Sauroposeidon and friends

February 24, 2014

Sauroposeidon and kin cervicals - DRAFTAs a break from photography posts, here are four pretty big vertebrae that swirl in the same thought-space in my head. All are shown to scale, in right lateral view. These are not the biggest sauropod cervical vertebrae–Supersaurus beats them all, and there are vertebrae of Puertasaurus, Alamosaurus, and Futalognkosaurus that rival the big Sauroposeidon vert, but those are either less well preserved or still awaiting detailed description.

Incidentally, I think BYU 12867 is a C10. The centrum proportions are about right, compared to Giraffatitan, and the neural spine looks good, too, like a geometric transformation of the big Giraffatitan C8. Also, the drawn-in prezyg outline for MIWG.7306 is a little short; the actual prezyg is a monster and would have overhung the condyle by another 10cm or so. I’m pretty sure that we had a composite photograph showing this at one point, but irritatingly none of us can find it at the moment. If it turns up, I’ll update the image.

For a long time I thought Sauroposeidon was a brachiosaurid. Now it seems to be a somphospondyl (D’Emic 2012) or possibly even a basal titanosaur (Mannion et al. 2013), even if we stick just to the holotype. But if it’s not a brachiosaurid, it’s cervical vertebrae are at least coarsely brachiosaur-y in outline.

You  may recall from Naish et al. (2004) that MIWG.7306 shares several derived characters with the holotype vertebrae of Sauroposeidon. Does that mean that Angloposeidon is a somphospondyl or titanosaur as well? I dunno–as always, we need more material–but it’s an interesting possibility.

References

Following on from Matt’s post about the difficulty of photographing big specimens without distortion, I thought I’d have a play with our best Sauroposeidon C8 photo, which I think is this one:

sauroposeidon-c8-alone

(That’s been the basis for classic SV-POW! posts such as Your neck is pathetic and Darren’s new indeterminate Wealden maniraptoran is inadequate.)

I was motivated by Andy Farke’s comment:

Another–and perhaps more important–area where surface models excel is when you can remove colors on the original specimen that wash out relevant details…I bet this is probably the case for the example vertebra of Sauroposeidon. How many fossae and foramina just don’t show up well on the photos above?

Andy was talking about completely colourless 3d surface models, in which the 3d shape allows a render to make shadows that bring out the subtle shapes. But it made me wonder whether we could get anywhere just by washing out the most prevalent colour in the photo.

I started by doing a big, fat Gaussian blur on a duplicate layer — 500 pixels in each direction — and sampling the colour in the middle, to get a rough-and-ready average. (There may be a better way — please shout if you know one.) That average colour was#7e6b2f. I used it to run Colour To Alpha on another duplicate of the original layer, so that we’d be left with only residual colours. Here’s the result:

sauroposeidon-c8-alone-colour-completely-removed

I’m in two minds about this. It may be informative, but it sure is ugly. To compromise, I reinstated the original layer underneath this mostly-transparent one, and turned its opacity down to 75%. Here’s the result — a nice compromise:

sauroposeidon-c8-alone-colour-removed

Of course, there are endless other approaches you can take — that’s the blessing and the curse of image-editing programs like GIMP. For example, here’s what I got doing a simple Colours → Auto → White Balance:

sauroposeidon-c8-alone-whitebalanced

I’m not sure that isn’t the best of the bunch, in terms of informativeness.

I also tried something else — not amazingly successfully, but I think it’s worth seeing. Since the two photos that Matt showed in the previous post were evidently taken from somewhat different angles, I thought I’d have a go at compositing them into a red-cyan anaglyph. Because the variation in camera position is mostly dorsoventral rather than anteroposterior, the vert has to be pointed upwards for the two eyes to see the two versions from different horizontal points. Here’s the best I could do:

c8-anaglyph

I would say this is of some value; but it’s nowhere near as good as, for example, the anaglyph of Cervical S of the Archbishop. I could sit and look at that one all day. The problems with this one arise for three reasons.

First, I had to reduce both parts of the Sauroposeidon anaglyph to monochrome (since one was already in that form), so all colour information was lost.

Second, I had to scale the high-resolution picture to the same size as the lower-resolution one, throwing away more detail.

