[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.]

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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!”

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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.

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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

A few bits and pieces about the PLOS Collection on sauropod gigantism that launched yesterday.

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First, there’s a nice write-up of one of our papers (Wedel and Taylor 2013b on pneumaticity in sauropod tails) in the Huffington Post today. It’s the work of PLOS blogger Brad Balukjian, a former student of Matt’s from Berkeley days. The introduction added by the PLOS blogs manager is one of those where you keep wanting to interrupt, “Well, actually it’s not quite like that …” but the post itself, once it kicks in, is good. Go read it.

Brad also has a guest-post on Discover magazine’s Crux blog: How Brachiosaurus (and Brethren) Became So Gigantic. He gives an overview of the sauropod gigantism collection as a whole. Well worth a read to get your bearings on the issue of sauropod gigantism in general, and the new collection in particular.

PLOS’s own community blog EveryONE also has its own brief introduction to the collection.

And PLOS and PeerJ editor Andy Farke, recently in these pages because of his sensational juvenile Parasaurolophus paper, contributes his own overview of the collection, How Big? How Tall? And…How Did It Happen?

Finally, if you’re at SVP, go and pick up your free copy of the collection. Matt was somehow under the impression that the PLOS USB drives with the sauropod gigantism collection would be distributed with the conference packet when people registered. In fact, people have to go by the PLOS table in the exhibitor area (booth 4 in the San Diego ballroom) to pick them up. There are plenty of them, but apparently a lot of people don’t know that they can get them.

References

This is an exciting day: the new PLOS Collection on sauropod gigantism is published to coincide with the start of this year’s SVP meeting! Like all PLOS papers, the contents are free to the world: free to read and to re-use.  (What is a Collection? It’s like an edited volume, but free online instead of printed on paper.)

There are fourteen papers in the new Collection, encompassing neck posture (yay!), nutrition (finally putting to bed the Nourishing Vomit Of Eucamerotus hypothesis), locomotion, physiology and evolutionary ecology. Lots every sauropod-lover to enjoy.

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Taylor and Wedel (2013c: 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.

Matt and I are particularly excited that we have two papers in this collection: Taylor and Wedel (2013c) on intervertebral cartilage in necks, and Wedel and Taylor (2013b) on pneumaticity in the tails of (particularly) Giraffatitan and Apatosaurus. So we have both ends of the animal covered. It also represents a long-overdue notch on our bed-post: for all our pro-PLOS rhetoric, this is the first time either of has had a paper published in a PLOS journal.

Wedel and Taylor (2013b: Figure 4). Giraffatitan brancai tail MB.R.5000 (‘Fund no’) in right lateral view. Dark blue vertebrae have pneumatic fossae on both sides, light blue vertebrae have pneumatic fossae only on the right side, and white vertebrae have no pneumatic fossae on either side. The first caudal vertebra (hatched) was not recovered and is reconstructed in plaster.

It’s a bit of a statistical anomaly that after a decade of collaboration in which there was never a Taylor & Wedel or Wedel & Taylor paper, suddenly we have five of them out in a single year (including the Barosaurus preprint, which we expect to eventually wind up as Taylor and Wedel 2014). Sorry about the alphabet soup.

Since Matt is away at SVP this week, I’ll be blogging mostly about the Taylor and Wedel paper this week. When Matt returns to civilian life, the stage should be clear for him to blog about pneumatic caudals.

Happy days!

References

Yesterday I announced that our new paper on Barosaurus was up as a PeerJ preprint and invited feedback.

I woke up this morning to find its third substantial review waiting for me.

That means that this paper has now accumulated as much useful feedback in the twenty-seven hours since I submitted it as any previous submission I’ve ever made.

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Taylor and Wedel (2013b: figure 7). Barosaurus lentus holotype YPM 429, Vertebra S (C?12). Left column from top to bottom: dorsal, right lateral and ventral views; right column: anterior view. Inset shows displaced fragment of broken prezygapophysis. Note the narrow span across the parapophyses in ventral view, and the lack of damage to the ventral surface of the centrum which would indicate transverse crushing.

