New paper out today: defining ‘cranial’ and ‘caudal’ for vertebrae

October 3, 2022

Vertebrae of Haplocanthosaurus (A-C) and a giraffe (D-F) illustrating three ways of orienting a vertebra: articular surfaces vertical — or at least the caudal articular surface vertical (A and D), floor of the neural canal horizontal (B and E), and similarity in articulation (C and F). See the paper for details! Taylor and Wedel (2002: fig. 6).

This is a lovely cosmic alignment: right after the 15th anniversary of this blog, Mike and I have our 11th coauthored publication (not counting abstracts and preprints) out today.

Taylor, Michael P., and Wedel, Mathew J. 2022. What do we mean by the directions “cranial” and “caudal” on a vertebra? Journal of Paleontological Techniques 25:1-24.

This one started back in 2018, with Mike’s post, What does it mean for a vertebra to be “horizontal”? That post and subsequent posts on the same topic (one, two, three) provoked interesting discussions in the comment threads, and convinced us that there was something here worth grappling with. We gave a presentation on the topic at the 1st Palaeontological Virtual Congress that December, which we made available as a preprint, which led to us writing the paper in the open, which led to another preprint (of the paper this time, not the talk).

Orienting vertebrae with the long axis of the centrum held horizontally seems simple enough, but choosing landmarks can be surprisingly complex. Taylor and Wedel (2022 fig. 5).

This project represented some interesting watersheds for us. It was not our first time turning a series of blog posts into a paper — see our 2013 paper on neural spine bifurcation for that — but it was our first time writing a joint paper in the open (Mike had started writing the Archbishop description in the open a few months earlier). It was also the last, or at least the most recent, manuscript that we released as a preprint, although we’ve released some conference presentations as preprints since then. I’m much less interested in preprints than I used to be, for reasons explained in this post, and I think Mike sees them as rather pointless if you’re writing the paper in the open anyway, which is his standard approach these days (Mike, feel free to correct me here or in the comments if I’m mischaracterizing your position).

So, we got it submitted, we got reviews, and then…we sat on them for a while. We have both struggled in the last few years with Getting Things Done, or at least Getting Things Finished (Mike’s account, my own), and this paper suffered from that. Part of the problem is that Mike and have far too many projects going at any one time. At last count, we have about 20 joint projects in various stages of gestation, and about 11 more that we’ve admitted we’re never going to get to (our To Don’t list), and that doesn’t count our collaborations with others (like the dozen or so papers I have planned with Jessie Atterholt). We simply can’t keep so many plates spinning, and we’re both working hard at pruning our project list and saying ‘no’ to new things — or, if we do think of new projects, we try to hand them off to others as quickly and cleanly as possible.

Two different ways of looking at a Haplocanthosaurus tail vertebra. Read on for a couple of recent real-life examples. Taylor and Wedel (2022: fig. 2).

Anyway, Mike got rolling on the revisions a few months ago, and it was accepted for publication sometime in late spring or early summer, I think. Normally it would have been published in days, but the Journal of Paleontological Techniques was moving between websites and servers, and that took a while. But Mike and I were in no tearing rush, and the paper is out today, so all is well.

One of the bits of the paper that I’m most proud of is the description of cheap and easy methods for determining the orientation of the neural canal. For neural canals that are open, either because they were fully prepped or never full of matrix to begin with, there’s the rolled-up-piece-of-paper method, which I believe first appeared on the blog back when I was posting photos of the tail vertebrae of the Brachiosaurus altithorax holotype. For neural canals that aren’t open, Mike came up with the Blu-tack-and-toothpick method, as shown in Figure 12 in the new paper:

A 3d print of NHMUK PV R2095, the holotype of Xenoposeidon, illustrating the toothpick method of determining neural canal orientation. Taylor and Wedel (2022: fig. 12).

I know both methods work because I recently had occasion to use them, studying the Haplocanthosaurus holotypes (see this post). For CM 572, the neural canal of the first caudal vertebra is full of matrix, so I used a variant of the toothpick method. I didn’t actually have Blu-tack or toothpicks, so I cut thin pieces of plastic from the edge of an SVP scale bar and stuck them in bits of kneadable eraser. It worked just fine:

The neural canal of caudal 2 was prepped, so I could use the rolled-up-piece-of-paper method:

(Incidentally, Mike and I refer to our low-tech orientation-visualizers as “neural-canal-inators”, in honor of Dr. Heinz Doofenshmirtz from Phineas and Ferb.)

