Some quick backstory: lots of sauropods have long, overlapping cervical ribs, like the ones shown here in Sauroposeidon (diagram from this old post):

These long cervical ribs are ossified tendons of ventral neck muscles, presumably longus colli ventralis. We know they’re ossified tendons because of their bone histology (Klein et al. 2012), and we suspect that they’re longus colli ventralis because those tendons look the same in birds, just less ossified, as in this rhea (same specimens as these even older posts: 1, 2):

Diplodocoids have apomorphically short cervical ribs, which never extend very far past the end of their respective centra and sometimes don’t overlap at all. Still, we assume the long ventral neck muscles were there, just without long ossified tendons. Which brings me to Apatosaurus, which has cervical ribs that are anteroposteriorly short but famously massive, extending very below and/or to the sides of the cervical centra — for a truly breathtaking example see this post. Here are C3 through C7 in CM 3018, the holotype of Apatosaurus lousiae (Gilmore 1936: plate 24):

At least for me, it’s hard to resist the temptation to mentally scoot those vertebrae together into articulation, and imagine that the very swoopy-looking and maybe even down-turned cervical ribs allowed the ventral tendon bundles to wrap around the bottom of each cervical rib protuberance, something like this:

But it’s just not so, because like all 2D images, Gilmore’s plate distorts 3D reality. If you get to see the mounted skeleton in person, it’s clear that the cervical ribs are all more or less in line, and none of them are pointed at the big protuberances, which stick way out ventrolaterally.

Here I’ve drawn in the likely trajectories of the longus colli ventralis tendons. My little red pathways don’t precisely match the cervical ribs as mounted, but there’s a lot of distortion and restoration going on. For example, comparing with Gilmore’s plate we can see that the cervical ribs of C5, which point downward compared to all the others, only do that because someone forced them to — the whole anterior portion of the rib, where the shaft would actually join to the capitulum and tuberculum, is reconstructed. Even if I’m a little off, it’s clear that the cervical ribs shafts point backward, they’re all more or less in two parallel lines, and none of them point down and out toward the ventrolateral processes. The photo contains a mountain of useful morphological information that you’d never get from the lateral views.

My takeaways from all this:

  1. If a person has only seen 2D images of a specimen, and especially if those 2D images have only been orthogonal views with no obliques, their little island of knowledge is surrounded by at least a sizeable lake of ignorance, if not a small ocean.
  2. That doesn’t mean that seeing specimens in person is the only antidote — 3D models and 3D prints are extremely useful, and for specimens that are difficult to manipulate because of their size or fragility, they may be more useful than seeing or handling the specimen, at least for some questions.
  3. For Apatosaurus specifically, those ventrolateral processes cry out for explanation. They’re fairly solid knobs of bone that stick way out past the ossified tendons of the ventral-most neck muscles. That’s a super-weird — and super-expensive — place to invest a bunch of bone if you’re not using it for something fairly important, especially in a lineage that had just spent the last 80-100 million years making their necks as light as possible.
  4. Pursuant to that last point, we’re now in — ugh-ouch-shame — our 8th year of BrontoSMASH!!, with still just the one conference presentation to show for it (Taylor et al. 2015). Prolly time we got moving on that again.

References

I am co-authoring a manuscript that, among other things, tries to trace the history of the molds made by the Carnegie Museum in the early 1900s, from which they cast numerous replica skeletons of the Diplodocus carnegii mount (CM 84, CM 94, CM 307 and other contributing specimens). This turns out to be quite a mystery, and I have become fascinated by it.

Below is the relevant section of the manuscript as it now stands. Can anyone out there shed any further light on the mystery?


So far as we have been able to determine, the casting of the concrete Diplodocus of Vernal was probably the last time the Carnegie Museum’s original molds were used. However, that was not Untermann’s intention. In his 1959 account, he wrote (p368–369):

Several museums in the United States and from lands as distant as Japan and Italy have expressed a desire to acquire the molds and cast a Diplodocus of their own from either plaster or some of the newer synthetics. To date no museum has apparently been able to make satisfactory arrangement for the acquisition of the molds and the casting of a skeleton. We still have the molds in Vernal, and any museum, anywhere, is welcome to them just for hauling them off. […] The Diplodocus on the lawn of the Utah Field House is the eleventh replica to be cast from the molds […] Does anyone wish to cast the twelfth?

