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

When I visited Dinosaur National Monument in October with Brian Engh and Yara Haridy, we spent a decent amount of time checking out DNM 28, a skull and associated bits of Camarasaurus. In particular, I got some shots of the axis (the second cervical vertebra behind the head), and it got me thinking about pneumaticity in this unusual element. Why I failed to get a full set of orthogonal shots is quite beyond my capacity, but we can roll with what I have. Before we go on, you might want to revisit Tutorial 36 to brush up on the general parts of the atlas-axis complex.


Here’s the axis in left lateral view (so, anterior to the left).

And a labeled version of the same. A few things to note:

  • One oddity of sauropod axes (and of axes of most critters) is that not only are the articular facets of the prezygapophyses not set forward of the neural arch, they’re set backward, well behind the forward point of the arch.
  • The dens epistrophei or odontoid process is sticking out immediately below the neural canal. This is the tongue of bone that articulated with the atlas (first cervical vertebra) in life.
  • Check out the prominent epipophysis above the postzygapophysis, which anchored the long dorsal neck muscles. (For more on epipophyses, see these posts, and especially this one.)
  • The diapophysis and parapophysis articulated with a cervical rib, which is not shown here. In fact, I don’t remember seeing it in the drawer that this vert came from. The atlantal and axial cervical ribs are small, apparently fused late in life if they fused at all, and are easily lost through taphonomic processes.
  • At least three pneumatic features are visible in this lateral view: the lateral fossa on the centrum, which is referred to as the “pleurocoel” in a lot of older literature; a ventral fossa that lies between the parapophysis and the midline ventral keel; and a fossa on the neural arch, behind the postzygodiapophyseal lamina. In the nomenclature of Wilson et al. (2011), this is the postzygocentrodiapophyseal fossa.

“Postzygocentrodiapophyseal fossa” is a mouthful, but I think it’s the only way to go. To be unambiguous, anatomical terminology needs to references specific landmarks, and the schemes proposed by Wilson (1999) for vertebral laminae and Wilson et al. (2011) for vertebral fossae are the bee’s knees in my book.

Nomenclatural issues aside, how do we know that these fossae were all pneumatic? Well, they’re invasive, there’s no other soft-tissue system that makes invasive fossae like that in archosaur vertebrae (although crocs sometimes have shallow fossae that are filled with cartilage or fat), and the same fossae sometimes have unambiguous pneumatic foramina in other vertebrae or in other sauropods.

Most of the features labeled above are also visible on the right side of the vertebra, although the ventral fossa is a little less well-defined in this view, and I can’t make out the prezyg facet. Admittedly, some of the uncertainty here is because of my dumb shadow falling across the vertebra. Specimen photography fail!

The paired ventral fossae are more prominent in this ventral view, on either side of the midline ventral keel (anterior is to the top).

And here’s a labeled version of the same ventral view.

Finally, here’s the posterior view. It’s apparent now that the neural spine is a proportionally huge slab of bone, like a broad, tilted shield between the postzygapophyses (which are also quite large for the size of this vertebra). The back side of the neural spine is deeply excavated by a complex fossa with several subfossae (kudos again to Jeff Wilson [1999] for that eminently useful term).

Here’s the same shot with some features of interest labeled. If I’ve read Wilson et al. (2011) correctly, the whole space on the back side of the neural spine and above the postzygs could be considered the spinopostzygapophyseal fossa, but here I’ve left the interspinous ligament scar (ILS) unshaded, on the expectation that the pneumatic diverticula that created that fossa were separated on the midline by the interspinous ligament. I might have drawn the ILS too conservatively, conceivably the whole space between the large deeply-shadowed subfossae was occupied by the interspinous ligament.

I’m particularly interested in those three paired subfossae, which for convenience I’m simply calling A, B, and C. Subfossa A may just be the leftover space between the spinopostzgyapophyseal laminae laterally and the interspinous ligament medially. I think subfossa B is invading the ramus of bone that goes to the epipophysis and postzygapophysis, but I didn’t think to check and see how far it goes (that might require CT anyway).

Subfossa C is the most intriguing. Together, those paired fossae form a couple of shallow pits, just on either side of the midline, and aimed straight forward. They can’t be centropostzygapophyseal fossae, which used to be called peduncular fossae, because they’re not in the peduncles on either side of the neural canal, they’re up above the lamina that connects the two postzygapophyses. Could they be ligament attachments? Maybe, but I’m skeptical for at least four reasons:

  1. Although interspinous ligament attachments often manifest as pits in the cervical vertebrae of birds, in sauropods they usually form rugosities or even spikes of bone that stick out, not inward. Furthermore, these pits are smooth, not rough like the interspinous ligament scars of birds.
  2. The interspinous ligament in tetrapods is typically a single, midline structure, and these pits are paired.
  3. Similar pits in front of the neural spine are present in some sauropod caudals, and they appear to be pneumatic (see Wedel 2009: p. 11 and figure 9).
  4. Pits at the base of the neural spine seem to be fairly uncommon in sauropod vertebrae. If they were attachment scars from the universally-present interspinous ligaments, we should expect them to be more prominent and more widespread.

But if these paired pits are not ligament scars, what are they? Why are they present, and why are they so distinct? Sometimes (often?) subfossae and accessory laminae look like the outcome of pneumatic diverticulum and bone reacting to each other (I almost wrote ‘playing together’), in what looks like a haphazard process of adaptation to local loading. But the symmetry of these pits argues against them being incidental or random. They don’t seem to be going anywhere, so maaaybe they are the first hoofbeats of the embossed laminae and “unfossae” that we see in the vertebrae of more derived sauropods (for which see this post), but again, their symmetry in size and placement isn’t really consistent with the “developmental program gone wild” appearance of “unfossae”. I really don’t know what to make of them, but if you have ideas, arguments, or observations to bring to bear, the comment field is open.