Finally, and most important, the two photos were not taken with the intention that they should be used to make an anaglyph. To work well, this has to be done with the images taken under the same lighting conditions, at the same distance from the specimen, from perspectives differing by about the distance between the pupils of the viewer, and with the camera-position difference being perfectly in the plane of the specimen. Needless to say, none of these conditions was met in this case, so it’s actually quite impressive that it works as well as it does.

We have a lot of options for illustrating specimens these days. Postage-stamp-sized greyscale photos really don’t cut it any more.

Here are two photos of what I infer to be C8 of OMNH 53062, the holotype of Sauroposeidon. The top one was taken by Mike during our visit to the OMNH in 2007. If you’re a regular you may recognize it from several older posts: 1, 2, 3. The bottom one was taken by Mike Callaghan, the former museum photographer at the OMNH, sometime in 1999 or 2000. I used it in Wedel et al. (2000) and Wedel and Cifelli (2005).

Sauroposeidon OMNH 53062 C8 photos compared

You’ll notice that the two photos are far from identical. In both cases, the photographers were up on ladders, as far above the vertebra as they could get, and there are still significant perspective effects. That’s just a fact of life when you’re taking photos of a vertebra that is 1.4 meters long, from anything lower than a helicopter. In Mike Taylor’s shot, the neural spine looms a little too large; in Mike Callaghan’s shot, the prezygapophysis looks a little too small, probably because it was curving off at the edge of the shot. So neither photograph is “right”; both distort the morphology of the specimen in different ways. Here’s how the two images stack up, with the outlines scaled to the same length:

Sauroposeidon OMNH 53062 C8 outlines compared

When I ran a draft of this post past Mike, he wrote (with permission to post):

I think the current draft misses an important point: the warning. We really can’t trust photos, however carefully taken, and however beautifully composited into TNFs*. You’re welcome to quote me as having said I’d have assumed the two C8s were different vertebrae. For that matter, I bet I could have worked up several taxonomically significant characters to distinguish them. Yikes.

* TNF = Taylor Normal Form, i.e., making multi-view photos like the ones here and here.

So the moral is, photos of big specimens almost always involve some distortion. This is clearly not ideal. But I have a plan for fixing it. I am hoping to get back to the OMNH this spring, and the next time I’m there, I’m going to take photos of this vertebra from a zillion angles and make a 3D model through photogrammetry. Happily, Heinrich Mallison has been producing a very helpful series of tutorials on that very topic over at dinosaurpaleo: 1, 2, 3, 4, with more on the way (I’ll update the links here later). Update: Don’t forget to check out Peter Falkingham’s (2012) paper in PE on making photogrammetric models with free software.

Armed with that model, it should be possible to produce a perspective-free lateral view image of the vertebra, to which all of the previous photos can be compared. I can’t use CT data because this vertebra has never been CTed; it’s too big to fit through a medical CT scanner, and probably too fragile to be packed up and shipped to an industrial CT machine like they used on Sue (not to mention that would require a significant chunk of money, which is probably not worth spending on a problem that can be solved in other ways).

So, photogrammetry to the rescue, or am I just deluding myself? Let me know what you think in the comments.

Finally, I should mention that the idea of superseding photographs with 3D photogrammetric models is not original. I got religion last week while I was having beers with Martin Sander and he was showing me some of the models he’s made. He said that going forward, he was going to forbid his students to illustrate their specimens only with photographs; as far as he was concerned, now that 3D models could be cheaply and easily produced by just about everyone, they should be the new standard. Inspiring stuff–now I must go do likewise.

Some previous posts on Sauroposeidon:

References

[This is part 4 in an ongoing series on our recent PLOS ONE paper on sauropod neck cartilage. See also part 1, part 2, and part 3.]

Big Bend Vanessa 182 small

Weird stuff on the ground, Big Bend, 2007.

Here’s a frequently-reproduced quote from Darwin:

About thirty years ago there was much talk that geologists ought only to observe and not theorise; and I well remember some one saying that at this rate a man might as well go into a gravel-pit and count the pebbles and describe the colours. How odd it is that anyone should not see that all observation must be for or against some view if it is to be of any service!

It’s from a letter to Henry Fawcett, dated September 18, 1861, and you can read the whole thing here.