From here, though, the story becomes contradictory. Sassaman (1988) reported that “the molds finally fell apart because of old age soon after it [the concrete Diplodocus] was made”. Similarly, Ilja Niewland (pers. comm., 2022) said that “The original moulds were thrown away somewhere during the 1960s (nobody at the [Carnegie Museum] could be more specific than that)”, suggesting that the molds may have been returned to their origin.

Both these accounts seem to be in error, as shown by a 1960 report in the Vernal Express newspaper (Anonymous 1960a; Figure H; see also Carr and Hansen 2005). This says that in the middle of July 1960, the molds were collected by the Rocky Mount Children’s Museum (now the Rocky Mount Imperial Center, Children’s Museum & Science Centre) in North Carolina, with the intention that they would be used to create a twelfth cast which would be mounted outside the museum building next to the Tar River in Rocky Mount’s Sunset Park. But was such a cast ever created? A sequence of reports in the Rocky Mount Evening Telegram from April to July 1960 (Williams 1960, Bell 1960a, Bell 1960b, Anonymous 1960b) enthusiastically announce and discuss the impeding arrival, and the later articles say that museum board president Harold Minges has left for Utah to collect to molds — but then the newspaper goes silent on the subject, and the project is never mentioned again. There is no positive evidence that the molds even arrived in Rocky Mount, far less that they were used to create a new mount. Thus newspaper reports from both Utah and North Carolina say that the molds set out on their journey from one to the other, but neither confirms that they ever arrived. On the other hand, there is also no report of the molds being lost or destroyed, so perhaps the most likely interpretation is that they arrived in Rocky Mount, but were found to be in worse condition than expected and quietly left in storage. This interpretation is supported by Rea (2001:210) who reported that “from Vernal the molds kept travelling — first, to the Rocky Mount Children’s Museum in Rocky Mount, North Carolina, although a cast was never made there”. Similarly, Moore (2014:234-235) stated that “From Vernal, Utah, [CM] molds of Diplodocus carnegii are shipped to Rocky Mount Children’s Museum in Rocky Mount, North Carolina. Because of the age-related damage to the molds, a cast was never prepared”.

Hurricane Floyd devastated Rocky Mount in 1999, with flooding from the River Tar destroying the original Children’s Museum along with all its exhibits and records (Leigh White, pers. comm., 2022), so no records survive that could confirm the molds’ arrival or any subsequent use. The museum was located next door to a municipal water treatment facility that also flooded and released unknown chemicals, so museum property that might have otherwise been salvageable in that area was deemed contaminated and required to be destroyed. If the molds were in storage at the Children’s Museum at this time, then this was likely the end of their story.

The Children’s Museum was re-established at the newly built Imperial Centre, where it still resides, but no trace exists there of molds or casts of Diplodocus. Corroborating the hypothesis that no cast ever existed, most staff who worked at the museum in the 1980s do not recall any such cast (Leigh White, pers. comm., 2022). Contradicting this, however, Jan Engle Hicks, Curator of Education at the Rocky Mount Children’s Museum from 1971–2002, has a memory of Diplodocus casts being on exhibit at the museum when she started work in 1971. She does not recall if they were still part of the museum collection in 1999 when the collection was destroyed.

Whether or not a cast was made at Rocky Mount, it is possible that this was not the end for the molds. Rea (2001:210) continues: “Eventually the molds found their way to the Houston Museum of Science, where they were used to fill in gaps in the Diplodocus hayi skeleton that had been swapped from Pittsburgh to Cleveland before ending up in Houston”, citing a personal communication from John S. McIntosh. (The skeleton in question is that of CM 662, which became CMNH 10670 in Cleveland, then HMNS 175 in Houston. Having been nominated as the holotype of the new species Diplodocus hayi by Holland (1924:399), the species was later moved to its own new genus Galeamopus by Tschopp et al. (2015:267).)

Due to the loss of the Rocky Mount Children’s Museum records, we cannot tell whether they ever shipped the molds to Houston; and we have not been able to obtain information from the Houston Museum. Brian Curtice (pers. comm., 2022) reports that he was in Houston in 1995 and did not see the molds in the collection, nor hear of their ever having been there. In the absence of evidence that the molds ever made it to Houston, it seems at least equally likely that the missing bones in HMNS 175 were cast and supplied by Dinolab, using the second-generation molds described blow, and that Rea (2001) misreported this.

