I’m scrambling to get everything done before I leave for England and SVPCA this weekend, so no time for a substantive post. Instead, some goodies from old papers I’ve been reading. Explanations will have to come in the comments, if at all.

Streeter (1904: fig. 3). Compare to the next image down, and note that in birds and other reptiles the spinal cord runs the whole length of the vertebral column, in contrast to the situation in mammals.

Streeter (1904: fig. 3). Compare to the next image down, and note that in birds and other reptiles the spinal cord runs the whole length of the vertebral column, in contrast to the situation in mammals.

Nieuwenhuys (1964: fig. 1)

Nieuwenhuys (1964: fig. 1)

Butler and Hodos (1996: fig. 16.27)

Butler and Hodos (1996: fig. 16.27)

For more noodling about nerves, please see:

References

  • Butler, A.B., and Hodos, W. 1996. Comparative Vertebrate Neuroanatomy: Evolution and Adaptation. 514 pp. Wiley–Liss, New York.
  • Nieuwenhuys, R. (1964). Comparative anatomy of the spinal cord. Progress in Brain Research, 11, 1-57.
  • Streeter, G. L. (1904). The structure of the spinal cord of the ostrich. American Journal of Anatomy, 3(1), 1-27.

 

Marble Mountains trilobites

 

These animals experienced days less than 23 hours long, and years with close to 400 days.

We feature a lot of Brian Engh’s stuff here–enough that he has his own category. But lately he has really been outdoing himself.

The wave of awesome started last year, when Brian started posting videos showing builds and suit tests for monsters–monster suits, monster puppets, monster you-name-its. Like this monster-sculpting timelapse from last August:

And this suit test from last October:

Brian even wrote a blog post about how he builds monsters.

Things really ramped up this May with the release of “In Mountains”, the first video in a three-part series from Brian’s Earth Beasts Awaken album (which is badass, and available for free here).

If you’re thinking that the Mountain Monster has some Estemmenosuchus in its background, you are correct–that astonishing real-world critter was one of Brian’s inspirations, among many others.

More awesomeness is coming in July, when the next video, “Call to Awaken”, is slated to be released. Here’s a teaser:

I have even more exciting Brian-Engh-related news, but I am not at liberty to discuss that just yet. Hopefully sometime this fall. Stay tuned, true believers.

 

Last time, we looked at how including intervertebral cartilage changes the neutral pose of a neck — or, more specifically, of the sequence of cervical vertebrae. The key finding (which is inexplicably missing from the actual paper, Taylor and Wedel 2013c) is that adding cartilage of thickness x between vertebrae whose zygapophyses are height y above the mid-height of the centra elevates the joint’s neutral posture by x/y radians.

Figure 14. Geometry of opisthocoelous intervertebral joints. Hypothetical models of the geometry of an opisthocoelous intervertebral joint compared with the actual morphology of the C5/C6 joint in Sauroposeidon OMNH 53062. A. Model in which the condyle and cotyle are concentric and the radial thickness of the intervertebral cartilage is constant. B. Model in which the condyle and cotyle have the same geometry, but the condyle is displaced posteriorly so the anteropos- terior thickness of the intervertebral cartilage is constant. C. the C5/C6 joint in Sauroposeidon in right lateral view, traced from the x-ray scout image (see Figure 12); dorsal is to the left. Except for one area in the ventral half of the cotyle, the anteroposterior separation between the C5 cotyle and C6 condyle is remarkably uniform. All of the arrows in part C are 52 mm long.

Figure 14. Geometry of opisthocoelous intervertebral joints. Hypothetical models of the geometry of an opisthocoelous intervertebral joint compared with the actual morphology of the C5/C6 joint in Sauroposeidon OMNH 53062. A. Model in which the condyle and cotyle are concentric and the radial thickness of the intervertebral cartilage is constant. B. Model in which the condyle and cotyle have the same geometry, but the condyle is displaced posteriorly so the anteroposterior thickness of the intervertebral cartilage is constant. C. the C5/C6 joint in Sauroposeidon in right lateral view, traced from the x-ray scout image (see Figure 12); dorsal is to the left. Except for one area in the ventral half of the cotyle, the anteroposterior separation between the C5 cotyle and C6 condyle is remarkably uniform. All of the arrows in part C are 52 mm long.

But how thick was the intervertebral cartilage in sauropods?

At the moment, no-one really knows. As Kent Stevens (2013) points out in his contribution to the PLOS ONE sauropod gigantism collection:

Determining the ONP of a sauropod’s cervical vertebral column given only its bones requires is necessarily speculative since the cartilage, and thus the intervertebral spacing, is unknown.