I’ve known this quote for ages, having been introduced to it at Berkeley–a copy used to be taped to the door of the Padian Lab, and may still be. It’s come back to haunt me recently, though. An even stronger version would run something like, “If you don’t know what you’re looking for, you won’t make the observation in the first place!”

OLYMPUS DIGITAL CAMERA

Kent Sanders looking at scans of BYU 12613, a posterior cervical of either Kaatedocus or an anomalously small Diplodocus, at the University of Utah in May, 2008.

For example: I started CT scanning sauropod vertebrae with Rich Cifelli and Kent Sanders back in January, 1998. Back then, I was interested in pneumaticity, so that’s what I looked for, and that’s what I found–work which culminated in Wedel et al. (2000) and Wedel (2003). It wasn’t until earlier this year that I wondered if it would be possible to determine the spacing of articulated vertebrae from CT scans. So everything I’m going to show you, I technically saw 15 years ago, but only in the sense of “it crossed my visual field.” None of it registered at the time, because I wasn’t looking for it.

A corollary I can’t help noting in passing: one of the under-appreciated benefits of expanding your knowledge base is that it allows you to actually make more observations. Many aspects of nature only appear noteworthy once you have a framework in which to see them.

OLYMPUS DIGITAL CAMERA

BYI 12613 going through a CT scanner at the University of Utah medical center. We were filming for the “Megasaurus” episode of Jurassic CSI. That shoot was crazy fun.

So anyway, the very first specimen we scanned way back when was the most anterior of the three plaster jackets that contain the four cervical vertebrae that make up OMNH 53062, which was destined to become the holotype of Sauroposeidon. I’ve written about the taphonomy of that specimen here, and you can read more about how it was excavated in Wedel and Cifelli (2005). We scanned that jacket first because, although the partial vertebrae it contains are by far the most incomplete of the four, the jacket is a lot smaller and lighter than the other two (which weigh hundreds of pounds apiece). Right away we saw internal chambers in the vertebrae, and that led to all of the pneumaticity work mentioned above.

Sauroposeidon C5 cross section Wedel 2007b fig 14

Internal structure of a cervical vertebra of Sauroposeidon, OMNH 53062. A, parts of two vertebrae from the middle of the neck. The field crew that dug up the bones cut though one of them to divide the specimen into manageable pieces. B, cross section of C6 in posterior view at the level of the break, traced from a CT image and photographs of the broken end. The left side of the specimen was facing up in the field and the bone on that side is badly weathered. Over most of the broken surface the internal structure is covered by plaster or too damaged to trace, but it is cleanly exposed on the upper right side (outlined). C, the internal structure of that part of the vertebra, traced from a photograph. The arrows indicate the thickness of the bone at several points, as measured with a pair of digital calipers. The camellae are filled with sandstone. Wedel (2007: fig. 14).

Happily for me, that first jacket contains not only the posterior two-thirds of the first vertebra (possibly C5), but also the front end of the second vertebra. Whoever decided to plow through the second vertebra to divide the specimen into manageable chunks in the field made a savvy choice. Way back in 2004 I realized that the cut edge of the second vertebra was not obscured by plaster, and therefore the internal structure could be seen and measured directly, which is a lot cleaner than relying on the artifact-heavy CT scans. (The CT scans are noisy because the hospital machines we had access to start to pant a bit when asked to punch x-rays through specimens this large and dense.) A figure derived from that work made it into a couple of papers and this post, and appears again above.

But that’s pneumaticity, which this post is allegedly not about. The cut through the second vertebra was also smart because it left the intervertebral joint intact.

Figure 11. Fifth and partial sixth cervical vertebrae of Sauroposeidon. Photograph and x-ray scout image of C5 and the anterior portion of C6 of Sauroposeidon OMNH 53062 in right lateral view. The anterior third of C5 eroded away before the vertebra was collected. C6 was deliberately cut through in the field to break the multi-meter specimen into manageable pieces for jacketing (see [37] for details). Note that the silhouettes of the cotyle of C5 and the condyle of C6 are visible in the x-ray.

