September 14, 2015
We’ve noted that the Taylor et al. SVPCA abstract and talk slides are up now up as part of the SVPCA 2015 PeerJ Collection, so anyone who’s interested has probably taken a look already to see what it was about. (As an aside, I am delighted to see that two more abstracts have been added to the collection since I wrote about it.)
It was my privilege to present a talk on our hypothesis that the distinctive and bizarre toblerone-shaped necks of apatosaurs were an adaptation for intraspecific combat. This talk was based on an in-progress manuscript that Matt is lead-authoring. Also on board is the third SV-POW!sketeer, the silent partner, Darren Naish; and artist/ethologist Brian Engh.
Here is our case, briefly summarised from five key slides. First, let’s take a look at what is distinctive in the morphology of apatosaur cervicals:
Here I’m using Brontosaurus, which is among the more extreme apatosaurs, but the same features are seen developed to nearly the same extent in Apatosaurus louisae, the best-known apatosaur, and to some extent in all apatosaurs.
Now we’ll look at the four key features separately.
First, the cervicals ribs of sauropods (and other saurischians, including birds) anchored the longus colli ventralis and flexor colli lateralis muscles — ventral muscles whose job is to pull the neck downwards. By shifting the attachments points of these muscles downwards, apatosaurs enabled them to work with improved mechanical advantage — that is, to bring more force to bear.
Second, by redirecting the diapophyses and parapophyses ventrally, and making them much more robust than in other sauropods, apatosaurs structured their neck skeletons to better resist ventral impacts.
Third, because the low-hanging cervical ribs created an inverted “V” shape below the centrum, they formed a protective cradle for the vulnerable soft-tissue that is otherwise exposed on the ventral aspect of the neck: trachea, oesophagus, major blood vessels. In apatosaurus, all of these would have been safely wrapped in layers of connective tissue and bubble-wrap-like pneumatic diverticula. The presence of diverticula ventral to the vertebral centrum is not speculative – most neosauropods have fossae on the ventral surfaces of their cervical centra, and apatosaurines tend to have foramina that connect to internal chambers as well (see Lovelace et al. 2007: fig. 4, which is reproduced in this post).
Fourth, most if not all apatosaurs have distinctive ventrally directed club-like processes on the front of their cervical ribs. (It’s hard to tell with Apatosaurus ajax, because the best cervical vertebra of that species is so very reconstructed.) How did these appear in life? It’s difficult to be sure. They might have appeared as a low boss; or, as with rhinoceros horns, they might even have carried keratinous spikes.
Putting it all together, we have an animal whose neck can be brought downwards with great force; whose neck was mechanically capable of resisting impacts on its ventral aspect; whose vulnerable ventral-side soft-tissue was well protected; and which probably had prominent clubs or spikes all along the ventral aspect of the neck. And all of this was accomplished at the cost of making the neck a lot heavier than it would have been otherwise. Off the cuff, it seems likely that the cervical series alone would have massed twice as much in apatosaurines as in diplodocines of the same neck length.
Doubling the mass of the neck is a very peculiar thing for a sauropod lineage to do – by the Late Jurassic, sauropods were the leading edge of an evolutionary trend to lengthen and lighten the neck that had been running for almost 100 million years, through basal ornithodirans, basal dinosauromorphs, basal saurischians, basal sauropodomorphs, and basal sauropods. Whatever the selective pressures that led apatosaurines to evolve such robust and heavy necks, they must have been compelling.
The possibility that apatosaurs were pushing or crashing their necks ventrally in some form of combat accounts for all of the weird morphology documented above, and we know that sexual selection is powerful force that underlies a lot of bizarre structures in extant animals, and probably in extinct ornithodirans as well (see Hone et al. 2012, Hone and Naish 2013).
What form of combat, exactly? There are various possibilities, which we’ll discuss another time. But I’ll leave you with Brian Engh’s beautiful illustration of one possible form of combat: a powerful impact of one neck brought down onto the dorsal aspect of another.