As recently as 1988, Rolfe (1988) wrote on behalf of the Royal Museum of Scotland, “At present I am exploring the possibility of re-using the Carnegie Museum, Pittsburgh moulds, although there is considerable doubt about whether they are up to the job, after so much previous use”. Sadly, his letter does not mention their then-current whereabouts.

In an unpublished manuscript, Madsen (1990:4) wrote that “The fate of the initial set of molds is somewhat in question, but Wann Langston (personal communication, 1989) suggests that they seem to have been lost, strayed, or stolen during transport from ? to ?. Principles contacted in regards to the disposition of the molds could not provide specific information.”. Infuriatingly, the question marks are in the original. Since both Langston and Madsen are now deceased, there is no way to discover on which of the molds’ journeys Langston thought they were lost or destroyed. It is unlikely, at least, that Langston had in mind the their initial journey from Vernal to Rocky Mount. Kirby (1998:4) wrote that “Somewhere along the line, as the story goes, the molds received from the Carnegie had been shipped to a school down south and never arrived. So they were lost”. Since Rocky mount is about 2000 miles east (not south) of Vernal, “a school down south” could not have referred, in a Utah publication, to a museum out east. The Houston museum also does not seems an especially likely candidate for this designation, being 1300 miles southeast of Vernal.

Putting it all together, there is no way that all the reports cited here can be accurate. Perhaps the most likely scenario is this: the molds were successfully shipped to Rocky Mount in July 1960 (Anonymous 1960a, Anonymous 1960b) but found to be unusable (Rea 2001:210, Moore 2014:234-235) and left in storage. At some later point there were shipped to a school in a southern state (Kirby 1998:4) but did not arrive (Langston cited in Madsen 1990:4). This may have happened in late 1988 or early 1989, between Rolfe’s (1988) letter that expressed an interest in using the molds and Langston’s personal communication to Madsen in 1989. Where the molds are now, and why they did not arrive, we can only speculate. As Madsen (1990:4) concluded, “It is truly a mystery that an estimated 3–6 tons of plaster molds could simply vanish!”

References

Anonymous. 1960a. Dinosaur molds take long ride to No. Carolina children’s home. Vernal Express, 14 July 1960, page 15. https://newspapers.lib.utah.edu/ark:/87278/s6zk6w6s/21338221

Anonymous. 1960b. Something ‘big’ for a fact. Rocky Mount Evening Telegram, 8 July 1960, page 4A. https://newspaperarchive.com/rocky-mount-evening-telegram-jul-08-1960-p-4/

Bell, Mae. 1960a. Dinosaur’s coming here brings questions galore. Rocky Mount Evening Telegram, 14 May 1960, page 2. https://newspaperarchive.com/rocky-mount-evening-telegram-may-14-1960-p-2/

Bell, Mae. 1960b. ‘Dinosaur’ soon to arrive here. Rocky Mount Evening Telegram, 3 July 1960, page 3A. https://newspaperarchive.com/rocky-mount-evening-telegram-jul-08-1960-p-8/

Carr, Elaine, and Aric Hansen. 2005. William Randolf Turnage, Dee Hall, and Ernest Untermann [archive photograph with metadata]. University of Utah, J. Willard Marriott Digital Library, image 1086142. https://collections.lib.utah.edu/details?id=1086142

Holland, William J. 1924. The skull of Diplodocus. Memoirs of the Carnegie Museum 9(3):379–403.

Kirby, Robert. 1998. Danny and the dinosaurs. Chamber Spirit (newsletter of the Vernal area Chamber of Commerce) 3(4):1–6.

Madsen, James H. 1990. Diplodocus carnegiei: Production and design of replica skeletons. Unpublished draft manuscript. (No author is named in the manuscript, but Madsen’s son Chris believes it is his work.)

Moore, Randy. 2014. Dinosaurs by the Decades: A Chronology of the Dinosaur in Science and Popular Culture. Greenwood, Westport, Connecticut.

Rea, Tom. 2001. Bone Wars: The Excavation and Celebrity of Andrew Carnegie’s Dinosaur. University of Pittsburgh Press, Pittsburgh, PA.

Rolfe, William D. I. 1988. Untitled letter to LuRae Caldwell (Utah Field House). 24 October 1988.

Sassaman, Richard. 1988. Carnegie had a dinosaur too. American Heritage 39(2):72–73.