Fifth and partial sixth cervical vertebrae of Sauroposeidon.
Photograph and x-ray scout image of C5 and the anterior portion of C6 of Sauroposeidon OMNH 53062 in right lateral view. The anterior third of C5 eroded away before the vertebra was collected. C6 was deliberately cut through in the field to break the multi-meter specimen into manageable pieces for jacketing (see Wedel and Cifelli 2005 for details). Note that the silhouettes of the cotyle of C5 and the condyle of C6 are visible in the x-ray. Taylor and Wedel (2013: figure 11).

Here are a photo of the jacket and a lateral scout x-ray. The weird rectangles toward the left and right ends of the x-ray are boards built into the bottom of the jacket to strengthen it.

Figure 12. CT slices from fifth cervical vertebrae of Sauroposeidon. X-ray scout image and three posterior-view CT slices through the C5/C6 intervertebral joint in Sauroposeidon OMNH 53062. In the bottom half of figure, structures from C6 are traced in red and those from C5 are traced in blue. Note that the condyle of C6 is centered in the cotyle of C5 and that the right zygapophyses are in articulation.

CT slices from fifth cervical vertebrae of Sauroposeidon.
X-ray scout image and three posterior-view CT slices through the C5/C6 intervertebral joint in Sauroposeidon OMNH 53062. In the bottom half of figure, structures from C6 are traced in red and those from C5 are traced in blue. Note that the condyle of C6 is centered in the cotyle of C5 and that the right zygapophyses are in articulation. Taylor and Wedel (2013: figure 12).

And here’s a closeup of the C5/C6 joint, with the relevant radiographs and tracing. The exciting thing here is that the condyle is centered almost perfectly in the cotyle, and the zygapophyses are in articulation. Together with the lack of disarticulation in the cervical rib bundle (read more about that here and in Wedel et al. 2000), these things suggest to us that the vertebrae are spaced pretty much as they were in life. If so, then the spacing between the vertebrae now tells us the thickness of the soft tissue that separated the vertebrae in life.

I should point out here that we can’t prove that the spacing between the vertebrae is still the same as it was in life. But if some mysterious force moved them closer together or farther apart, it did so (1) without  decentering the condyle of C6 within the cotyle of C5, (2) without moving the one surviving zygapophyseal joint out of contact, and (3) without disarticulating the cervical ribs. The cervical ribs were each over 3 meters long in life and they formed vertically-stacked bundles on either side below the vertebrae; that’s a lot of stuff to move just through any hypothetical contraction or expansion of the intervertebral soft tissues after death. In fact, I would not be surprised if the intervertebral soft tissues did contract or expand after death–but I don’t think they moved the vertebrae, which are comparatively immense. The cartilage probably pulled away from the bone as it rotted, allowing sediment in. Certainly every nook and cranny of the specimen is packed with fine-grained sandstone now.

Anyway, barring actual preserved cartilage, this is a best-case scenario for trying to infer intervertebral spacing in a fossil. If articulation of the centra, zygs, and cervical ribs doesn’t indicate legitimate geometry, nothing ever will. So if we’re going to use the fossils to help settle this at all, we’re never going to have a better place to start.

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.

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. Taylor and Wedel (2013: figure 14).

So, by now, you know I’m a doofus. I have been thinking about this problem literally for years and the data I needed to address it was sitting on my hard drive the entire time. One of the things I pondered during those lost years is what the best shape for a concave-to-convex intervertebral joint might be. Would the best spacing be radially constant (A in the figure above), or antero-posteriorly constant (B), or some other, more complicated arrangement? The answer in this case surprised me–although the condyle is a lot smaller in diameter than the cotyle, the anteroposterior separation between them in almost constant, as you can see in part C of the above figure.

Figure 13. Joint between sixth and seventh cervicals vertebrae of Sauroposeidon. X-ray scout image of the C6/C7 intervertebral joint in Sauroposeidon OMNH 53062, in right lateral view. The silhouette of the condyle is traced in blue and the cotyle in red. The scale on the right is marked off in centimeters, although the numbers next to each mark are in millimeters.

Joint between sixth and seventh cervicals vertebrae of Sauroposeidon.
X-ray scout image of the C6/C7 intervertebral joint in Sauroposeidon OMNH 53062, in right lateral view. The silhouette of the condyle is traced in blue and the cotyle in red. The scale on the right is marked off in centimeters, although the numbers next to each mark are in millimeters. Taylor and Wedel (2013: figure 13).