We’re aware that this proposal is necessarily somewhat speculative. But we’re just not able to see any other explanation for the distinctive apatosaur neck. Even if we’re wrong about the ventrolateral processes on the cervical ribs supporting bosses or spikes, the first three points remain true, and given how they fly in the face of sauropods’ long history of making their necks lighter, they fairly cry out for explanation. If anyone has other proposals, we’ll be happy to hear them.
- Hone, D. W., Naish, D., & Cuthill, I. C. (2012). Does mutual sexual selection explain the evolution of head crests in pterosaurs and dinosaurs?. Lethaia 45(2):139-156.
- Hone, D. W. E., & Naish, D. (2013). The ‘species recognition hypothesis’ does not explain the presence and evolution of exaggerated structures in non‐avialan dinosaurs. Journal of Zoology 290(3):172-180.
- Lovelace, D. M., Hartman, S. A., & Wahl, W. R. (2007). Morphology of a specimen of Supersaurus (Dinosauria, Sauropoda) from the Morrison Formation of Wyoming, and a re-evaluation of diplodocid phylogeny. Arquivos do Museu Nacional, Rio de Janeiro 65(4):527-544.
September 22, 2014
We have good descriptions of the proximal parts of the cervical ribs for lots of sauropods. We also have histological cross-sections of a few, mostly thanks to the work of Nicole Klein and colleagues (Klein et al. 2012, Preuschoft and Klein 2013), although histological cross-sections of ribs were also figured as long ago as 1999, by Dalla Vecchia (1999: figs. 29 and 30), and as recently as this month, by Lacovara et al. (2014: supplementary figure 4).
What we have very, very few of is series of cross-sections that show how the cr0ss-section of a cervical rib changes along its length. There may be more out there (and if I have forgotten any, please remind me!), but at the moment I can only think of three such figures: two in Janensch (1950: figs. 83 and 85), both on Giraffatitan, and one in Klein et al. (2012: fig. 1), with cross-sections from Mamenchisaurus, Giraffatitan, and Diplodocus (shown at the top of the post).
Rarer still are images that show cross-sections of overlapped cervical ribs, stacked in situ. You could use the information in Janensch (1950: figs. 83 and 85) to generate the stacked cross-sections, but you wouldn’t know the spacing between the ribs as they were in the ground. I think the image just above, of the cervical rib bundles in the Sauroposeidon holotype, OMNH 53062, may be the first of its kind–again, if you know of any others, please let me know. I took the notes for this figure back in 2004, sitting down with the holotype and some digital calipers to make sure I could scale everything correctly, I just hadn’t ever put it into a presentable form until now. The first C6 section (blue V-shape) is from right at the root where the capitulum and tuberculum meet and the posterior shaft of the rib begins.
It is by now well-understood that the long cervical ribs of sauropods and other dinosaurs are ossified tendons of the long hypaxial neck muscles, specifically the longus colli ventralis and flexor colli lateralis. We argued this back in 200o on comparative anatomical grounds (Wedel et al. 2000b: pp. 378-379), and it has now been demonstrated histologically (Klein et al. 2012, Lacovara et al. 2014). The system of stacked tendons is also found in most birds. Here’s the bundle of stacked tendons in a rhea neck, only slightly fanned out:
And the same neck, with both the epaxial and hypaxial muscles more fully separated:
What I’d really like is an MRI of a rhea or ostrich neck, showing the stacked tendons and their associated belts of muscle, to compare with the stacked cervical ribs of Sauroposeidon and other sauropods. Anyone know of any?
