Caudal pneumaticity in saltasaurines. Cerda et al. (2012: fig. 1).
Earlier this month I was amazed to see the new paper by Cerda et al. (2012), “Extreme postcranial pneumaticity in sauropod dinosaurs from South America.” The title is dramatic, but the paper delivers the promised extremeness in spades. Almost every figure in the paper is a gobsmacker, starting with Figure 1, which shows pneumatic foramina and cavities in the middle and even distal caudals of Rocasaurus, Neuquensaurus, and Saltasaurus. This is most welcome. Since the 1990s there have been reports of saltasaurs with “spongy bone” in their tail vertebrae, but it hasn’t been clear until now whether that “spongy bone” meant pneumatic air cells or just normal marrow-filled trabecular bone. The answer is air cells, loads of ‘em, way farther down the tail than I expected.
Caudal pneumaticity in diplodocines. Top, transverse cross-section through an anterior caudal of Tornieria, from Janensch (1947: fig. 9). Bottom, caudals of Diplodocus, from Osborn (1899: fig. 13).
Here’s why this is awesome. Lateral fossae occur in the proximal caudals of lots of neosauropods, maybe most, but only a few taxa go in for really invasive caudal pneumaticity with big internal chambers. In fact, the only other sauropod clade with such extensive pneumaticity so far down the tail are the diplodocines, including Diplodocus, Barosaurus, and Tornieria. But they do things differently, with BIG, “pleurocoel”-type foramina on the lateral surfaces of the centra, leading to BIG–but simple–camerae inside, and vertebral cross-sections that look like I-beams. In contrast, the saltasaurines have numerous small foramina on the centrum and neural arch that lead to complexes of small pneumatic camellae, giving their vertebrae honeycomb cross-sections. So caudal pneumaticity in diplodocines and saltsaurines is convergent in its presence and extent but clade-specific in its development. Pneumaticity doesn’t get much cooler than that.
Pneumatic ilia in saltasaurines. Cerda et al. (2012: fig. 3).
But it does get a little cooler. Because the stuff in the rest of the paper is even more mind-blowing. Cerda et al. (2012) go on to describe and illustrate–compellingly, with photos–pneumatic cavities in the ilia, scapulae, and coracoids of saltasaurines. And, crucially, these cavities are connected to the outside by pneumatic foramina. This is important. Chambers have been reported in the ilia of several sauropods, mostly somphospondyls but also in the diplodocoid Amazonsaurus. But it hasn’t been clear until now whether those chambers connected to the outside. No patent foramen, no pneumaticity. It seemed unlikely that these sauropods had big marrow-filled vacuities in their ilia–as far as I know, all of the non-pneumatic ilia out there in Tetrapoda are filled with trabecular bone, and big open marrow spaces only occur in the long bones of the limbs. And, as I noted in my 2009 paper, the phylogenetic distribution of iliac chambers is consistent with pneumaticity, in that the chambers are only found in those sauropods that already have sacral pneumaticity (showing that pneumatic diverticula were already loose in their rear ends). But it’s nice to have confirmation.
So, the pneumatic ilia in Rocasaurus, Neuquensaurus, and Saltasaurus are cool because they suggest that all the other big chambers in sauropod ilia were pneumatic as well. And for those of you keeping score at home, that’s another parallel acquisition in Diplodocoidea and Somphospondyli (given the apparent absence of iliac chambers in Camarasaurus and the brachiosaurids, although maybe we should bust open a few brachiosaur ilia just to be sure*).
* I kid, I kid.**
** Seriously, though, if you “drop” one and find some chambers, call me!
Pectoral pneumaticity in saltasaurines. Cerda et al. (2012: fig. 2).
But that’s not all. The possibility of pneumatic ilia has been floating around for a while now, and most of us who were aware of the iliac chambers in sauropods probably assumed that eventually someone would find the specimens that would show that they were pneumatic. At least, that was my assumption, and as far as I know no-one ever floated an alternative hypothesis to explain the chambers. But I certainly did not expect pneumaticity in the shoulder girdle. And yet there they are: chambers with associated foramina in the scap and coracoid of Saltasaurus and in the coracoid of Neuquensaurus. Wacky. And extremely important, because this is the first evidence that sauropods had clavicular air sacs like those of theropods and pterosaurs. So either all three clades evolved a shedload of air sacs independently, or the basic layout of the avian respiratory system was already present in the ancestral ornithodiran. I know where I’d put my money.