Don’t get too worked up about that, though, because the next joint is very different! Here’s the C6/C7 joint, again in a lateral scout x-ray, with the ends of the bones highlighted. Here the condyle is almost as big in diameter as the cotyle, but it is weirdly flat. This isn’t a result of overzealous prep–most of the condyle is still covered in matrix, and I only found its actual extent by looking at the x-ray. This is flatter than most anterior dorsal vertebrae of Apatosaurus–I’ve never seen a sauropod cervical with such a flat condyle. Has anyone else?

The condyle of C6 is a bit flatter than expected, too–certainly a lot flatter than the cervical condyles in Giraffatitan and the BYU Brachiosaurus vertebrae. As we said in the paper,

It is tempting to speculate that the flattened condyles and nearly constant thickness of the intervertebral cartilage are adaptations to bearing weight, which must have been an important consideration in a cervical series more than 11 meters long, no matter how lightly built.

Anyway, obviously here the anteroposterior distance between condyle and cotyle could not have been uniform because they are such different shapes. Wacky. The zygs are missing, so they’re no help, and clearly the condyle is not centered in the cotyle. Whether this posture was attainable in life is debatable; I’ve seen some pretty weird stuff. In any case, we didn’t use this joint for estimating cartilage thickness because we had no reason to trust the results.

Figure 15. First and second dorsal vertebrae of Apatosaurus CM 3390. Articulated first and second dorsal vertebrae of Apatosaurus CM 3390. A. Digital model showing the two vertebrae in articulation, in left lateral (top) and ventral (bottom) views. B-G. Representative slices illustrating the cross-sectional anatomy of the specimen, all in posterior view. B. Slice 25. C. Slice 31. D. Slice 33. E. Slice 37. F. Slice 46. G. Slice 61. Orthogonal gaps are highlighted where the margins of the condyle and cotyle are parallel to each other and at right angles to the plane of the CT slice. 'Zygs' is short for 'zygapophyses', and NCS denotes the neurocentral synchondroses.

First and second dorsal vertebrae of Apatosaurus CM 3390.
Articulated first and second dorsal vertebrae of Apatosaurus CM 3390. A. Digital model showing the two vertebrae in articulation, in left lateral (top) and ventral (bottom) views. B-G. Representative slices illustrating the cross-sectional anatomy of the specimen, all in posterior view. B. Slice 25. C. Slice 31. D. Slice 33. E. Slice 37. F. Slice 46. G. Slice 61. Orthogonal gaps are highlighted where the margins of the condyle and cotyle are parallel to each other and at right angles to the plane of the CT slice. ‘Zygs’ is short for ‘zygapophyses’, and NCS denotes the neurocentral synchondroses. Taylor and Wedel (2013: figure 15).

Kent Sanders and I had also scanned several of the smaller sauropod vertebrae from the Carnegie collection (basically, the ones that would fit in the trunk of my car for the drive back to Oklahoma). Crucially, we’d scanned a couple of sets of articulated vertebrae, CM 3390 and CM 11339, both from juvenile individuals of Apatosaurus. In both cases, the condyles and cotyles are concentric (that’s what the ‘orthogonal gaps’ are all about in the above figure) and the zygs are in articulation, just as in Sauroposeidon. These are dorsals, so we don’t have any cervical ribs here to provide a third line of evidence that the articulation is legit, but all of the evidence that we do have is at least consistent with that interpretation.

So, here’s an interesting thing: in CM 3390, above, the first dorsal is cranked up pretty sharply compared to the next one, but the condyle is still centered in the cotyle and the zygs are in articulation. Now, the vertebrae have obviously been sheared by taphonomic deformation, but that seems to have affected both vertebrae to the same extent, and it’s hard to imagine some kind of taphonomic pressure moving one vertebra around relative to the next. So I think it’s at least plausible that this range of motion was achievable in life. Using various views and landmarks, we estimate the degree of extension here somewhere between 31 and 36 degrees. That’s a lot more than the ~6 degrees estimated by Stevens and Parrish (1999, 2005). And, as we mentioned in the paper, it nicely reinforces the point made by Upchurch (2000), that flexibility in the anterior dorsals should be taken into account in estimating neck posture and ROM.