Incidentally, I think the cervical ribs and cervical rib bundles of sauropods are one line of evidence for sauropod necks having been rather slenderly-muscled. The long, multi-segment muscles like the longus colli ventralis are the outermost components of the muscular envelope that surrounds the vertebrae, as you can see in the rhea dissection photos. In sauropod specimens with articulated cervical ribs, the ribs do not deviate from one another or fan out. Rather, they lie in vertically stacked bundles that run from one capitulum-tuberculum intersection to the next. So the depth of that intersection–the “root” of the cervical rib of any given vertebra–plus the thickness of the ribs stacked underneath it, is pretty much the thickness of the muscular envelope around the neck, or at least around the ventral half. And the cervical ribs are typically pretty close to the vertebral centra–only weirdos like Apatosaurus and Erketu displace them very far ventrally (see Taylor and Wedel 2013a: fig. 7 and this post). So, thin jackets of muscle around proportionally large vertebrae–or, if you like, corn-on-the-cob rather than shish-kebabs.
As for why sauropods have long cervical ribs, Mike and I discussed some possibilities in our 2013 PeerJ paper (Taylor and Wedel 2013a), and Preuschoft and Klein addressed the issue last fall in PLOS ONE (Preuschoft and Klein 2013). My favorite hypothesis is that long tendons allow an animal to shift the bulk of the muscle–and therefore the center of gravity–toward the base of the neck, but that long unossified tendons can be distorted through stretching, which wastes muscular energy. Ossifying those long tendons is like putting bony wheelbarrow handles on each vertebra, allowing the muscles to move the vertebra from a distance without so much wasted energy, and probably with finer positional control.
That’s a nifty hypothesis in need of testing, anyway. In fact, cervical ribs and their associated muscles could stand a lot more attention on both the descriptive and analytical fronts. I know that Liguo Li has some research in the works on different conformations of hypaxial muscles, tendons, and cervical ribs in birds (you know, when she’s not describing bizarre new titanosaurs like Yongjinglong — see Li et al. 2014). If you saw Peter Dodson give their talk at SVP last fall, you probably remember some stunning images of dissected bird necks. As a famous legislator once said, we shall watch her career with great interest.
- Dalla Vecchia, F.M. 1999. Atlas of the sauropod bones from the Upper Hauterivian – Lower Barremian of Bale/Valle (SW Istria, Croatia). Natura Nacosta 18:6-41.
- Janensch, Werner. 1950. Die Wirbelsaule von Brachiosaurus brancai. Palaeontographica (Suppl. 7) 3: 27-93.
- Klein, N., Christian, A., & Sander, P. M. (2012). Histology shows that elongated neck ribs in sauropod dinosaurs are ossified tendons. Biology letters, 8(6), 1032-1035.
- Lacovara, Kenneth J.; Ibiricu, L.M.; Lamanna, M.C.; Poole, J.C.; Schroeter, E.R.; Ullmann, P.V.; Voegele, K.K.; Boles, Z.M.; Egerton, V.M.; Harris, J.D.; Martínez, R.D.; Novas, F.E. (September 4, 2014). A Gigantic, Exceptionally Complete Titanosaurian Sauropod Dinosaur from Southern Patagonia, Argentina. Scientific Reports. doi:10.1038/srep06196.
- Li L-G, Li D-Q, You H-L, Dodson P (2014) A New Titanosaurian Sauropod from the Hekou Group (Lower Cretaceous) of the Lanzhou-Minhe Basin, Gansu Province, China. PLoS ONE 9(1): e85979. doi:10.1371/journal.pone.0085979
- Preuschoft, H., & Klein, N. (2013). Torsion and Bending in the Neck and Tail of Sauropod Dinosaurs and the Function of Cervical Ribs: Insights from Functional Morphology and Biomechanics. PloS one, 8(10), e78574.
- Taylor, Michael P., and Mathew J. Wedel. 2013a. Why sauropods had long necks; and why giraffes have short necks. PeerJ 1:e36. 41 pages, 11 figures, 3 tables. doi:10.7717/peerj.36
- Wedel, M.J., Cifelli, R.L., and Sanders, R.K. 2000b. Osteology, paleobiology, and relationships of the sauropod dinosaurSauroposeidon. Acta Palaeontologica Polonica 45(4): 343-388.
