Supersaurus scapulae further strengthening the suggestion all of the large diplodocid elements belong to a single individual.

(Yes, it really does say “a distal proximal caudal vertebra”.)

Curtice and Stadtman say that the pelvis consists of left ilium plus four sacral vertebrae; but as the photo above clearly shows, it is the right ilium that is preserved.

Here is a summary table, in standard anatomical order:

Specimen Element Referred by
9024 Mid-cervical vertebra Jensen 1987
4503 Anterior dorsal vertebra Curtice & Stadtman 2001
9044 Posterior dorsal vertebra Curtice et al. 1996
12814 Posterior dorsal vertebra Curtice & Stadtman 2001
13018 Pelvis (right ilium, four sacral vertebrae) Curtice & Stadtman 2001
9045 Proximal caudal vertebra Curtice & Curtice 1996
12843 “Distal proximal” caudal vertebra Jensen 1985
9084 Twelve articulated mid-caudal vertebrae Jensen 1985
9077 Caudal vertebra Jensen 1985
4839 Caudal vertebra Curtice & Stadtman 2001
9045 Caudal vertebra Curtice & Stadtman 2001
12639 Caudal vertebra Curtice & Stadtman 2001
12819 Caudal vertebra Curtice & Stadtman 2001
12843 Caudal vertebra Curtice & Stadtman 2001
9025 Left scapulocoracoid Holotype
12962 Right scapulocoracoid Jensen 1985
13744 Left ulna Curtice & Stadtman 2001
12390 Carpal Curtice & Stadtman 2001
12424 Right pubis Curtice & Stadtman 2001
12946 Right ischium Jensen 1985
12555 Left ischium Curtice & Stadtman 2001
9000 Phalanx Curtice & Stadtman 2001

This is an impressively complete specimen — especially for a giant sauropod, as these tend only to survive in the form of isolated elements.

But is it really one specimen? That’s the subject of the next post.

(This post is rather slender by recent standards. That’s because I accidentally hit Publish when it was only half written. Rather than leave it to slowly change as I write more, I think it’s better to let this first half stand as its own post, and write the rest as its own post next time.)

References

  • Curtice, Brian D. and Linda J. Curtice. 1996. Death of a dinosaur: a reevaluation of Ultrasauros macintoshi (Jensen 1985). Journal of Vertebrae Paleontology 16(3):26A.
  • 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.
  • 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.

In part 2, we concluded that BYU 9024, the large cervical vertebra assigned by Jensen to the Supersaurus holotype individual, is in fact a perfectly well-behaved Barosaurus cervical — just a much, much bigger one than we’ve been used to seeing. Although we heavily disclaimered our size estimates, Andrea Cau quite rightly commented:

Thanks for the disclaimer: unfortunately, it is going to be ignored by the Internet.
[…]
So, my boring-conservative mind asks: what is the smallest size that is a valid alternative explanation? I mean, if we combine all possible factors (position misinterpretation, deformation effects, allometry and so on) what could be the smallest plausible size? Only the latter should be taken as “the size” of this animal, pending more material.

Andrea is right that we should take a moment to think a bit more about the possible size implications of BYU 9024.

BYU 9024, the huge cervical vertebra assigned to Supersaurus but which is actually Barosaurus, in left dorsolateral view, lying on its right side with anterior to the right. In front of it, for scale, a Diplodocus cervical from about the same serial position. (Note that the Diplodocus vertebra here appears proportionally bigger than it really is, due to being much closer to the camera.)

What we know for sure is that the vertebra is 1380 mm long (give or take a centimeter or two due to the difficulties of measuring big complex bones in an objective way, something we should write about separately some time.)

We are 99% certain that the bone is a Barosaurus cervical.

We are much less certain about the serial position of that bone. When we were at BYU, we concluded that it most resembled C9 of the AMNH specimen, but I honestly can’t remember the detail of our reasoning (can you, Matt?) and our scanned notebooks don’t offer much in the way of help. We know from McIntosh (2005) that the neural spine of C8 is unsplit and that C9 has the first hint of a cleft.  How does that compare with BYU 9024? Here’s a photo to help you decide:

BYU 9024, large cervical vertebra in left dorsolateral view, inverted (i.e. with dorsal towards us and anterior to the right). Note the shallow cleft between metapophyses at bottom left.

And here’s an anaglyph, to help you appreciate the 3D structure. (Don’t have any red-cyan glasses? GET SOME!)

BYU 9024, oriented similarly to the previous photograph.