It’s worth reviewing the timeline here:

  • Monday 23rd September, 1:19 am: I completed the submission process.
  • 7:03 am: the preprint was published. It took less than six hours.
  • 10:52 am: received a careful, detailed review from Emanuel Tschopp. It took less than four hours from publication, and so of course less than ten from submission.
  • About 5:00 pm: received a second review, this one from Mark Robinson. (I don’t know the exact time because PeerJ’s page doesn’t show an actual timestamp, just “21 hours ago”.)
  • Tuesday 24th September, about 4:00 am: received a third review, this from ceratopsian-jockey and open-science guru Andy Farke.

Total time from submission to receiving three substantial reviews: about 27 hours.

It’s worth contrasting that with the times taken to get from submission to the receipt of reviews — usually only two of them — when going through the traditional journal route. Here are a few of mine:

  • Diplodocoid phylogenetic nomenclature at the Journal of Paleontology, 2004-5 (the first reviews I ever received): three months and 14 days.
  • Revised version of the same paper at PaleoBios, 2005 (my first published paper): one month and 10 days.
  • Xenoposeidon description at Palaeontology, 2006: three months and 19 days, although that included a delay as the handling editor sent it to a third, tie-breaking, reviewer.
  • Brachiosaurus revision at the Journal of Vertebrate Paleontology, 2008: one month and 11 days.
  • Sauropod neck anatomy (eventually to be published in a very different form in PeerJ) at Paleobiologyfive months and two days.
  • Trivial correction to the Brachiosaurus revision at the Journal of Vertebrate Paleontology, 2010: five months and 11 days, bizarrely for a half-page paper.

Despite the wide variations in submission-to-review time at these journals, it’s clear that you can expect to wait at least a month before getting any feedback at all on your submission at traditional journals. Even PeerJ took 19 days to get the reviews of our neck-anatomy paper back to us.

So I am now pretty such sold on the pre-printing route. As well as getting this early version of the paper out there early so that other palaeontologists can benefit from it (and so that we can’t be pre-emptively plagiarised), issuing a preprint has meant that we’ve got really useful feedback very quickly.

I highly recommend this route.

By the way, in case anyone’s wondering, PeerJ Preprints is not only for manuscripts that are destined for PeerJ proper. They’re perfectly happy for you to use their service as a place to gather feedback for your work before submitting it elsewhere. So even if your work is destined for, say, JVP, there’s a lot to be gained by preprinting it first.

I was very pleased, on checking my email this morning, to see that my and Matt’s new paper, The neck of Barosaurus was not only longer but also wider than those of Diplodocus and other diplodocines, is now up as a PeerJ preprint!

Figure6-vertebra-q-composite

Taylor and Wedel (2013b: figure 6). Barosaurus lentus holotype YPM 429, Vertebra Q (C?13). Top row: left ventrolateral view. Middle row, from left to right: anterior view, with ventral to the right; ventral view; posterior view, with ventral to the left. Bottom row: right lateral view, inverted. Inset shows diapophyseal facet on right side of vertebra, indicating that the cervical ribs were unfused in this individual despite its great size. Note the broad, flat prezygapophyseal facet visible in anterior view.

I was pleased partly because of the very quick work on PeerJ’s part. I submitted the preprint at 1:22am last night, then went to bed. Almost immediately I got an automatic email from PeerJ saying:

Thank you for submitting your manuscript, “The neck of Barosaurus was not only longer but also wider than those of Diplodocus and other diplodocines” (#2013:09:838:0:0:CHECK:P) – it has now been received by PeerJ PrePrints.

Next, it will be checked by PeerJ staff, who will notify you if any alterations are required to the manuscript or accompanying files.

If the PrePrint successfully passes these checks, it will be made public.