Figure 16. Dorsal vertebrae of Apatosaurus CM 11339. Articulated middle or posterior dorsal vertebrae of Apatosaurus CM 11339. A. X-ray scout image showing the two vertebrae in articulation, in left lateral view. B–D. Slices 39, 43 and and 70 in posterior view, showing the most anterior appearance of the condyles and cotyles.

Dorsal vertebrae of Apatosaurus CM 11339.
Articulated middle or posterior dorsal vertebrae of Apatosaurus CM 11339. A. X-ray scout image showing the two vertebrae in articulation, in left lateral view. B–D. Slices 39, 43 and and 70 in posterior view, showing the most anterior appearance of the condyles and cotyles. Taylor and Wedel (2013: figure 16).

Here’s our last specimen, CM 11339. No big surprises here, although if you ever had a hard time visualizing how hyposphenes and hypantra fit together, you can see them in articulation in parts C and D (near the top of the specimen). Once again, by paging through slices we were able to estimate the separation between the vertebrae. Incidentally, the condyle IS centered in the cotyle here, it just doesn’t look that way because the CT slice is at an angle to the joint–see the lateral scout in part A of the figure to see what I mean.

So, what did we find? In Sauroposeidon the spacing between C5 and C6 is 52mm. That’s pretty darn thick in absolute terms–a shade over two inches–but really thin in relative terms–only a little over 4% of the length of each vertebra. In both of the juvenile Apatosaurus specimens, the spacing between the vertebrae was about 14mm (give or take a few because of the inherent thickness of the slices; see the paper for details on these uncertainties).

Now, here’s an interesting thing: we can try to estimate the intervertebral spacing in an adult Apatosaurus in two ways–by scaling up from the juvenile apatosaurus, or by scaling sideways from Sauroposeidon (since a big Apatosaurus was in the same ballpark, size-wise)–and we get similar answers either way.

Scaling sideways from Sauroposeidon (I’m too lazy to write anymore so I’m just copying and pasting from  the paper):

Centrum shape is conventionally quantified by Elongation Index (EI), which is defined as the total centrum length divided by the dorsoventral height of the posterior articular surface. Sauroposeidon has proportionally very long vertebrae: the EI of C6 is 6.1. If instead it were 3, as in the mid-cervicals of Apatosaurus, the centrum length would be 600 mm. That 600 mm minus 67 mm for the cotyle would give a functional length of 533 mm, not 1153, and 52 mm of cartilage would account for 9.8% of the length of that segment.

Scaling up from the juveniles: juvenile sauropods have proportionally short cervicals (Wedel et al. 2000). The scanned vertebrae are anterior dorsals with an EI of about 1.5. Mid-cervical vertebrae of this specimen would have EIs about 2, so the same thickness of cartilage would give 12mm of cartilage and 80mm of bone per segment, or 15% cartilage per segment. Over ontogeny the mid-cervicals telescoped to achieve EIs of 2.3–3.3. Assuming the cartilage did not also telescope in length (i.e., didn’t get any thicker than it got taller or wider), the ratio of cartilage to bone would be 12:120 (120 from 80*1.5), so the cartilage would account for 10% of the length of the segment–almost exactly what we got from the based-on-Sauroposeidon estimate. So either we got lucky here with our tiny sample size and truckloads of assumptions, or–just maybe–we discovered a Thing. At least we can say that the intervertebral spacing in the Apatosaurus and Sauroposeidon vertebrae is about the same, once the effects of scaling and EI are removed.

Finally, we’re aware that our sample size here is tiny and heavily skewed toward juveniles. That’s because we were just collecting targets of opportunity. Finding sauropod vertebrae that will fit through a medical-grade CT scanner is not easy, and it’s just pure dumb luck that Kent Sanders and I had gotten scans of even this many articulated vertebrae way back when, since at the time we were on the hunt for pneumaticity, not intervertebral joints or their soft tissues. As Mike has said before, we don’t think of this paper as the last word on anything. It is, explicitly, exploratory. Hopefully in a few years we’ll be buried in new data on in-vivo intervertebral spacing in both extant and extinct animals. If and when that avalanche comes, we’ll just be happy to have tossed a snowball.