The morphology around the crown of the neural spine is difficult to interpret, partly just because the fossil itself is a bit smashed up and partly because the bone, the (minimal) restoration and the matrix are such similar colours. But here’s my best attempt to draw out what’s happening, zoomed in from last non-anaglyph photo:

As you start at the prezygapophyses and work backwards, the SPRLs fade out some way before you reach the crown, and disappear at or before what appears to be an ossified midline ligament scar projecting anteriorly from very near the top of the vertebra. Posterior to that are two small, tab-like metapophyses that appear almost like separate osteological features.

Now this is a very strange arrangement. Nothing like it occurs in any of the cervicals of Diplodocus, where all the way from C3 back to the last cervical, the SPRLs run continuously all the way up from the prezygs to the metapophyses:

Hatcher (1901:plate V). Diplodocus carnegii holotype CM 84, cervical vertebra 2-15 in anterior view.

What we’d love to do of course is compare this morphology with a similar plate of the AMNH Barosaurus cervicals in anterior view, but no such plate exists and no such photos can be taken due to the ongoing entombment of the vertebrae. So we’re reduced to feeding on scraps. McIntosh (2005:47) says:

The neural spine of cervical 8 is flat across the top, and that of cervical 9 shows the first trace of a divided spine (Fig. 2.2A). This division increases gradually in sequential vertebrae, being moderately developed in cervicals 12 and 13, and as a deep V-shape in cervicals 15 and 16.

Sadly, McIntosh illustrates only cervicals 8 and 13 in anterior view: Fig 2.2A does not illustrate C9, as the text implies. And neither of the illustrated vertebrae much resembles what we see in BYU 9024.

So while in 2016 we interpreted BYU 9024 is having “the first trace of a divided spine”, we do hold open the possibility that what we’re seeing is a vertebra in which the spine bifurcation is a little more developed than we’d realised, but with strange morphology that does not correspond closely to any well-preserved vertebra we’ve seen of any sauropod. (Most Barosaurus cervicals are either crushed and damaged; the well preserved ones outside of the AMNH walkway tomb are from a more anterior part of the neck where there is no bifurcation of the spine.)

There is one more possibility. Here is a truly lovely (privately owned) Barosaurus cervical in the prep lab at the North American Museum of Ancient Life (NAMAL):

Uncrushed Barosaurus cervical vertebra, serial position uncertain, in the NAMAL prep lab.

In this blessedly undistorted vertebra, we can see that the summit of the neural spine is flared, with laterally projecting laminae that are likely homologous with metapophyses. (The vertebra is symmetrical in this respect.) Might it be possible that the tab-like metapophyses of BYU 9024 were like this in life, but have been folded upwards post-mortem?

All of this leaves the serial position of the vertebra far from certain. But what we can do is compare it with the lengths of all the known AMNH Barosaurus vertebrae. Columns 1 and 2 in the table below show the serial position and total length of the AMNH cervicals. Column 3 shows the factor by which the 1370 mm length of BYU 9024 exceeds the relevant cervical, and column 4 shows the corresponding estimate for total neck length, based on 8.5 m (Wedel 2007:206–207) for AMNH Barosaurus.

Cv# Length (mm) BYU 9224 ratio BYU 9024 neck length
8 618 2.217 18.84
9 685 2.000 17.00
10 737 1.859 15.80
11 775 1.768 15.03
12 813 1.685 14.32
13 850 1.612 13.70
14 865 1.584 13.46
15 840 1.631 13.86
16 750 1.827 15.53

So to finally answer Andrea’s question from waaay back at the start of this post, the smallest possible interpretation of the BYU 9024 animal gives it a neck 1.584 times as long as that of the AMNH individual, which comes out around 13.5 m (and implies a total length of maybe 43 m).

But I don’t at all think that’s right: I am confident that the serial position of BYU 9024 is some way anterior to C14, likely no further back than C11 — which gives us a neck at least 15 m long (and a total length of maybe 48 m and a mass of maybe 12 × 1.768^3 = 66 tonnes).

 

References

  • Hatcher, Jonathan B. 1901. Diplodocus (Marsh): its osteology, taxonomy and probable habits, with a restoration of the skeleton. Memoirs of the Carnegie Museum 1:1-63 and plates I-XIII.
  • McIntosh, John S. 2005. The genus Barosaurus Marsh (Sauropoda, Diplodocidae). pp. 38-77 in: Virginia Tidwell and Ken Carpenter (eds.), Thunder Lizards: the Sauropodomorph Dinosaurs. Indiana University Press, Bloomington, Indiana. 495 pp.
  • Wedel, Mathew J. 2007. Postcranial pneumaticity in dinosaurs and the origin of the avian lung. Ph.D dissertation, Integrative Biology, University of California, Berkeley, CA. Advisors: Kevin Padian and Bill Clemens. 290 pages.