There’s loads more interesting stuff to talk about, like the fact that the ultra-pneumatic saltasaurines are among the smallest sauropods, or the way that fossae and camerae are evolutionary antecedent to camellae in the vertebrae of sauropods, so maybe we should start looking for fossae and camerae in the girdle bones of other sauropods, or further macroevolutionary parallels in the evolution of pneumaticity in pterosaurs, sauropods, and theropods. Each one of those things could be a blog post or maybe a whole dissertation. But my mind is already thoroughly blown. I’m going to go lie down for a while. Congratulations to Cerda et al. on what is probably the most important paper ever written on sauropod pneumaticity.
References
- Cerda, I.A., Salgado, L., and Powell, J.E. 2012. Extreme postcranial pneumaticity in sauropod dinosaurs from South America. Palaeontologische Zeitschrift. DOI 10.1007/s12542-012-0140-6
- Janensch, W. 1947. Pneumatizitat bei Wirbeln von Sauropoden und anderen Saurischien. Palaeontographica, Supplement 7, 3:1–25.
- Osborn, H. F. 1899. A skeleton of Diplodocus. Memoirs of the American Museum of Natural History 1:191–214.
… but red-cyan anaglyphs are cooler
December 14, 2010
Over at his truly unique blog Paleo Errata, Jeff Martz is claiming that Stereopairs Are Cool. This assertion he supports with the following figure that he put together, showing a set of five stereopairs of a Longosuchus braincase:
Unfortunately, I am one of those who can’t “see” stereopairs, so these images are uninformative to me — or, at least, no more informative than your average inch-wide braincase photo.
So how else can we envisage the stereo information in these pairs of photos that Jeff took? My favourite way is using red-cyan anaglyphs — those goofy 3d images that you look at through 3d glasses. To compare, I did this to Jeff’s image. The process is simple: take two copies of the stereopair image, cut out all the right-eye views from one set and all the left-eye views from the other, then edit the colour levels of both layers. In one, take the red right down to zero, so you only have blue+green=cyan; in the other take the green and blue down so you only have red. Then stack one layer on top of the other and change its mode to “Lighten only”. Export the result as a JPEG and you get this result:
Armed with my red-cyan glasses (which, remember, I got as a freebie with a Lego catalogue), I can now make out the 3d structure really easily. Positives for the anaglyph approach:
- The 3D image is much easier to see
- The result takes up less space on the page
- Most importantly, the size limitation is removed: I have some beautiful whole-screen anaglyphs (e.g. Archbishop cervical, wallaby skull), whereas stereograms are restricted to a couple of inches’ separation.
The downside is, of course, that you need special equipment to see them –albeit equipment so laughably minimal that Amazon.com will sell you THREE PAIRS for $1.39, you cheap gits. But for those of who who are too poor to find $1.39, and who don’t have two friends with whom you can form an ad-hoc 3D-glasses buying consortium at a cost of $0.47 each, there is one further approach: a low-rent technique that I call a “wigglegram” for want of a better term. Here it is:
I discovered this approach by accident, when flipping through a bunch of photographs that I’d taken of, I think, the Archbishop. As a matter of policy, I take most of my photos twice, so that if I shake slightly or the auto exposure gets it wrong, I have a good copy that I can retain. I was trying to decide which of two nearly identical pictures to keep. But as it happened, I’d moved the camera slightly to the side between taking the first and the second, so as I skipped back and forth between them, I was seeing two slightly different perspectives.
So there you have it: three different ways to visualise 3d structure, each built from the same basic set of photos. They each have their merits, and I hope we’ll increasingly see more of all three of them, as we move into the Shiny Digital Future, and arbitrary limits on manuscript length and numbers of figures get lifted.
I leave y0u with an actual application of all this. Matt and I have, for some time, been working on a manuscript about caudal pneumaticity in sauropods, and we wanted to include a brief survey of which genera it’s been reported in. Among the candidates was Saltasaurus, which has a candidate pneumatic caudal vertebra that was illustrated thus by Powell (2003: plate 53, part 3):
Matt can “see” stereograms, and insisted that the dark patch on the side of the centrum is a pneumatic fossa. I wasn’t so sure, and in fact we got into quite an argument over whether or not to include this specimen in our list. The argument was neatly concluded when I had the obvious idea of converting Powell’s stereogram into an anaglyph:
As soon as I saw this, I recognised what the structure is: the crescent moon-shaped dark patch is indeed a deep, invasive fossa, and the broad, roughly circular object above it and to the right is a lumpen lateral process sticking right out into the camera (and partially hiding the fossa). So Matt was right, the vertebra is pneumatic, and a beautiful friendship was saved by the power of red-cyan anaglyphys. Yay!
References
- Powell, Jaime E. 2003. Revision of South American Titanosaurid dinosaurs: palaeobiological, palaeobiogeographical and phylogenetic aspects. Records of the Queen Victoria Museum 111: 1-94.