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

Untermann, G. Ernest. 1959. A replica of Diplodocus. Curator 2(4):364–369. doi:10.1111/j.2151-6952.1959.tb00520.x

Williams, Oliver. Pre-historic dinosaur to tower over city; giant animal four times taller than man. Rocky Mount Evening Telegram, 24 April 1960, page 3B. https://newspaperarchive.com/rocky-mount-evening-telegram-apr-24-1960-p-11/

We’ve shown you the Apatosaurus louisae holotype mounted skeleton CM 3018 several times: shot from the hip, posing with another massive vertebrate, photographed from above, and more. Today we bring you a world first: Apatosaurus from below. Scroll and enjoy!

Obviously there’s a lot of perspective distortion here. You have to imagine yourself lying underneath the skeleton and looking up — as I was, when I took the short video that was converted into this image.

Many thanks to special-effects wizard Jarrod Davis for stitching the video into the glorious image you see here.

The most obvious effect of the perspective distortion is that the neck and tail both look tiny: we are effectively looking along them, the neck in posteroventral view and the tail in anteroventral. The ribs are also flared in this perspective, making Apato look even broader than it is in real life. Which is pretty broad. One odd effect of this is that this makes the scapulae look as though they are sitting on top of the ribcage rather than appressed to its sides.

 

Yes, we’ve touched on a similar subject in a previous tutorial, but today I want to make a really important point about writing anything of substance, whether it’s a scientific paper, a novel or the manual for a piece of software. It’s this: you have to actually do the work. And the way you do that is by first doing a bit of the work, then doing a bit more, and iterating until it’s all done. This is the only way to complete a project.

Yes, this is very basic advice. Yes, it’s almost tautological. But I think it needs saying because it’s a lesson that we seem to be hardwired to avoid learning. This, I assume, is why so many wise sayings have been coined on the subject. Everyone has heard that “A journey of a thousand miles begins with a single step”, attributed to Lao Tzu in maybe the 5th Century BC. More pithily, I recently discovered that Williams Wordsworth is supposed to have said:

To begin, begin.

I love that. In just three words, it makes the point that there is no secret to be learned here, no special thing that you can do to make beginning easier. You just have to do it. Fire up your favourite word processor. Create a new document. Start typing.

And to Wordsworth’s injunction, I would add this:

To continue, continue

Because, again, there is no secret. You just have to do it.

Mounted skeleton of Diplodocus carnegii holotype CM 84 in the rare dorsal view.

At the moment I am working on four separate but related papers. Honestly, sometimes it’s hard even keeping them straight in my head. Sometimes I forget which one I am editing. It would be easy to get overwhelmed and … just not finish. I don’t mean it would be easy to give up: that would be a decision, and I don’t think I would do that. But if I listened to my inner sluggard, I would just keep on not making progress until the matter become moot.

So here is what I do instead:

  • I pick one of the papers, which is the one I’m going to work on that evening, and I try not to think too much about the others.
  • I figure out what needs to be in that paper, in what order.
  • I write the headings into a document, and I put an empty paragraph below each, which just says “XXX”. That’s the marker I use to mean “work needed here”.
  • I use my word-processor’s document-structuring facilities to set the style of each of the headings accordingly — 1st, 2nd or occasionally 3rd level.
  • I auto-generate a table of contents so I can see if it all makes sense. If it doesn’t, I move my headings around and regenerate the table of contents, and I keep doing that until it does make sense.
  • I now have a manuscript that is 100% complete except in the tiny detail that it has no content. This is a big step! Now all I have to do is write the content, and I’ll be finished.
  • I write the content, one section at a time. I search for “XXX” to find an unwritten section, and I write it.
  • When all the “XXX” markers have been replaced by text, the paper is done — or, at least, ready to be submitted.

Caveats:

First, that list makes it sound like I am really good at this. I’m not. I suck. I get distracted. For example, I am writing this blog-post as a distraction from writing a section of the paper I’m currently working on. I check what’s new on Tweetdeck. I read an article or two. I go and make myself a cup of tea. I play a bit of guitar. But then I go back and write a bit more. I could be a lot more efficient. But the thing is, if you keep writing a bit more over and over again, in the end you finish.

Second, the path is rarely linear. Often I’m not able to complete the section I want to work on because I am waiting on someone else to get back to me about some technical point, or I need to find relevant literature, or I realise I’m going to need to make a big digression. That’s fine. I just leave an “XXX” at each point that I know I’m going to have to revisit. Then when the email comes in, or I find the paper, or I figure out how to handle the digression, I return to the “XXX” and fix it up.