You will receive notification by email at each stage of this process; you can also check the status of your manuscript at any time.

Lots to like here: the quickness of the response, the promise of automatic email updates, and the one-click link to check on progress (as opposed to the usual maze of Manuscript Central options to navigate).

Sure enough, a couple of hours later the next automatic email arrived, telling me that Matt had accepted PeerJ’s email invitation to be recognised as the co-author of the submission.

And one hour ago, just as I was crawling out of bed, I got the notification that the preprint is up. That simple.

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Taylor and Wedel (2013b: Figure 9). Partial reconstruction of the Barosaurus lentus holotype YPM 429, cervical vertebra R, approximating its undamaged state by allowing for dorsoventral crushing, shearing and loss of some extremities. Anterior and posterior views scaled to 125% of uncorrected width and 80% of uncorrected height. Dorsal view scaled to 80% of uncorrected height; condyle moved forward and cotyle scaled to 50% of uncorrected width to allow for shearing. Lateral view scaled to 125% of uncorrected height, and sheared backwards 15 degrees. Metapophyses and postzygapophyses drawn in multiple views based on vertebrae Q and S and AMNH 6341 material.

I’m also pleased because we managed to get this baby written so quickly. It started life as our talk at SVPCA in Edinburgh (Taylor and Wedel 2013a), which we delivered 25 days ago having put it together mostly in a few days running up to the conference — so it’s zero to sixty in less than a month. Every year we promise ourselves that we’ll write up our talks, and we never seem to get around to it, but this year I started writing on the train back from Edinburgh. By the time I got home I had enough of a hunk of text to keep me working on it, and so we were able to push through in what, for us, is record time.

Now here’s what we’d like:

We want this paper’s time as a preprint to be time well spent — which means that we want to improve it. To do that, we need your reviews. Assuming we get some useful comments, we plan to release an updated version pretty soon; and after some number of iterations, we’ll submit the resulting paper as a full-fledged PeerJ paper.

So if you know anything about sauropods, about vertebra, about deformation, about ecology, or even about grammar or punctuation, please do us a favour: read the preprint, then get over to its PeerJ page and leave your feedback. You’ll be helping us to improve the scientific record. We’ll acknowledge substantial comments in the final paper, but even the pickiest comments are appreciated.

Because we want to encourage this approach to bringing papers to publication, we’d ask you please do not post comments about the paper here on SV-POW!. Please post them on the PeerJ preprint page. We’ve leaving comments here open for discussion of the preprinting processes, but not the scientific content.

References

  • Taylor, Michael P., and Mathew J. Wedel. 2013a. Barosaurus revisited: the concept of Barosaurus (Dinosauria: Sauropoda) is based on erroneously referred specimens. (Talk given as: Barosaurus revisited: the concept of Barosaurus (Dinosauria: Sauropoda) is not based on erroneously referred specimens.) pp. 37-38 in Stig Walsh, Nick Fraser, Stephen Brusatte, Jeff Liston and Vicen Carrió (eds.), Programme and Abstracts, 61st Symposium on Vertebrae Palaeontology and Comparative Anatomy, Edinburgh, UK, 27th-30th August 2013. 33 pp.
  • Taylor, Michael P., and Mathew J. Wedel. 2013b. The neck of Barosaurus was not only longer but also wider than those of Diplodocus and other diplodocines. PeerJ PrePrints 1:e67v1 http://dx.doi.org/10.7287/peerj.preprints.67v1

Let’s take another look at that Giraffatitan cervical. MB.R.2180:C5, from a few days ago:

FigureA-Giraffatitan-SI-C5

That’s a pretty elongate vertebra, right? But how elongate, exactly? How can we quantify whether it’s more or less elongate than some other vertebra?

The traditional answer is that we quantify elongation using the elongation index, or EI. This was originally defined by Upchurch (1998:47) as “the length of a vertebral centrum divided by the width across its caudal face”. Measuring from the full-resolution version of the image above, I make that 1779/529 pixels, or 3.36.