More on the internal structure of Alamosaurus vertebrae
January 18, 2010
A section of the cotyle of a presacral vertebra of Alamosaurus (Woodward and Lehman 2009:fig. 6A).
The last time we talked about Alamosaurus, I promised to explain what the arrow in the above image is all about. The image above is a section through the cotyle (the bony socket of a ball-and-socket joint) at the end of one of the presacral vertebra. The external bone surface would have been over on the left; it was either very thin (which happens) or a bit eroded, or both. The arrow is pointing at something weird–a plate of bone inside the vertebra that forms a sort of shadow cotyle deep to the articular surface.
This is weird for a couple of reasons. First, once camellate (small-chambered) vertebrae get above a certain level of complexity, it’s hard to make any sense of the orientation of individual bony struts. Possibly I haven’t seen enough vertebrae, or played with enough 3D models, to figure it out. You would certainly expect that the struts would be oriented to resist biomechanical loads, just like the struts in the long bones of your limbs; the fact that sauropod verts were filled with air whereas your long bones are filled with marrow shouldn’t make any difference. Back in the day, Kent Sanders–who is second author on that super-important paper on unidirectional air flow in croc lungs that you’ve probably heard about (Farmer and Sanders 2010)–speculated to me that the complex of laminae we see in the vertebrae of most sauropods are still there in the inflated-looking vertebrae of titanosaurs and birds, they’re just incarnated in internal struts rather than external laminae. Cool hypothesis for somebody to test.
The other reason that this is weird is that the plate of bone is parallel to the articular surface. One place where I have seen some regularity in terms of strut orientation is in zygapophyses, where in both camerae and camellate vertebrae the internal struts are oriented at right angles to the articular surfaces of the zygs, like beams propping up a wall. In this Alamosaurus section, there are indeed smaller struts that run at right angles to both the cotyle and the internal plate, but I have no idea why they’re so wimpy and the plate is so thick; a priori I would have expected the reverse.
It turns out that this isn’t even the first time that an internal “shadow” of the cotyle has been figured–check out this figure that I redrew from Powell’s (1992:fig. 16) Saltasaurus osteology. But don’t credit me with the discovery. I’d looked at this section a hundred times and even drawn it and never noticed the shadow cotyle, until it was pointed out by Woodward and Lehman (2009)–another reason to read that paper if you haven’t yet. Kudos to Holly Woodward for spotting this and making the connection.
Now that I’ve drawn attention to the weirdness and given credit where it’s due, this is one of those times I’m going to throw up my hands in confusion and open the floor for comments.
References
- Farmer, C.G., and Sanders, K. 2010. Unidirectional airflow in the lungs of alligators. Science 327:338-340.
- Powell, J.E. 1992. Osteologia de Saltasaurus loricatus (Sauropoda – Titanosauridae) del Cretacico Superior del noroeste Argentino; pp. 165-230 in J.L. Sanz and A.D. Buscalioni (editors), Los Dinosaurios y Su Entorno Biotico: Actas del Segundo Curso de Paleontologia in Cuenca. Institutio Juan de Valdes.
- Woodward, H.N., and Lehman, T.M. 2009. Bone histology and microanatomy of Alamosaurus sanjuanensis (Sauropoda: Titanosauria) from the Maastrichtian of Big Bend National Park, Texas. Journal of Vertebrate Paleontology 29(3):807-821.
ASPs for Alamosaurus
January 4, 2010
A section of the cotyle of a presacral vertebra of Alamosaurus (Woodward and Lehman 2009:fig. 6A). The arrow will be explained in a future post!
Last year was good for sauropod pneumaticity. In the past few months we’ve had the publication of the first FEA of pneumatic sauropod vertebrae by Schwarz-Wings et al (2009), as well as a substantial section on pneumaticity in the big Alamosaurus histology paper by Woodward and Lehman (2009). I won’t repeat here everything that Woodward and Lehman have to say about pneumaticity, I just want to draw attention to a little piece of it. Their work is observant, up-to-date, and worth reading, so if you can get access to the paper, read it.
The major brake on the growth of our knowledge and understanding of pneumaticity is sample size. I harped on this in 2005 (Wedel 2005), and Mike just brought it up again in a comment on a previous post. In fact, what he had to say is so relevant that I’m going to just cut and paste it here:
How does degree of pneumatisation vary between individuals? Here are three more: how does it vary along the neck, how does it vary long the length of an individual vertebra, and how does it vary through ontogeny? Then of course there is variation between taxa across the tree. So what we have here is a five-and-half-dimensional space that we want to fill with observations so that we can start to deduce conclusions. Trouble is, there are, so far, 22 published observations (neatly summarised by Wedel 2005:table 7.2), which is not really enough to let us map out 5.5-space! That’s one reason why, at the moment, each observation is valuable — it adds 4% to the total knowledge in the world.