Part of the our goal in our own PLOS collection paper (Taylor and Wedel 2013c) was to take some very tentative first steps towards estimating the cartilage thickness. To do this, we used two approaches. First, we looked at CT scans of articulated vertebrae; and second, we measured the cartilage thickness in a selection of extant animals and thought about what we could extrapolate.

Since the CT scans were Matt’s domain, I’m going to pass over those for now, in the hope that he’ll blog about that part of the paper. Here, I want to look at the extant-animal survey.

Figure 18. Cartilage in the neck of a rhea. Joint between cervicals 11 (left) and 10 (right) of a rhea, sagittally bisected. Left half of neck in medial view. The thin layers of cartilage lining the C11 condyle and C10 cotyle are clearly visible.

Figure 18. Cartilage in the neck of a rhea. Joint between cervicals 11 (left) and 10 (right) of a rhea, sagittally bisected. Left half of neck in medial view. The thin layers of cartilage lining the C11 condyle and C10 cotyle are clearly visible.

The first thing to say is that our survey is inadequate in many ways. We worked with the specimens we could get hold of, in the state we had them. This means that:

  • we have a very arbitrary selection of different animals,
  • they are at different ontogenetic stages, and
  • their cartilage thickness was measured by a variety of methods.

Our goal was not at all to reach anything like a definitive answer, but just to get the question properly asked, and so hopefully to catalyse much a more detailed survey.

With that proviso out of the way, here are our main results (from Table 4 of the paper, though here I have removed the sauropod CT-scan rows since we’ll be writing about those separately).

Taxon Thickness Reference Notes
Turkey 4.56% This study Difference in measurements of intact neck and articulated sequence of cleaned, degreased and dried vertebrae.
Ostrich 6.30% Cobley et al. (2013) Difference in measurements of individual vertebrae with and without cartilage.
Rhea 2.59% This study Measurement of in situ cartilage in bisected neck.
Alligator 14.90% This study Measurement of in situ cartilage from photograph of cross section.
Horse 6.90% This study Measurement of in situ cartilage from photograph of cross section.
Camel 13.00% This study Crude measurement from condyle margin to cotyle lip of lateral-view X-ray. This is an interim measurement, which we hope to improve on when we obtain better images.
Dog 17.00% This study Measurement of intervertebral gaps in lateral-view X-ray, uncorrected for likely concavity of cotyles.
Giraffe 24.00% This study Difference in measurement of intact neck and closely articulated sequence of cleaned vertebrae. Young juvenile specimen.
Muraenosaurus 14.00% Evans (1993) Measurement of in situ cartilage in fossils.
Cryptoclidus 20.00% Evans (1993) Measurement of in situ cartilage in fossils.

We’ve expressed the measurements as a ratio between cartilage thickness and the length of the bone itself — that is, cartilage/bone. Another way to think of this is that the percentage is a correction factor which you need to add onto bone length to get whole-segment length. Note that this is not the same ratio as the proportion of total segment length that consists of cartilage: that would be (cartilage thickness + bone length) / bone length.

(We also tossed in some measurements of plesiosaur neck cartilage that Mark Evans made way back when. Get that thing properly published, Mark!)

Even this small survey throws up some interesting points.

First, there is a huge range of proportional cartilage thicknesses: almost an order of magnitude from the 2.59% of the Rhea up to the 24% of the juvenile giraffe — or, even if you discard that because of its ontogenetic stage, up to 17% for the dog. And note that the 17% for the dog is probably an under-estimate, since we were working from an X-ray that doesn’t show the concavity of the vertebral cotyles.

Figure 22. Dog neck in X-ray. Neck of a dog (dachsund), in X-ray, with the seven cervical vertebrae indicated. This photo has been used with permission from the Cuyahoga Falls Veterinary Clinic.

Figure 22. Dog neck in X-ray. Neck of a dog (dachsund), in X-ray, with the seven cervical vertebrae indicated. This photo has been used with permission from the Cuyahoga Falls Veterinary Clinic.

(Two reviewers expressed scepticism that this is the usual condition for dogs, but this X-ray is consistent with those of other dogs illustrated in the veterinary literature.)

The second thing to note is that the cartilage measurements for birds (average 4.5%) are are much lower than those of crocodilians (14.9%) or mammals (15.2%). What does this mean? Among these groups, sauropods are most closely related to birds; but birds and crocs form the extant phylogenetic bracket, so we can’t tell from phylogeny alone whether to expect them to more closely approach the avian or crocodilian condition. Furthermore, in being opisthocoelous (condyle in front, cotyle at the back) sauropod cervicals most closely resemble those of mammals in gross structure — and they have the thickest cartilage of all.