You know the drill: lotsa pretty pix, not much yap.
Our first stop of the day was the Fruita Paleontological Area, which has a fanstastic diversity of Morrison animals, including the mammal Fruitafossor and the tiny ornithopod Fruitadens.
Plus it’s a pretty epic landscape, especially with the clouds and broken light we had this morning.
I found a bone! Several bits, actually, a few meters away from the Fruitadens type quarry. I’d like to think that this proximal femur might be Fruitadens, but I don’t know the diagnostic characters and haven’t had time to look them up. Anyone know how diagnostic this honorary shard of excellence might be?
After lunch, John Foster took us on a short hike to the quarry where Elmer Riggs got the back half of the Field Museum Apatosaurus. The front half came from a site in southern Utah, several decades later.
The locals brought Riggs out in the 1930s for the dedication of two monuments–this one at the Apatosaurus quarry, and another like it at the Brachiosaurus quarry some miles away. Tragically, both monuments have the names of the dinosaurs misspelled!
In the afternoon we visited the Mygatt-Moore Quarry and the Camarasaurus site in Rabbit Valley. Can you see the articulated Camarasaurus neck in this photo?
Here’s a hint: the neural arches of two posterior cervical vertebrae in
transverse horizontal cross-section.
This Camarasaurus is apparently a permanent feature. If you’re wondering why no-one has excavated it, it’s because it’s buried in sandstone that is stupid-dense. The expenditure of time and resources just isn’t worth it, when right down the hill dinosaurs are pouring out of the much softer sediments of the Mygatt-Moore Quarry like water from a hydrant. This is the lesson I am learning about the Morrison: finding dinosaurs is easy. Finding dinosaurs you can get out of the ground and prepare–that’s something else.
Our last stop of the day was Gaston Design, where Rob Gaston showed us how he molds, casts, and mounts everything from tiny teeth to good-sized skeletons.
Like this Deinosuchus that is about to chomp on Jim Kirkland. Jim doesn’t look too worried.
Here’s a nice cast of a busted sauropod dorsal, probably from Apatosaurus or Diplodocus, showing the pneumatic internal structure. Compare to similar views of dorsals in this post and this one. This is actually one half of a matched set that includes both halves of the centrum. I left with one of those sets of my own, a few dollars poorer and a whole lot happier.
The end–for now.
That last one really hurts. Here’s the original image, which should have gone in the paper with the interpretive trace next to it rather than on top of it:
Papers referenced in these slides:
- Taylor, M.P., and Wedel, M.J. 2013b. The effect of intervertebral cartilage on neutral posture and range of motion in the necks of sauropod dinosaurs. PLOS ONE 8(10): e78214. 17 pages. doi:10.1371/journal.pone.0078214
- Wedel, M.J. 2007a. What pneumaticity tells us about ‘prosauropods’, and vice versa. Special Papers in Palaeontology 77:207-222.
- 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.
On that last slide, I also talked about two further elaborations: figures that take up the entire page, with the caption on a separate (usually facing) page, and side title figures, which are wider than tall and get turned on their sides to better use the space on the page.
Also, if I was doing this over I’d amend the statement on the last slide with, “but it doesn’t hurt you at all to be cognizant of these things, partly because they’re easy, and partly because your paper may end up at an outlet you didn’t anticipate when you wrote it.”
And I just noticed that the first slide in this group has the word ‘without’ duplicated. Jeez, what a maroon. I’ll try to remember to fix that before I post the whole slide set at the end of this exercise.
A final point: because I am picking illustrations from my whole career to illustrate these various points, almost all fail in some obvious way. The photos from the second slide should be in color, for example. When I actually gave this talk, I passed out reprints of several of my papers and said, “I am certain that every single figure I have ever made could be improved. So as you look through these papers, be thinking about how each one could be made better.”