But then those doofuses Wedel et al. (2000:346) came along and said:

When discussing vertebral proportions Upchurch (1998) used the term elongation index (EI), defined as the length of the centrum divided by the width of the cotyle. Although they did not suggest a term for the proportion, Wilson & Sereno (1998) used centrum length divided by the height of the cotyle as a character in their analysis. We prefer the latter definition of this proportion, as the height of the cotyle is directly related to the range of motion of the intervertebral joint in the dorsoventral plane. For the purposes of the following discussion, we therefore redefine the EI of Upchurch (1998) as the anteroposterior length of the centrum divided by the midline height of the cotyle.

Since then, the term EI has mostly been used in this redefined sense — but I think we all agree now that it would have been better for Wedel et al to have given a new name to Wilson and Sereno’s ratio rather than apply Upchurch’s name to it.

Aaaanyway, measuring from the image again, I give that vertebra an EI (sensu Wedel et al. 2000) of 1779/334 = 5.33. Which is 58% more elongate than when using the Upchurch definition! This of course follows directly from the cotyle being 58% wider than tall (529/334 pixels).

So one of principal factors determining how elongate a vertebra seems to be is the shape of its cotyle. And that’s troublesome, because the cotyle is particularly subject to crushing — and it’s not unusual for even consecutive vertebrae from the same column to be crushed in opposite directions, giving them (apparently) wildly different EIs.

Here’s an example (though not at all an extreme one): cervicals 4 and 6 of the same specimen, MB.R.2180 (formerly HM SI), as the multi-view photo above:

DSCN5527-5535-SI-c4-and-c6-posterior

Measuring from the photos as before, I make the width:height ratio of C4 683/722 pixels = 0.95, and that of C6  1190/820 pixels = 1.45. So these two vertebrae — from the same neck, and with only one other vertebrae coming in between them — differ in preserved cotyle shape by a factor of 1.53.

And by the way, this is one of the best preserved of all sauropod neck series.

Let’s take a look at the canonical well-preserved sauropod neck: the Carnegie Diplodocus, CM 84. Here are the adjacent cervicals 13 and 14, in posterior view, from Hatcher (1901: plate VI):

Hatcher1901-plate-VI-C13-C14-posterior

For C14 (on the left), I get a width:height ratio of 342/245 pixels = 1.40. For C13 (on the right), I get 264/256 pixels = 1.03. So C14 is apparently 35% broader than its immediate predecessor. I absolutely don’t buy that this represents how the vertebrae were in life.

FOR EXTRA CREDIT: what does this tell us about the reliability of computer models that purport to tell us about neck posture and flexibility, based on the preserved shapes of their constituent vertebrae?

So what’s to be done?

The first thing, as always in science, is to be explicit about what statements we’re making. Whenever we report an elongation index, we need to clearly state whether it’s EI sensu Upchurch 1998 or EI sensu Wedel et al. 2000. Since that’s so cumbersome, I’m going propose that we introduce two new abbreviations: EIH (Elongation Index Horizonal), which is Upchurch’s original measure (length over horizontal width of cotyle) and EIV (Elongation Index Vertical), which is Wilson and Sereno’s measure (length over vertical height of cotyle). If we’re careful to report EIH and EIV (or better still both) rather than an unspecified EI, then at least we can avoid comparing apples with oranges.

But I think we can do better, by combining the horizontal and vertical cotyle measurements in some way, and dividing the length by the that composite. This would give us an EIA (Elongation Index Average), which we could reasonably expect to preserve the original cotyle size, and so to give a more reliable indication of “true” elongation.

The question is, how to combine the cotyle width and height? There are two obvious candidates: either take the arithmetic mean (half the sum) or the geometric mean (the square root of the product). Note that for round cotyles, both these methods will give the same result as each other and as EIH and EIV — which is what we want.