Third, sometimes writing a section blows up into something bigger. That’s OK. Just make a decision. That’s how I ended up working on these four papers at the moment. I started with one, but a section of it kept growing and I realised it really wanted to be its own paper — so I cut it out of the first one and made it its own project. But then a section of that one grew into a third paper, and then a section of that one grew into a fourth. Not a problem. Sometimes, that’s the best way to generate new ideas for what to work on: just see what come spiralling out of what you’re already working on.

None of these caveats change the basic observation here, which is simply this: in order to get a piece of work completed, you first have to start, and then have to carry on until it’s done.

 

Here at SV-POW! Towers, we like to show you iconic mounted skeletons from unusual perspectives. Here’s one:

Apatosaurus louisae holotype CM 3018, mounted skeleton in the public gallery of the Carnegie Museum of Natural History: head, neck, torso and hip in right posterolateral view. Photograph by Matt Wedel, 12th March 2019 (my birthday!)

Oh, man, I love that museum. And I love that specimen. And I love the one that’s standing next to it (Diplodocus CM 82, natch.) I’ve got to find a way to get myself back out there.

That’s all: just enjoy.

For this forthcoming Barosaurus paper, we would like to include an establishing photo of the AMNH Barosaurus mount. There are two strong candidate photos which we’ve used before in an SVPCA talk, but since this is a formal publication we need to be more careful about copyright. Here are the photos, which are the property of their respective rightsholders:

This one is hard to find at all, at least using Google’s reverse image search. Whereas this one …

… is sprinkled all over the Internet, but (in all the cases I’ve seen so far) without attribution.

Does anyone have the necessary skills to track down who the rightsholder is for either of these? Thank you!

Here are some blank diagrams I whipped up for drawing in spinal cord pathways.

This one shows the whole cord, brainstem, thalamus, and cerebral cortex in coronal section, in cartoon form.

It’s for drawing in ascending sensory and descending motor pathways, as shown in this office hours sketch. DC-ML is dorsal column/medial lemniscus, which carries discriminative touch and conscious proprioception. ALS is anterolateral system, which carries pain, temperature, pressure, and itch. The lateral corticospinal tract carries fibers for voluntary control of major muscle groups. Each pathway differs in terms of where it decussates (crosses the midline, left-to-right and vice versa) and synapses (relays from one neuron to the next). The sensory pathways involve primary, secondary, and tertiary sensory neurons, and the motor pathways involve upper motor neurons (UMNs) and lower motor neurons (LMNs).

This one shows cross-sections of the cord at cervical, thoracic, lumbar, and sacral levels, for drawing ascending and descending pathways and thinking about how patterns of somatotopy come to exist.

Somatotopy is the physical representation of the body in the central nervous system. A common abbreviation scheme is A-T-L for arm-trunk-leg, as shown here for ascending sensory and descending motor pathways.

Finally, this one shows the spinal cord and spinal nerve roots at four adjacent spinal levels, for tracking the specific fates of sensory and motor neurons at each spinal level.

This is particularly useful when working out the consequences of an injury, like the spinal cord hemisection (Brown-Sequard syndrome) shown here in pink. The little human figure only shows the zone in which pain and temperature sensation are lost. There would also be losses of discriminative touch, conscious proprioception, and voluntary motor control on the same side as the injury.

Finally, since we’ve had a bit of a sauropod drought lately, here are a couple of photos of the mounted cast skeleton of Patagotitan in Stanley Field Hall at the Field Museum of Natural History in Chicago.

I gotta say, this mount beats the one at the AMNH in every way, because it’s well lit and you can move all the way around it and even look down on it from above. In fact, in terms of getting to move all the way around it, get well back from it to see the whole thing at once, and even walk directly underneath it (without having to ask permission to hop the fence), it might be the best-mounted sauropod skeleton in the world. The Brachiosaurus outside is also pretty great (evidence), but it loses points because you can’t walk around it on an upstairs balcony. Every other mounted sauropod I know of is either in more cramped surroundings, or you can’t get underneath it, or is less well-lit, or some combination of the above. Am I forgetting any worthy contenders? Feel free to make your case in the comments.