- Wedel, M.J. 2003b. The evolution of vertebral pneumaticity in sauropod dinosaurs. Journal of Vertebrate Paleontology 23:344-357.
- Wedel, M.J., and Sanders, R.K. 2002. Osteological correlates of cervical musculature in Aves and Sauropoda (Dinosauria: Saurischia), with comments on the cervical ribs of Apatosaurus. PaleoBios 22(3):1-6.
- Wedel, Mathew J., Richard L. Cifelli and R. Kent Sanders. 2000b. Osteology, paleobiology, and relationships of the sauropod dinosaurSauroposeidon. Acta Palaeontologica Polonica 45(4): 343-388.
January 15, 2014
[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.]
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!”
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.
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.
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.
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.
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.
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.
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.
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.
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.
- Stevens, K.A. and Parrish, J.M. 1999. Neck posture and feeding habits of two Jurassic sauropod dinosaurs. Science 284: 798-800. [Free subscription required]
- Stevens, Kent A., and J. Michael Parrish. 2005. Neck posture, dentition, and feeding strategies in Jurassic sauropod dinosaurs. pp. 212-232 in: Virginia Tidwell and Ken Carpenter (eds.), Thunder Lizards: the Sauropodomorph Dinosaurs. Indiana University Press, Bloomington, Indiana. 495 pp.
- Taylor, Michael P., and Mathew J. Wedel. 2013c. The effect of intervertebral cartilage on neutral posture and range of motion in the necks of sauropod dinosaurs. PLOS ONE 8(10): e78214. 17 pages. doi:10.1371/journal.pone.0078214 [PDF]
- Upchurch, P. 2000. Neck posture of sauropod dinosaurs. Science 287: 547b.
- Wedel, M.J. 2003b. The evolution of vertebral pneumaticity in sauropod dinosaurs. Journal of Vertebrate Paleontology 23:344-357.
- Wedel, M.J. 2007. Aligerando a los gigantes (Lightening the giants). ¡Fundamental! 12:1-84. [in Spanish, with English translation]
- Wedel, M.J., and Cifelli, R.L. 2005. Sauroposeidon: Oklahoma’s native giant. Oklahoma Geology Notes 65 (2):40-57.
- Wedel, M.J., R.L. Cifelli and R.K. Sanders. 2000. Osteology, paleobiology, and relationships of the sauropod dinosaur Sauroposeidon. Acta Palaeontologica Polonica 45(4): 343-388.
My spouse, Vicki, the other Dr. Wedel, is a physical and forensic anthropologist. And she’s one of a very small number of scientists who have (a) learned something new about the human body, and (b) used it to help identify dead people. And since that process involves the sciences of hard-tissue histology and skeletochronology–not to mention lots of dead folks–I reckon it might be of interest here. Hence this post.
This started about a decade ago, when Vicki was working on her PhD under Alison Galloway at UC Santa Cruz. Vicki worked with Alison on a ton of forensic cases, including some you probably heard of–they analyzed the remains of Laci Peterson and her unborn baby, Connor, for Scott Peterson’s murder trial. I had the unusual privilege of assisting a couple of times, on other cases, once to take some pictures in the lab while Vicki fished the skeleton out of the bag of skin that was all that was left of the body, and once to crawl around on my hands and knees picking human finger bones out of a muddy slough near Santa Cruz. All in all, I’m happy that my usual victims have been dead a lot longer.
Incidentally, the only show with forensic content that Vicki will watch voluntarily is Dexter. She cannot stand CSI, NCIS, or the other “behind the scenes” forensic investigation shows. We’ve tried watching them, but the inaccuracies drive her crazy (paleo people: imagine getting the Clockwork Orange therapy and being forced to watch Clash of the Dinosaurs). Real cases are solved by teams of specialists, not two omnicompetent protagonists; it takes weeks or months, not half an hour; and if the forensics people carry guns, it’s because they know waaaay too much about how some very bad, very organized people dispose of bodies (the short answer is, not thoroughly enough*).