In summary, sauropod axes are more interesting than I thought, even in a derpasaurus like Cam. I’ll have to pay more attention to them going forward.

References

 

Unworn:

Worn:

Spent some time last week just admiring these things. They’re pretty cool.

EDIT: in answer to Mike’s question in the first comment below, here’s a photo of some more worn teeth, showing that the level of wear in the one shown above is not unusual. Also, all of these worn teeth still had full roots, with no sign of the root resorption that would have preceded shedding of the tooth, so they were evidently going to be used for a while yet, probably a few months at least — BUT see the very useful comment from Jens Kosch below on the likely rapidity of tooth replacement in Camarasaurus.

DINO collections - more worn Camarasaurus teeth

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 am still building up to a big post on vertebral orientation, but in the meantime, check out this caudal vertebra of a Komodo dragon, Varanus komodoensis. This is right lateral view–the vert is strongly procoelous, and the articular ends of the centrum are really tilted relative to the long axis. I find this encouraging, for two reasons. First, it helped me clarify my thinking on how we ought to orient vertebrae, which Mike wrote about here and here. And second, it gives me some hope, because if we can figure out why tilting your articular surfaces makes functional sense in extant critters like monitors, maybe we can apply those lessons to sauropods and other extinct animals.

This is LACM Herpetology specimen 121971. Many thanks again to Neftali Camacho for access and assistance, and to Jessie Atterholt for basically doing all the other jobs while I was faffing about with this Komodo dragon.

No time right now for me to dig into the interesting and important discussion on how we should orient vertebrae (here and here so far) – that will be coming soon. In the meantime, here’s something else.

As printed, in one of WesternU’s 3D printers.

Coming off the tray.

Cleaned up and in my hand. This is a 70% scale print, so a little smaller than the original, but all the important morphology is clear enough. For one thing, I can finally make sense of the dorsal views of the vertebra.

I have been astonished at how useful a 3D print can be as an aid to thought. The caudals of the Snowmass Haplocanthosaurus are among the smallest sauropod vertebrae I’ve spent a lot of time with, and they’re still heavy enough and fragile enough that I don’t just whip them out and twirl them around in my fingers. But I can do that with the 3D prints, and it really helps ram the morphology home in my brain. There are a thousand subtle things I might not otherwise have noticed if I hadn’t been able to turn those shapes over easily in my hand. Not to mention the other things you can do with prints, like physically sculpt on them without gooping up your fossils (we’re midway through step #8 from that post, BTW).

Anyway, back to Xeno. Mike reminded me that I have seen the actual specimen in person exactly once, very briefly during our 2005 visit to the NHM collections when I was over there for SVPCA. But it wasn’t Xeno yet, and we had other fish to fry, including a lot of pneumatic and possibly-pneumatic stuff for me to see and photograph for my dissertation. So I have to admit that it didn’t register. Being able to handle it now, so much that Mike has written about it snaps into focus. Not that his writing isn’t clear, there’s just a huge gulf between the best written description and holding a thing in your hands.

Why do I have this thing? Partly to educate myself, partly because it’s relevant to a current project, and partly because we may not be done with Xeno. Stay tuned.

Many thanks to Gary Wisser for setting up the print, and to Jeff Macalino for pulling it for me.

We don’t post on pterosaurs very often, but I’m making an exception for Caelestiventus. Mostly because I had the unusual experience of holding a life-size 3D print of its skull a few days before it was published. Brooks Britt and George Engelmann are both attending Flugsaurier 2018 in Los Angeles, and Brooks gave a talk on the new pterosaur on Friday. It’s from the Upper Triassic Saints & Sinners Quarry in far northeastern Utah, which has also produced theropods, sphenosuchian crocs (like 80 individuals to date, no exaggeration), drepanosaurs (I’ve seen the material and that paper is going to be mind-blowing whenever it arrives), and other assorted hellasaurs. Some of that material is figured in the Britt et al. (2016) paper on the Saints & Sinners Quarry (a free download from the link below). As far as I know, the Caelestiventus paper is the second big volley on the Saints & Sinners material, out of what will probably be a long stream of important papers.

Anyway, since we’ve just been discussing the utility of 3D printing in paleontology (1, 2), I thought you’d like to see this. Brooks did caution us that the 3D model was a work in progress, and he now thinks that Caelestiventus had a more convex dorsal skull margin, with the downward forehead dip in the version that got printed being less prominent or absent. You can see a slightly different version in the skull recon drawn by second author Fabio M. Dalla Vecchia, which he kindly released into the public domain here.

Otherwise the 3D print is pretty good. The big plate below the orbit is weird and from what I gather not present in other dimorphodontids. Because the Saints & Sinners material was buried in sand, which is relatively incompressible compared to mud and clay, it’s all preserved in three dimensions with essentially no crushing. Caelestiventus therefore yields new information about Dimorphodon micronyx, which has been known since 1859 but mostly from pancaked material.

Stay tuned (in general, not here necessarily) for more on the remarkable tetrapods of the Sants & Sinners Quarry – the next few years are going to be very exciting. And since this may be my first and last Flugsaurier post, many thanks to the organizers for making it such an engaging and enjoyable experience, especially Mike Habib, Liz Martin-Silverstone, and Dave Hone.

References