Incidentally, the spinal cord of Patagotitan was something like 120 feet long, and the longest DC-ML primary sensory neurons ran all the way from tail-tip to brainstem before they synapsed, making them among the longest cells in the history of life.

A belated thank-you to Josh Matthews and the rest of the Burpee PaleoFest crew for a fun day at the FMNH back in March. I got home from that trip about 3 days before the pandemic quarantine started, so it’s waaaaay past time for me to blog about how awesome that trip was. Watch this space. UPDATE: hey, look, it only took me a third of a year this time! Link.

Daniel Vidal et al.’s new paper in Scientific Reports (Vidal et al. 2020) has been out for a couple of days now. Dealing as it does with sauropod neck posture, it’s obviously of interest to me, and to Matt. (See our earlier relevant papers Taylor et al. 2009, Taylor and Wedel 2013 and Taylor 2014.)

Overview

To brutally over-summarise Vidal et al.’s paper, it comes down to this: they digitized the beautifully preserved and nearly complete skeleton of Spinophorosaurus, and digitally articulated the scans of the bones to make a virtual skeletal mount. In doing this, they were careful to consider the neutral pose of consecutive vertebrae in isolation, looking at only one pair at a time, so as to avoid any unconscious biases as to how the articulated column “should” look.

Then they took the resulting pose, objectively arrived at — shown above in their figure 1 — and looked to see what it told them. And as you can well see, it showed a dramatically different pose from that of the original reconstruction.

Original skeletal reconstruction of Spinophorosaurus nigerensis (Remes et al. 2009:figure 5, reversed for ease of comparison). Dimensions are based on GCP-CV-4229/NMB-1699-R, elements that are not represented are shaded. Scale bar = 1 m.

In particular, they found that as the sacrum is distinctly “wedged” (i.e. its anteroposterior length is greater ventrally than it is dorsally, giving it a functionally trapezoidal shape, shown in their figure 1A), so that the column of the torso is inclined 20 degrees dorsally relative to that of the tail. They also found lesser but still significant wedging in the last two dorsal vertebrae (figure 1B) and apparently some slight wedging in the first dorsal (figure 1C) and last cervical (figure 1D).

The upshot of all this is that their new reconstruction of Spinophorosaurus has a strongly inclined dorsal column, and consequently a strongly inclined cervical column in neutral pose.

Vidal et al. also note that all eusauropods have wedged sacra to a greater or lesser extent, and conclude that to varying degrees all eusauropods had a more inclined torso and neck than we have been used to reconstructing them with.

Response

I have to be careful about this paper, because its results flatter my preconceptions. I have always been a raised-neck advocate, and there is a temptation to leap onto any paper that reaches the same conclusion and see it as corroboration of my position.

The first thing to say is that the core observation is absolutely right, — and it’s one of those things that once it’s pointed out it’s so obvious that you wonder why you never made anything of it yourself. Yes, it’s true that sauropod sacra are wedged. It’s often difficult to see in lateral view because the ilia are usually fused to the sacral ribs, but when you see them in three dimensions it’s obvious. Occasionally you find a sacrum without its ilium, and then the wedging can hardly be missed … yet somehow, we’ve all been missing its implications for a century and a half.

Sacrum of Diplodocus AMNH 516 in left lateral and (for our purposes irrelevant) ventral views. (Osborn 1904 figure 3)

Of course this means that, other thing being equal, the tail and torso will not be parallel with each other, but will project in such a way that the angle between them, measured dorsally, is less than 180 degrees. And to be fair, Greg Paul has long been illustrating diplodocids with an upward kink to the tail, and some other palaeoartists have picked up on this — notably Scott Hartman with his very uncomfortable-looking Mamenchisaurus.

But I do have three important caveats that mean I can’t just take the conclusions of the Vidal et al. paper at face value.

1. Intervertebral cartilage

I know that we have rather banged on about this (Taylor and Wedel 2013, Taylor 2014) but it remains true that bones alone can tell us almost nothing about how vertebrae articulated. Unless we incorporate intervertebral cartilage into our models, they can only mislead us. To their credit, Vidal et al. are aware of this — though you wouldn’t know it from the actual paper, whose single mention of cartilage is in respect of a hypothesised cartilaginous suprascapula. But buried away the supplementary information is this rather despairing paragraph:

Cartilaginous Neutral Pose (CNP): the term was coined by Taylor for “the pose found when intervertebral cartilage [that separates the centra of adjacent vertebrae] is included”. Since the amount of inter-vertebral space cannot be certainly known for most fossil vertebrate taxa, true CNP will likely remain unknown for most taxa or always based on estimates.