Which mean should we use for EIA? to my mind, it depends which is best preserved when a vertebra is crushed. If a 20 cm circular cotyle is crushed vertically to 10cm, does it tend to smoosh outwards to 30 cm (so that 10+30 = the original 20+20) or to 40 cm (so that 10 x 40 = the original 20 x 20)? If the former, then we should use arithmetic mean; if the latter, then geometric mean.

Does anyone know how crushing works in practice? Which of these models most closely approximates reality? Or can we do better than either?

Update (8:48am): thanks for Emanuel Tschopp for pointing out (below) what I should have remembered: that Chure et al.’s (2010) description of Abydosaurus introduces “aEI”, which is the same as one of my proposed definitons of EIA. So we should ignore the last four paragraphs of this post and just use aEI. (Their abbreviation is better, too.)

 

References

  • Hatcher, Jonathan B. 1901. Diplodocus (Marsh): its osteology, taxonomy and probable habits, with a restoration of the skeleton. Memoirs of the Carnegie Museum 1:1-63 and plates I-XIII.
  • Upchurch, Paul. 1998. The phylogenetic relationships of sauropod dinosaurs. Zoological Journal of the Linnean Society 124:43-103.
  • Wedel, Mathew J., Richard L. Cifelli and R. Kent Sanders. 2000b. Osteology, paleobiology, and relationships of the sauropod dinosaur Sauroposeidon. Acta Palaeontologica Polonica 45(4):343-388.
  • Wilson, J. A. and Paul C. Sereno. 1998. Early evolution and higher-level phylogeny of sauropod dinosaurs. Society of Vertebrate Paleontology, Memoir 5:1-68.

Apatosaurus1B

We’ve blogged a lot of Bob Nicholls‘ art (here, here, and here) and we’ll probably continue to do so for the foreseeable future. We don’t have much choice: he keeps drawing awesome things and giving us permission to post them. Like this defiantly shaggy Apatosaurus, which was probably the star of the Morrison version of Duck Dynasty. Writes Bob:

On my way home at the airport I did a sketch of your giant Apatosaurus* – see attachment.  My thought was that massive thick necks were probably pretty sexy things to apatosaurs, so maybe sexually mature individuals used simple feathers (stage 1, 2 or 3?) to accentuate the neck profile.  The biggest males would of course have the most impressive growths so in the attached sketch your giant has one of the biggest beards in Earth’s history!  What do you think of this idea?

Well, I think it’s awesome. And entirely plausible, for reasons already explained in this post.

“Now, wait,” you may be thinking, “I thought you guys said that sauropod necks weren’t sexually selected.” Actually we made a slightly different point: that the available evidence does not suggest that sexual selection was the primary driver of sauropod neck elongation. But we also acknowledged that biological structures are almost never single-purpose, and although the long necks of sauropods probably evolved to help them gather more food, there is no reason that long necks couldn’t have been co-opted as social billboards. This seems especially likely in Apatosaurus, where the neck length is unremarkable** but the neck fatness is frankly bizarre (and even inspired a Star Wars starfighter!).

I also love the “mobile ecosystem” of birds, other small dinosaurs, and insects riding on this Apatosaurus or following in its train. It’s a useful reminder that we have no real idea what effect millions of sauropods would have on the landscape. But it’s not hard to imagine that most Mesozoic terrestrial ecosystems were sauropod-driven in a thousand cascading and ramifying chains of cause and effect. I’d love to know how that worked. At heart, I’m still a wannabe chrononaut, and all my noodlings on pneumaticity and sauropod nerves and neural spines and so on are just baby steps toward trying to understand sauropod lives. Safari by way of pedantry: tally-ho!

For other speculative apatosaurs, see:

* “My” giant is the big Oklahoma Apatosaurus, which I gave a talk on at SVPCA a couple of weeks ago. See these posts for more details (123).