References

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

Sauroposeidon OMNH 53062 C7-C8 left side

Sauroposeidon holotype OMNH 53062, posterior half of ?C7 and all of ?C8 in left lateral view. Scale bar is in inches.

There’s a lot more Sauroposeidon material these days than there used to be, thanks to the referral by D’Emic and Foreman (2012) of Paluxysaurus and Ostrom’s Cloverly material and the new Cloverly material to my favorite sauropod genus. I’ve seen almost all of this material firsthand, but obviously the specimen I’m most familiar with is the holotype, OMNH 53062. It was the primary thing occupying my mind from the summer of 1996 through the spring of 2000, and it has remained a frequent object of wonder ever since.

The specimen was found lying on its right side in the field, so that side is in better shape, by virtue of having been more deeply buried and thus protected from the ravages of freezing and thawing and other erosional processes. When the jackets were taken out of the ground and prepared, the not-so-well-preserved left sides were prepped first. Then permanent support jackets were made on the left sides, the vertebrae were flipped onto their left sides, the field jackets were removed from their right sides, and the vertebrae were prepped on the right. They’ve been lying in their support jackets, left side down and right side up, ever since. (For more on the taphonomy and recovery of the specimen, see this post and Wedel and Cifelli 2005 [free PDF linked below].)

Now, if I had known what I was doing, I would have photographed the crap out of the left sides before the verts were flipped. But it was my first project and I was learning on the job, and that didn’t occur to me until later.

It also didn’t occur to me that, once flipped, the left sides would be effectively out of reach forever. But the vertebrae are extremely fragile. The bigger verts have cracks running through them, and the jackets flexed noticeably when we took them for CT scanning. I am worried that if we tried to flip the bigger verts today, they might just crumble. Even the surface bone is fragile. I remember once trying to get some dust off one of the verts with a vacuum cleaner hose, and watching in horror as some of the millimeter-thin external bone just flaked off and flew away. That was in the late 1990s, when the verts were still stored in the dusty, drafty WWII-era buildings that had housed the museum collections for ages. Now they’re in what I still think of as the “new” building, which opened in 2000, in a really nice modern collection room with climate and dust control, and I’ve never seen them with any noticeable dust.

Anyway, the left sides are now obscured by their supporting jackets and will remain that way for the foreseeable future. And I don’t have a complete set of photos of the left sides of the verts. But I do have one, of the back half of ?C7 and all of ?C8, and a scan of it appears at the top of this post. It’s a scan of a physical photograph because it was taken in late 1996 or early 1997–no-one I knew had a digital camera, and if you wanted a digital version of a photograph, you shot it on a film camera, had a big print made, and scanned that on a flatbed scanner.

Here’s another version with the vertebrae outlined:

Sauroposeidon OMNH 53062 C7-C8 left side - outlined

When I and everyone else thought that Sauroposeidon was a brachiosaur, I was pretty sure that these were C7 and C8, out of a total of 13 cervicals, just like Giraffatitan. And it still might be so–a future analysis might find that the newly-expanded Sauroposeidon is a brachiosaurid after all, and even if not, Gomani (2005) posited a primitive cervical count of 13 for titanosaurs. If that’s true, then possibly 13 cervicals are primitive for all titanosauriforms, and the increases beyond that–to 17 in Euhelopus and 14-17 in more derived titanosaurs like Futalognkosaurus and Rapetosaurus–were deviations from that primitive pattern.

But.

If Sauroposeidon was a basal somphospondyl, as posited by D’Emic and Foreman (2012) and as found in the phylogenetic analysis of D’Emic (2012), then maybe it was more like Euhelopus than Giraffatitan, and maybe it had more than 13 cervicals. (Note that D’Emic [2012] found Sauroposeidon to be a basal somphospondyl but outside the Euhelopodidae, so even in his analysis, Euhelopus could have gotten its extra cervicals independently of Sauroposeidon.) That’s an interesting prospect, since the 11.5-meter neck estimate for Sauroposeidon I made back in 2000 was based on the conservative assumption of 13 cervicals. If Sauroposeidon had more cervicals, they were probably mid-cervicals (nobody adds more dinky C3s, or stubby cervico-dorsals*–that would be silly), and therefore between 1 and 1.25 meters long. So if the individual represented by OMNH 53062 had 15 cervicals, as Mike hypothetically illustrated in this post, its neck might was probably more like 14 meters long, and if it had 17 cervicals, like Euhelopus and Rapetosaurus, its neck might have topped 16 meters–as long or longer than that of Supersaurus.