Now this is true, so far as it goes: it’s usually impossible to know how much cartilage there was, and what shape it took, as only very unusual preservational conditions give us this information. But I don’t think that lets us out from the duty of recognising how crucial that cartilage is. It’s not enough just to say “It’s too hard to measure” and assume it didn’t exist. We need to be saying “Here are the results if we assume zero-thickness cartilage, here’s what we get if we assume cartilage thickness equal to 5% centrum length, and here’s what we get if we assume 10%”.

I really don’t think it’s good enough in 2020 to say “We know there was some intervertebral cartilage, but since we don’t know exactly how much we’re going to assume there was none at all”.

The thing about incorporating cartilage into articulating models is that we would, quite possibly, get crazy results. I refer you to the disturbing figure 4 in my 2014 paper:

Figure 4. Effect of adding cartilage to the neutral pose of the neck of Diplodocus carnegii CM 84. Images of vertebra from Hatcher (1901:plate III). At the bottom, the vertebrae are composed in a horizontal posture. Superimposed, the same vertebrae are shown inclined by the additional extension angles indicated in Table 2.

I imagine that taking cartilage into account for the Spinophorosaurus reconstruction might have given rise to equally crazy “neutral” postures. I can see why Vidal et al. might have been reluctant to open that can of worms; but the thing is, it’s a can that really needs opening.

2. Sacrum orientation

As Vidal et al.’s figure 1A clearly shows, the sacrum of Spinophorosaurus is indeed wedge-shaped, with the anterior articular surface of the first sacral forming an angle of 20 degrees relative to the posterior articular surface of the last:

But I don’t see why it follows that “the coalesced sacrum is situated so that the posterior face of the last sacral centrum is sub-vertical. This makes the presacral series slope dorsally and allows the tail to be subhorizontal (Figs. 1 and 4S)”. Vidal et al. justify this by saying:

Since a subhorizontal tail has been known to be present in the majority of known sauropods[27, 28, 29], the [osteologically induced curvature] of the tail of Spinophorosaurus is therefore compatible with this condition.

But those three numbered references are to Gilmore 1932, Coombs 1975 and Bakker 1968 — three venerable papers, all over fifty years old, dating from a period long before the current understanding of sauropod posture. What’s more, each of those three was about disproving the previously widespread assumption of tail-dragging in sauropods, but the wedged sacrum of Spinophorosaurus if anything suggests the opposite posture.

So my question is, given that the dorsal and caudal portions of the vertebral column are at some specific angle to each other, how do we decide which (if either) is horizontal, and which is inclined?

Three interpretations of the wedged sacrum of Spinophorosaurus, in right lateral view. In all three, the green line represents the trajectory of the dorsal column in the torso, and the red line that of the caudal column. At the top, the tail is horizontal (as favoured by Vidal et al. 2020) resulting in an inclined torso; at the bottom, the torso is horizontal, resulting in a dorsally inclined tail; in the middle, an intermediate posture shows both the torso and the tail slightly inclined.

I am not convinced that the evidence presented by Vidal et al. persuasively favours any of these possibilities over the others. (They restore the forequarters of Spinophorosaurus with a very vertical and ventrally positioned scapula in order to enable the forefeet to reach the ground; this may be correct or it may not, but it’s by no means certain — especially as the humeri are cross-scaled from a referred specimen and the radius, ulna and manus completely unknown.)

3. Distortion

Finally, we should mention the problem of distortion. This is not really a criticism of the paper, just a warning that sacra as preserved should not be taken as gospel. I have no statistics or even systematic observations to back up this assertion, but the impression I have, from having looked closely at quite a lot of sauropod vertebra, is the sacra are perhaps more prone to distortion than most vertebrae. So, for example, the very extreme almost 30-degree wedging that Vidal et al. observed in the sacrum of the Brachiosaurus altithorax holotype FMNH PR 25107 should perhaps not be taken at face value.

Now what?

Vidal el al. are obviously onto something. Sauropod sacra are screwy, and I’m glad they have drawn attention in a systematic way to something that had only been alluded to in passing previously, and often in a way that made it seems as though the wedging they describe was unique to a few special specimens. So it’s good that this paper is out there.