The third thing to note is that there is considerable variation within groups. Although the cartilage is proportionally thin for all three birds, it’s more than twice as thick in the ostrich as in the rhea (although some of this could be due to the different measurement methods used for these two birds). More interestingly, among mammals the cartilage is twice as thick in camels as in horses. In the horse, the condyles are deeply inserted into the cotyles of the preceding vertebrae; but in camels, they don’t reach even the lip of the cotyle. This should worry us, as horse and camel cervicals are grossly similar, and no osteological correlates have been identified that would allow us to determine from the bones alone how very different the cartilage is between these two mammals. So it seems possible that there were similarly dramatic differences in the neck-cartilage thickness of different sauropods.

Note: I said that no osteological correlates have been identified. That doesn’t mean they don’t exist. One thing I would love to see is a serious attempt to analyse cartilage thickness across a broad range of mammals, and to examine the corresponding dry bones to see whether in fact there are correlates that could be informative in this respect. One lesson that Matt and I have learned over and over again is that there’s often plenty of data in places that are out in the open, but where no-one’s thought to look.

Next time: more on searching for osteological correlates of cartilage. Then, measurements of sauropod-neck cartilage from CT scans, and likely implications for cartilage thickness in life.

References

For a palaeontology blog, we don’t talk a lot about geology. Time to fix that, courtesy of my middle son Matthew, currently 13 years old, who made this helpful guide to the rock cycle as Geology homework.

rocky1

rocky2

rocky3

Mark Witton, pterosaur-wrangler, Cthulhu-conjurer, globe-trotting paleo playboy and all-around scientific badass, drew this (and blogged about it):

Buzzed small

I liked it, but I thought it could use some color, so I hacked a crude version in GIMP and sent it to Mark with a, “Hey, please put this on a t-shirt so I can throw money at you” plea. Lo and behold, he did just that.

Buzzed for Wedel - 480

You can get your own from Mark’s Zazzle store. And apparently he will have more sauropod-themed merch coming soon.

If your museum doesn't look like this, you should reconsider your existence.

If your museum doesn’t look like this, you should reconsider your existence.

We’re just back from SVPCA 2013 in Edinburgh. The first part of the meeting was held at the Royal Society of Edinburgh, but on Friday we moved to the National Museums Scotland. Which is awesome. And free to the public. The design process for the museum seems to have been, “Okay, let’s get one of, oh, every interesting thing in the world, and put it right here.” We have tons more photos of amazing things from the museum, and maybe we’ll get around to posting them sooner or later, but today I have other things to do.

This pathetic, racially senescent freak is destined for evolution's dustbin.

This pathetic, racially senescent freak is destined for evolution’s dustbin. And he knows it.

Like make fun of Mike. And talk about vomiting dinosaurs.

Dude, this party totally ro-BLAAAUUGGH!!

Dude, this party totally ro-BLAAAUUGGH!!

This groovy stuffed fulmar, Fulmarus glacialis, is shown in the act of puking, which it does to dissuade predators. And probably everyone else. I am reliably informed by Darren that this is unrealistic fulmar vomit, and that the real thing is  more of a thin stream, like the world’s nastiest water gun, which can be directed with considerable accuracy. Note to self: don’t piss off the fulmars.

Vomiting sauropod by Wedel and NichollsLast year cemented “drawing goofy sauropods down at the pub” as a regular SVPCA Thing. So one night I was out with Mike and Darren and paleoartist Bob Nicholls, who is famous around these parts as the creator of the Greatest. Paleoart. Ever. I did a goofy sketch in my notebook illustrating the “defensive vomit” hypothesis, which Brian Engh and I cooked up during this alligator dissection. More on that another time, maybe. Anyway, after bashing out a fairly pathetic sauropod-puking-on-theropod scene, I passed the notebook to Bob and said, “Make this not suck”. Which he did. (Seriously, if you could see my original scrawl, you’d be the one throwing up.)

So now I have an original Bob Nicholls sketch–heck, the world’s first Wedel-Nicholls artist collaboration!–in my notebook, of one of evolution’s most majestic successes responding appropriately to a vulgar, overstudied theropod. Bob drew it right in front of me and I got to drink good beer while I watched him work.

And that, more or less, is why I attend SVPCA.

Giant Irish Mike - cut out

I couldn’t sign off without giving you another version of Giant Irish Mike, with the background cropped out so he can be dropped right into posters, slide shows, and other works of science and art. I really, really hope that he turns up in conference talks and other presentations in the months and years to come. If so, send us a photo documenting his miraculous apparition and we’ll show it to the world.

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