* Once a guy who was threatening to testify against a certain criminal organization was shot in the head, his body burned, and his burnt remains scattered along the side of the road. Vicki and Alison picked the bone shards out of the roadside gravel, identified some of them as bits of skull, and found bevelling diagnostic of ballistics trauma on some of those. The way the bone had shattered showed that the gunshot had been inflicted perimortem–around the time of death–and before the body was burned. Bottom line, whatever plan you have to get rid of the body, it is probably not going to be enough to keep someone like Vicki from figuring out how you did it. That much, the TV shows do get right.
Not only is hard to really, truly get rid of a human body, it’s also hard to tell exactly when a person died, especially if all you have is a body in the woods. Insects are good–there’s a whole field of forensic entomology, whose practitioners age cadavers based on what insects are present and what stages of their life cycles they’re in. But what if all that is left is a pile of bones in the woods (which happens more often that you might think, and sometimes for completely innocuous reasons)? I’m preaching to the choir here, but bones can survive for a long time, so general wear-and-tear doesn’t tell you much. Rapetosaurus looks like it died last year.
There’s another side to this, which is figuring out how old someone was at the time of death based on their skeleton. Tooth eruption is good, and fusion of the epiphyseal growth plates, but both of those processes are basically done by the time people are in their mid-20s (teeth) to mid-30s (epiphyseal fusion). After that, there are methods based on the morphology of the auricular surface of the ilium and the public symphyses, but these only narrow things down to intervals of 5 to 15 years, and that’s a lot of missing persons reports to sift through. And none of the regular skeletal methods work past the age of 55 or 60. After that, no matter how healthy you are, the primary skeletal changes are attritional (i.e., you’re wearing out), and that process varies so much among individuals and populations that there are basically no predictive guidelines.
All of this was on Vicki’s mind when she was a grad student, so she was alert to anything that might help forensic anthropologists narrow down the possibilities for identifying dead folks. She was teaching in an osteology course and one of her students, Josh Peabody, brought up dental cementum increment analysis (DCIA), which is used in zooarcheology to determine the age and season at death of animal remains found at archaeological sites. Josh wanted to know if the method worked on humans.
At the time–2004–DCIA was being tested for age at death in some historical human populations from archaeological sites, but no-one had tried using it for season at death. So Vicki and Josh set out to see if it would work.
Our teeth, like those of other mammals, are held in their sockets by periodontal ligaments. The periodontal ligament of each tooth attaches via Sharpey’s fibers to the dental cementum on the tooth root(s). Cementum is laid down in annual bands, so you can count the number of bands on a tooth, add the normal age at which that tooth erupts, and get a pretty tight estimate of when the animal died. So much for age at death, which was already being done on humans in a limited way in the early 2000s, albeit in archaeological rather than forensic contexts.
But wait, there’s more. Actually two bands of cementum are laid down every year–a dark band in the winter (roughly October to March) and a light band in the summer (roughly April to September). ‘Dark’ and ‘light’ describe the appearance of the bands under polarized light microscopy. In the summer months, the collagen fibrils that make up the cementum are aligned parallel to the tooth root, so more light comes through. In the winter, the collagen is aligned perpendicular to the root, so less light is transmitted, and the winter bands appear darker by comparison. So not only does the number of pairs of light-and-dark bands tell you the number of years since the tooth erupted, the color of the outermost band tells you in which six-month period the individual died, and the thickness of the outermost band might help you narrow that down even further.
At least, that’s how it works in other mammals. Would it hold up in humans? After all, we’re pretty good at adjusting our environments to suit us, rather than vice versa. If the winter-summer banding pattern was present in humans, it would be a huge boon to forensic science. Even people in their 40s and beyond with no very reliable skeletal indicators of age could be aged to within a year or two, and their season at death narrowed down to a 2-3 month window.