Now, I’m not saying that Sauroposeidon had a 16-meter neck. The conservative estimate is still 13 cervicals adding up to 11.5 meters. But the possibility of a longer neck is tantalizing, and can’t be ruled out based on current evidence. As usual, we need more fossils.

Happily, now that Sauroposeidon is known from Oklahoma, Texas, and Wyoming, and is one of the best-represented EKNApods instead of one of the scrappiest, the chances that we’ll find more of it–and recognize it–are looking good. I will keep my fingers firmly crossed–as they have been for the last 17 years.

* Radical pedantry note: of course we have very good evidence of sauropods getting more cervical vertebrae by recruiting dorsals into the cervical series. So, for example, 13 cervicals and 12 dorsals are supposed to be primitive for neosauropods, but diplodocids have 15 and 10, respectively–the obvious inference being that the first two dorsals got cervicalized. So in this narrow meristic sense, sauropods definitely did add cervicodorsals. But my point above is about the morphology of the verts themselves–once diplodocids had those two extra cervicals at the end, the former cervicodorsals were free to become more “cervicalized” in form. So effectively–in terms of the shapes of their necks–diplodocids added mid-cervicals.

References

DEmic 2012 figure 5 titanosauriform phylogeny

D’Emic (2012: figure 5)

Now this is super-freakin’ cool, and I’ve been meaning to blog about it for a while. In Mike D’Emic’s recent titanosauriform phylogeny (D’Emic 2012), he (correctly) included Brachiosaurus and Giraffatitan as separate OTUs, and, hey, whaddayaknow, they’re not sister taxa anymore: Brachiosaurus is more closely related to a trio of Early Cretaceous North American brachiosaurids than it is to Giraffatitan.

The potential for someone to find this result was there ever since Mike broke Brachiosaurus and Giraffatitan apart, as a previously composite OTU, in his 2009 paper. It just hadn’t materialized. In fact, some authors have gone out of their way to not find this out, by keeping the old composite coding. That seems…unwise, in retrospect. Whether one agreed with Mike on the nomenclatural point of generic separation or not, not coding the two taxa as separate OTUs (especially after Mike had done that work for them) was a poor phylogenetic decision–in essence, it constrained Brachiosaurus and Giraffatitan to be sister taxa in the analysis, and outlawed any more interesting results–like the one obtained by D’Emic (2012)–before the software even started crunching trees.

So anyway, back to the coolness inherent in D’Emic’s tree. Of course, like all phylogenetic results this is just a hypothesis and it is subject to revision based on new information blah blah blah…but it is really interesting that there is now some phylogenetic support for an endemic radiation of brachiosaurids in North America (bonus goofy observation–you can’t spell ‘endemic’ without D’Emic). Or perhaps Lauriasia–I would kill to know where the British brachiosaurids (or basal titanosauriforms) fit into this story, and Lusotitan, and the apparently tiny Croatian carbonate platform brachiosaurs.

Also super-interesting that, if this tree is accurate, these endemic Early Cretaceous brachiosaurids were living alongside a giant basal somphospondyl in the form of Sauroposeidon, which came from heaven knows where. Look who it’s surrounded by–Ligabuesaurus is from Argentina, Tastavinsaurus is from Spain, and the euhelopodids are from eastern Asia. Evidently there was also a global radiation of basal somphospondyls. And why are all the Early Cretaceous North American brachiosaurids small–smaller than Brachiosaurus and Giraffatitan, anyway (at least until we find bigger individuals of the former)–while Sauroposeidon is so big? Or is that just an effect of tiny sample sizes, and one lucky strike in the form of the Sauroposeidon holotype?

So much cool stuff to think about. I don’t usually get this much enjoyment out of a tree unless it has lights and ornaments.

References

Follow

Get every new post delivered to your Inbox.

Join 414 other followers