** Assuming we can be blasé about a neck that is more than twice as long (5 m) as a world-record giraffe neck (2.4 m), for garden variety Apatosaurus, or three times that length for the giant Oklahoma Apatosaurus (maybe 7 m).

OMNH baby Apatosaurus

I was at the Oklahoma Museum of Natural History in March to look at their Apatosaurus material, so I got to see the newly-mounted baby apatosaur in the “Clash of the Titans” exhibit (more photos of that exhibit in this post). How much of this is real (i.e., cast from real bones, rather than sculpted)? Most of the vertebral centra, a few of the neural arches, some of the limb girdle bones, and most of the long bones of the limbs. All of the missing elements–skull, neural arches, ribs, appendicular bits–were sculpted by the OMNH head preparator, Kyle Davies. Kyle is one of those frighteningly talented people who, if they don’t have what they need, will just freaking build it from scratch. Over the years he has helped me out a LOT with the OMNH sauropod material–including building a clamshell storage jacket for the referred scapula of Brontomerus so we could photograph it from the lateral side–so it’s about time I gave him some props.

Atlas-axis model with Kyle

Case in point: this sweet atlas-axis complex that Kyle sculpted for the juvenile Apatosaurus mount.

Atlas-axis model by Kyle Davies

Most fish, amphibians, and other non-amniote tetrapods only have a single specialized vertebra for attaching to the skull. But amniotes have two: a ring- or doughnut-shaped first cervical vertebra (the atlas) that articulates with the occipital condyle(s) of the skull, and a second cervical vertebra (the axis) that articulates with the atlas and sometimes with the skull as well. Mammals have paired occipital condyles on the backs or bottoms of our skulls, so our skulls rock up and down on the atlas (nodding “yes” motion), and our skull+atlas rotates around a peg of bone on the axis called the odontoid process or dens epistrophei (shaking head “no” motion). As shown in the photos and diagrams below, the dens of the axis is actually part of the atlas that fuses to the second vertebra instead of the first. Also, reptiles, including dinosaurs and birds, tend to have a single ball-shaped occipital condyle that fits into the round socket formed by the atlas, so their “yes” and “no” motions are less segregated by location.

Anyway, the whole shebang is often referred to as the atlas-axis complex, and that’s the reconstructed setup for a baby Apatosaurus in the photo above.  In addition to making a dull-colored one for the mount, Kyle made this festive version for the vert paleo teaching collection. Why so polychromatic?

Atlas-axis model key

Because in fact he built two: the fully assembled one two photos above, and a completely disassembled one, some of which is shown in this photo (I had to move the bigger bits out of the tray so they wouldn’t block the key card at the back). I originally composed this post as a tutorial. But frankly, since Kyle did all of the heavy lifting of (a) making the thing in the first place, (2) making a color-coded key to it, and (d) giving me permission to post these photos, it would be redundant to walk through every element. So think of this as a self-study rather than a tutorial.

Atlas-axis model by Kyle Davies - labeled

Oh, all right, here’s a labeled version. Note that normally in an adult animal the single piece of bone called the atlas would consist of the paired atlas neural arches (na1) and single atlas intercentrum (ic1), and would probably have a pair of fused cervical ribs (r1). Everything else would be fused together to form the axis, including the atlas pleurocentrum (c1), which forms the odontoid process or dens epistrophei (etymologically the “tooth” of the axis).

Romer 1956 fig 119 atlas-axis complex

Here’s the complete Romer (1956) figure from the key card, with a mammalian atlas-axis complex  for comparison. Incidentally, the entire book this is drawn from, Osteology of the Reptiles, is freely available online.

Apatosaurus axis-atlas complex Gilmore 1936 figs 5 and 6

And here’s the complete Gilmore (1936) figure. Sorry for the craptastic scan–amazingly, this one is NOT freely available online as far as I can tell, and Mike and I have been trying to get good scans of the plates for years. Getting back on topic, single-headed atlantal cervical ribs have been found in several sauropods, especially Camarasaurus where several examples are known, so they were probably a regular feature, even though they aren’t always preserved.