In the above photos, notice how terribly thin the base of the neural arch is, antero-posteriorly. Both of these vertebrae are in pretty good shape, without much breakage or missing material, and their morphology is broadly consistent with that of other proximal caudals of Haplocanthosaurus, so we can’t write this off as distortion. As weird as it looks, this is just what Haplo proximal caudals were like. And with the neural canals held horizontally, the first two caudals end up oriented like so:

Now, as we pointed out in the paper, the titular question is not about determining the posture of the vertebrae in life, it’s about defining the directions ‘cranial’ and ‘caudal’ for isolated vertebrae — Mike asked the question back when for the holotype (single) dorsal vertebra of Xenoposeidon. But an interesting spin-off for me has been getting confronted with the weirdness of vertebrae whose articular surfaces are nowhere near orthogonal with their neural canals. I tilted those CM 572 Haplo caudals so that their neural canals were horizontal partly because that’s the preferred orientation that Mike and I landed on in the course of this work, but also partly because to me, that’s a more arresting image than the preceding ones with the articular faces held vertically. I’m both freaked out and fascinated, and that seems like a promising combination — there are mysteries here that cry out to be solved.

As usual, we have loads of people to thank. In addition to all those listed in the Acknowledgments of the new paper, I’m grateful to Matt Lamanna and Amy Henrici of the Carnegie Museum of Natural History for letting me play with study the Haplo specimens in their care. Mike and I also owe a huge thanks to the editorial team at the Journal of Paleontological Techniques. We reached out to them a few days ago to ask if it might be possible to get our in-press paper done and out in time for SV-POW!’s anniversary weekend, and they pitched in to make it happen.

What’s next? We weighed the evidence and formulated what the best solution we could think of. Now it’s up to the world to decide if that was a useful contribution. The comment thread is open — let’s find out.

6 Responses to “New paper out today: defining ‘cranial’ and ‘caudal’ for vertebrae”

  1. liviuurziceanu Says:

    There is recorded any footprint from a very big brachiosaurid? I have heard about a 90 cm diameter print from Spain, late Jurassic, made by forefront leg of a sauropod. Do you know if it is from a brachiosaurid? Is there any footprint which point to a dinosaur larger than Argentinosaurus? Kind regards

  2. Matt Wedel Says:

    There is recorded any footprint from a very big brachiosaurid?

    Not to my knowledge.

    I have heard about a 90 cm diameter print from Spain, late Jurassic, made by forefront leg of a sauropod. Do you know if it is from a brachiosaurid?

    I don’t.

    Is there any footprint which point to a dinosaur larger than Argentinosaurus?

    The giant tracks from the Australian Broome Sandstone are the biggest I know of, but no-one has published a rigorous comparison with Argentinosaurus, for which we have zero preserved feet.

  3. llewelly Says:

    Congratulations on the paper. It’s nice to have precise definitions of words. Now if only people will use them : )

  4. llewelly Says:

    In reply to liviuurziceanu , there’s a Maltese et al 2018 paper describing what they claim is the largest sauropod pes ever reported, with what is definitely among the best sauropod paper titles:

    _The real Bigfoot: a pes from Wyoming, USA is the largest sauropod pes ever reported and the northern-most occurrence of brachiosaurids in the Upper Jurassic Morrison Formation_

    which is only tangentially related to your question, but I’m so entertained by the paper title I decided to crowbar it in anyway.

  5. liviuurziceanu Says:

    @llewelly: very interesting document

  6. llewelly Says:

    Another paper liviuurziceanu may find interesting:

    _A three-dimensionally preserved sauropod manus impression from the Upper Jurassic of Portugal: Implications for sauropod manus shape and locomotor mechanics_

    Milan et al 2005

    From the Discussion:

    “Additionally, the impressions of digit I in these tracks indicate a very short pollex claw.
    In the cast described in this study, the impression of the pollex claw is proportionally smaller relative to manus size than in similar sauropod tracks from the Middle Jurassic of Portugal (Santos et al., 1994). This shows that the trackmaker possesed a relatively
    small pollex claw, consistent with the brachiosaurid
    manual morphology ”

    I think there’s another place in the same paper where they say it’s also consistent with other basal titanosauriforms.

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