But we really do need to see it as only a beginning. Some of the things I want to see:

  • Taking cartilage into account. If this results in silly postures, we need to understand why that is the case, not just pretend the problem doesn’t exist.
  • Comparison of sauropod sacra with those of other animals — most important, extant animals whose actual posture we can observe. This might be able to tell us whether wedging really has the implications for posture that we’re assuming.
  • Better justification of the claim that the torso rather than the tail was inclined.
  • An emerging consensus on sauropod shoulder articulation, since this also bears on torso orientation. (I don’t really have a position on this, but I think Matt does.)
  • The digital Spinophorosaurus model used in this study. (The paper says “The digital fossils used to build the virtual skeleton are deposited and accessioned at the Museo Paleontológico de Elche” but there is no link, I can’t easily find them on the website and they really should be published alongside the paper.)

Anyway, this is a good beginning. Onward and upward!

References

  • Bakker, Robert T. 1968. The Superiority of Dinosaurs. Discovery 3:11–22.
  • Coombs, Walter P. 1975. Sauropod habits and habitats. Palaeogeography, Palaeoclimatology, Palaeoecology 17:1-33.
  • Gilmore, Charles W. 1932. On a newly mounted skeleton of Diplodocus in the United States National Museum. Proceedings of the United States National Museum 81:1-21.
  • Hatcher, John Bell. 1901. Diplodocus (Marsh): its osteology, taxonomy, and probable habits, with a restoration of the skeleton. Memoirs of the Carnegie Museum 1:1-63.
  • Osborn, Henry F. 1904. Manus, sacrum and caudals of Sauropoda. Bulletin of the American Museum of Natural History 20:181-190.
  • Taylor, Michael P. 2014. Quantifying the effect of intervertebral cartilage on neutral posture in the necks of sauropod dinosaurs. PeerJ 2:e712. doi:10.7717/peerj.712
  • Taylor, Michael P., and Mathew J. Wedel. 2013c. The effect of intervertebral cartilage on neutral posture and range of motion in the necks of sauropod dinosaurs. PLOS ONE 8(10):e78214. 17 pages. doi:10.1371/journal.pone.0078214
  • Taylor, Michael P., Mathew J. Wedel and Darren Naish. 2009. Head and neck posture in sauropod dinosaurs inferred from extant animals. Acta Palaeontologica Polonica 54(2):213-230.
  • Vidal, Daniel, P Mocho, A. Aberasturi, J. L. Sanz and F. Ortega. 2020. High browsing skeletal adaptations in Spinophorosaurus reveal an evolutionary innovation in sauropod dinosaurs. Scientific Reports 10(6638). Indispensible supplementary information at https://static-content.springer.com/esm/art%3A10.1038%2Fs41598-020-63439-0/MediaObjects/41598_2020_63439_MOESM1_ESM.pdf
    doi:10.1038/s41598-020-63439-0

Storm Giant

March 12, 2020

No, not his new Brachiosaurus humerus — his photograph of the Chicago Brachiosaurus mount, which he cut out and cleaned up seven years ago:

This image has been on quite a journey. Since Matt published this cleaned-up photo, and furnished it under the Creative Commons Attribution (CC By) licence, it has been adopted as the lead image of Wikipedia’s Brachiosaurus page [archvied]:

Consequently (I assume) it has now become Google’s top hit for brachiosaurus skeleton:

Last Saturday, Fiona and I went to Birdland, a birds-only zoo in the Cotswolds, about an hour away from where we live. The admission price also includes “Jurassic Journey”, a walking tour of a dozen or so not-very-good dinosaur models. In an interpretive centre in this area, I found this Brachiosaurus skeletal reconstruction stencilled on the wall:

I immediately knew it was the Chicago mount due to the combination of Giraffatitan anterior dorsals and Brachiosaurus posterior dorsals; but I found it more hauntingly familiar than that. A quick hunt turned up Matt’s seven-year-old post, and when I told Matt about my discovery he filled me in on its use in Wikipedia.

So this is 99% of a good story: we’re delighted that this work is out there, and has resulted in a much better Brachiosaurus image at Birdland than the rather sad-looking Stegosaurus next to it. The only slight disappointment is that I couldn’t find any sign of credit, which they really should have included given that Matt put the image out under CC By rather than in the public domain.

But as Matt said: “Even though I didn’t get credited, I’m always chuffed to see my stuff out in the world.” So true.