Also, as noted in this post, it is odd that in this specimen of Apatosaurus the cervical ribs had not fused to the first two vertebrae, even though they normally do, and despite the fact that the vertebrae had fused to each other, even though they normally don’t. Further demonstration, if any were needed, that sauropod skeletal fusions were wacky.

Varanops atlas-axis complex Campione and Reisz 2011 fig 2C3

For comparison to the above images, here is the atlas-axis complex in the synapsid Varanops, from Campione and Reisz (2011: fig. 2C).

Those proatlas thingies are present in some sauropods, but that’s about all I know about them, so I’ll say no more for now.

There is a good overview of the atlas-axis complex with lots of photos of vertebrae of extant animals on this page.

Previous SV-POW! posts dealing with atlantes and axes (that’s right) include:

References

IMG_0465

Those familiar with Lull (1919: plate II: figure 2) will recognise this as “vertebra Q” of the Barosarus lentus holotype YPM 429, in ventral view.

Stay tuned for more exciting Barosaurus-related news!

References

Lull, R. S. 1919. The sauropod dinosaur Barosaurus Marsh. Memoirs of the Connecticut Academy of Arts and Sciences 6:1-42 and plates I-VII.

What is an ad-hominem attack?

September 4, 2013

I recently handled the revisions on a paper that hopefully will be in press very soon. One of the review comments was “Be very careful not to make ad hominem attacks”.

I was a bit surprised to see that — I wasn’t aware that I’d made any — so I went back over the manuscript, and sure enough, there were no ad homs in there.

There was criticism, though, and I think that’s what the reviewer meant.

Folks, “ad hominem” has a specific meaning. An “ad hominem attack” doesn’t just mean criticising something strongly, it means criticising the author rather than the work. The phrase is Latin for “to the man”. Here’s a pair of examples:

  • “This paper by Wedel is terrible, because the data don’t support the conclusion” — not ad hominem.
  • “Wedel is a terrible scientist, so this paper can’t be trusted” – ad hominem.

What’s wrong with ad hominem criticism? Simply, it’s irrelevant to evaluation of the paper being reviewed. It doesn’t matter (to me as a scientist) whether Wedel strangles small defenceless animals for pleasure in his spare time; what matters is the quality of his work.

Note that ad hominems can also be positive — and they are just as useless there. Here’s another pair of examples:

  • “I recommend publication of Naish’s paper because his work is explained carefully and in detail” — not ad hominem.
  • “I recommend publication of Naish’s paper because he is a careful and detailed worker” — ad hominem.

It makes no difference whether Naish is a careful and detailed worker, or if he always buys his wife flowers on their anniversary, or even if he has a track-record of careful and detailed work. What matters is whether this paper, the one I’m reviewing, is good. That’s all.

As it happens the very first peer-review I ever received — for the paper that eventually became Taylor and Naish (2005) on diplodocoid phylogenetic nomenclature — contained a classic ad hominem, which I’ll go ahead and quote:

It seems to me perfectly reasonable to expect revisers of a major clade to have some prior experience/expertise in the group or in phylogenetic taxonomy before presenting what is intended to be the definitive phylogenetic taxonomy of that group. I do not wish to demean the capabilities of either author – certainly Naish’s “Dinosaurs of the Isle of Wight” is a praiseworthy and useful publication in my opinion – but I question whether he and Taylor can meet their own desiderata of presenting a revised nomenclature that balances elegance, consistency, and stability.

You see what’s happening here? The reviewer was not reviewing the paper, but the authors. There was no need for him or her to question whether we could meet our desiderata: he or she could just have read the manuscript and found out.

(Happy ending: that paper was rejected at the journal we first sent it to, but published at PaleoBios in revised form, and bizarrely is my equal third most-cited paper. I never saw that coming.)

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