To be fair, there are a few more published observations. Schwarz and Fritsch (2006) published ASPs for cervicals of Giraffatitan and Dicraeosaurus, and I have a gnawing feeling that there are a couple here and there that I’ve seen but not remembered. I’ve got some more of my own data in the as-yet-unpublished fourth chapter of my diss, which I failed to get out as part of the Paleo Paper Challenge. And, getting back to the subject of the post, Woodward and Lehman (2009:819) have some tasty new data to report:
Digital images of sections of vertebrae and ribs were imported into ArcGIS 8.1 (Dangermond, 2001; for methods see Woodward, 2005). A unitless value for the total area of the image was calculated, using the outline of the bone as a perimeter. Subtracted from this was the area value taken up by bone, as determined by color differences (lighter areas are camellate cavities, darker areas are bone). Using this method, longitudinal sections of centra are estimated to be roughly 65% air filled. The amount of open space similarly calculated for the pneumatic proximal and medial rib sections is about 52%, whereas the cancellous spongiosa in distal rib transverse sections yields an average estimate of about 44% of their cross sectional area. Hence, the camellate cavities result in an appreciably lower bone volume compared to spongiosa.
The ASP of 0.65 for centra is right in line with the numbers I’ve gotten for neosauropods, and with the results of Schwarz and Fritsch (2006) for Giraffatitan (Dicraosaurus had a much lower ASP, around 0.2 IIRC). The stuff about the ribs is particularly interesting. Using densities of 0.95 for bone marrow, 1.8 for avian (and sauropod) compact bone, and 1.9 for mammalian compact bone we get the following:
- Pneumatic Alamosaurus vertebrae – ASP of 0.65, density of 0.63 g/cm^3.
- Pneumatic Alamosaurus ribs – ASP of 0.52, density of 0.86 g/cm^3.
- Apneumatic Alamosaurus ribs – MSP (marrow space proportion) of 0.44, density of 1.43 g/cm^3.
- Pneumatic bird long bones – ASP of 0.59, density of 0.74 g/cm^3.
- Apneumatic bird long bones – MSP of 0.42, density of 1.44 g/cm^3.
- Apneumatic mammal long bones – MSP of 0.28, density of 1.63 g/cm^3.
ASPs and MSPs of bird and mammal bones are calculated from K values reported by Cubo and Casinos (2000) for birds and Currey and Alexander (1985) for mammals. I don’t know what the in vivo density of sauropod compact bone was; changing it from the avian value of 1.8 to the mammalian value of 1.9 would have a negligible effect on the outcome.
At least with the data in hand, we can make the following generalizations:
- The apneumatic bones of birds are thinner-walled than those of mammals, on average. (This has been known for a long time.)
- The apneumatic ribs of Alamosaurus were more similar in density to apneumatic bird bones than to apneumatic mammal bones.
- In both birds and Alamosaurus, pneumatization reduces the amount of bone tissue present by 15-30% in the same elements (long bones for birds, ribs for Alamosaurus). Pneumatic bones are light not just because the marrow is replaced by air, but because there is less bone tissue than in apneumatic bones, as bird people have been observing for ages.
There’s loads more work to be done on this sort of thing, so I’m going to stop blogging now and get back to it. Stay tuned!
References
- Cubo, J., and Casinos, A. 2000. Incidence and mechanical significance of pneumatization in the long bones of birds. Zoological Journal of the Linnean Society 130: 499–510.
- Currey, J. D., and Alexander, R. McN. 1985. The thickness of the walls of tubular bones. Journal of Zoology 206:453–468.
- Schwarz D, and Fritsch G. 2006. Pneumatic structures in the cervical vertebrae of the Late Jurassic Tendaguru sauropods Brachiosaurus brancai and Dicraeosaurus. Eclogae Geologicae Helvetiae 99:65–78.
- Schwarz-Wings, D., Meyer, C.A., Frey, E., Manz-Steiner, H.-R., and Schumacher, R. 2009. Mechanical implications of pneumatic neck vertebrae in sauropod dinosaurs. Proceedings of the Royal Society B. doi: 10.1098/rspb.2009.1275
- Wedel, Mathew J. 2005. Postcranial skeletal pneumaticity in sauropods and its implications for mass estimates. pp. 201-228 in Wilson, J. A., and Curry-Rogers, K. (eds.), The Sauropods: Evolution and Paleobiology. University of California Press, Berkeley.
- Woodward, H.N., and Lehman, T.M. 2009. Bone histology and microanatomy of Alamosaurus sanjuanensis (Sauropoda: Titanosauria) from the Maastrichtian of Big Bend National Park, Texas. Journal of Vertebrate Paleontology 29(3):807-821.