To find out, Vicki and Josh had a dentist in Santa Cruz collect 112 teeth pulled from patients over the course of a year (with full IRB approval and informed consent from the dental patients). For their purposes, a tooth pulled from a live person is just as good as one from a cadaver or skeleton–extraction kills the tooth as surely as death of the body. Better, even, in that it was easier to quickly get lots of teeth with very precise extraction data.
Vicki and Josh cut a few teeth together and they found dark and light bands right away. They presented those preliminary results at the American Academy of Forensic Sciences meeting in 2005. After that, Josh got busy with his own research, but Vicki pressed on (while finishing a dissertation on different project, and being a first-time mom).
If this was a movie, this is the part where there would be a montage of inspirational music to get us quickly past a lot of hard, boring work. Each of the 112 teeth had to be embedded in plastic, a section through the root cut out with a saw, that section mounted on a slide and ground down until it was translucent (this process will be familiar to bone histologists of all stripes, paleo or neo). Then Vicki had to go all the way around the perimeter of the each root to find the place where the cementum bands showed the most clearly, and count them. This part is trickier than it sounds, unless you’ve done some histo and know just how butt-ugly some sections can be under the scope.
The results? In the words of the Bloodhound Gang, which Vicki quotes in her DCIA talks, “You and me baby ain’t nothin’ but mammals”. Here’s the payoff graph:
The one out-of-place measurement was probably caused by the dark band not being thick enough to register clearly on the image.
Now that she knew that DCIA could be used to determine season at death in humans, Vicki started applying it in her forensic cases, of which there have been many. The vast majority of the work of forensic anthropologists is invisible to the public: after analyzing a set of remains, a forensic anthropologist writes a case report for whatever law enforcement office (or, much less frequently, law firm or other entity) brought them in, and that’s that. The case reports are almost always confidential, but they have to be written to exacting standards since they may be used as evidence in court. So forensic anthropologists spend a lot of time toiling over papers that hardly anyone gets to read.
However, sometimes a case is written up for journal publication–if it’s sufficiently novel or unusual, and if permission can be secured from all of the relevant parties. In 2008, Vicki was approached by the Merced County sheriff’s office to help try to identify the remains of a young woman who had been murdered in 1971. That’s the 37-year-old cold case mentioned in the title of this post, and rather than tell you about it, I’ll point you to Vicki’s case report (Wedel et al. 2013), published last month in the Journal of Forensic Identification and freely available here.
I wasn’t sure whether to post about this or not–as cool as they are, murder cases are not our normal stock in trade on this blog. What decided me was talking with Andy Farke. He read Vicki’s paper as soon as it came out, and he said that he really enjoyed getting to see how forensic anthropologists work in the real world. I sometimes take for granted that, since I am married to a forensic anthropologist, I get to see how this works all the time. But that’s a pretty rare experience–if paleontology is a small field, forensic anthropology is positively tiny. So if you want to see an example of the real science that CSI and the like are based on, here’s your window.
What’s next? Vicki has several validation studies on DCIA in progress, for which she and her collaborators have collected a much larger sample size–over 1000 teeth–to try to answer questions like: what tooth is best to use for DCIA? Should the histological sections be made longitudinally or transversely through the tooth root? Does cementum banding vary with latitude? And since banding patterns are reversed in the Southern Hemisphere, following the flip-flopped season, what happens at the equator? Watch this space, and keep an eye out for Vicki’s future publications–including a book due out next year–at her website, Bodies, Bugs, and Bones.
- Wedel, V.L. 2007b. Determination of season at death using dental cementum increment analysis. Journal of Forensic Sciences 52(6): 1334-1337.
- Wedel, V.L., G. Found, and G.L. Nusse. 2013. A 37 year-old cold case identification using novel and collaborative methods. Journal of Forensic Identification 63(1): 5-21.