March 12, 2011
UPDATE April 16, 2012: The paper is officially published now. I’ve updated the citation and link below accordingly.
More new goodies:
Yates, A.M., Wedel, M.J., and Bonnan, M.F. 2012. The early evolution of postcranial skeletal pneumaticity in sauropodomorph dinosaurs. Acta Palaeontologica Polonica 57(1):85-100. doi: http://dx.doi.org/10.4202/app.2010.0075
This is only kinda sorta published. The accepted manuscript is now posted on the APP website, and it has a DOI, but it’s not formatted or available in print yet. But after discussing it amongst ourselves, we authors agreed that (1) the paper is globally available and it’s silly to pretend otherwise, (2) there are no nomenclatural ramifications of that fact, and (3) we’re tired of not being able to talk about this stuff. So we’re gonna, starting…now.
A brief tale of Serendipity in Science (TM):
Back in 2004 I was in my third year of grad school at Berkeley. My fellow grad student, Brian Kraatz, gave me a heads up about the 19th International Congress of Zoology coming up in Beijing. Attendees could submit 500-word abstracts or 2000-word short papers. I didn’t plan on doing either one, until the night before they were due, when I changed my mind and wrote almost all of what would become this paper in a single six-hour session (don’t be too impressed; I’ve been trying to replicate that feat for seven years with no success).
That summer, I met up with Brian in Beijing a week before the congress, and we spent the extra time working in the collections of the Institute of Vertebrate Paleontology and Paleoanthropology (IVPP). Paul Barrett was there, working on prosauropods, and he and I had some long and fascinating conversations. We also gave our talks in the same session at the congress. Paul must have decided I was not a complete moron because he invited me to give a talk in the basal sauropodomorph symposium at SVP in 2005.
A brief aside: many of the animals I grew up calling prosauropods ended up outside of the monophyletic Prosauropoda that is anchored on Plateosaurus. Some are now basal sauropods, some are closer to sauropods than to Plateosaurus but outside of Sauropoda, and some are outside of Prosauropoda + Sauropoda. The phylogenetically correct term encompassing all of the nonsauropods is ’basal sauropodomorphs’, and it means roughly what ‘prosauropods’ did until a decade or so ago. I often slip into informally using ‘prosauropods’, but I try to remember to put the term in quotes so as not to mislead anyone.
I had been to England in 2004 and 2005 and seen the putatively pneumatic vertebrae of Erythrosuchus and what was then known as Thecodontosaurus caducus (and is currently trading under the name Pantydraco caducus for reasons that it would be otiose, for the moment, to rehearse)–and, not incidentally, had finally met Mike in person, although we’d been corresponding since 2000. I’d also been to Stuttgart primarily to see the appendicular material of Janenschia and ended up spending some quality time with Plateosaurus. (Since the theme here is serendipity, note that the Janenschia work–my raison d’etre for going to Germany–died on the table, whereas I’ve now been an author on three ‘prosauropod’ papers and have more in the works. Weird!)
Anyway, with all of that accidental experience with ‘prosauropods’ and other interesting critters like Erythrosuchus, I found that I actually had something to say in 2005 SVP symposium. I titled my talk, ‘What pneumaticity tells us about “prosauropods”, and vice versa’, and it turned into the 2007 paper of the same title.
None of this would have happened if Brian hadn’t hounded me about going to Beijing, and if I hadn’t ended up talking so much with Paul on that trip, and if I hadn’t finished up with Janenschia on my first day in Stuttgart and spent the rest of the week playing with Plateosaurus. And so on. Science is unpredictable, especially for scientists.
When I sent around the PDF of the paper to friends and colleagues, I included this quip: “Were prosauropods pneumatic? The fossils don’t say. Somehow I stretched that out to 16 pages.” Mike claims that because of this quip he’s never been able to take that paper seriously. But it is my favorite among my solo efforts. It includes loads of stuff on the origins of air sacs and pneumaticity that I wasn’t able to get into my earlier papers, either because it wasn’t directly relevant or because some reviewer forced me to excise it.
Almost immediately after the paper came out, Adam Yates and Matt Bonnan went and found roughly a zillion pneumatic ‘prosauropods’, which was a bit embarrassing since I’d just concluded that the evidence for ‘prosauropod’ pneumaticity was thin to nonexistent. So it is a damn good thing for me that I was already on friendly terms with both of them, because instead of taking the opportunity to smack me down, they invited me on board. Which led to Adam’s talk at SVP in Bristol in 2009, and to the new paper.
And actually, the depth of my incorrectness was even greater than I had thought. I reckon that literally millions of people have seen the mounted Plateosaurus skeleton in the AMNH, and any of them who have looked closely have seen this:
(Click for full size, unlabeled version.)
You see the problem here, I’m sure: the semi-big, semi-obvious fossa divided by an accessory lamina, not consistent with a muscle attachment point or fat pad or cartilage or infection, but very consistent in both form and location with the pneumatic fossae of other, more derived sauropodomorphs. On the lateral face of the vertebra, probably seen by millions, obvious to anyone who cares to look. A pneumatic prosauropod, in other words, right out in public for decades and decades (this time I don’t have to use the scare quotes because Plateosaurus actually IS a prosauropod sensu stricto). I didn’t even notice the first time I visited the AMNH back in 2006. I took the above photos, which are the basis for Figure 4 in the paper, in 2009.
So: ‘prosauropods’ were pneumatic. Some of them. A little bit. If you’d like to know more, please read the paper–it’s free.
Finally, a big thank-you to Adam and Matt for inviting me to be part of this. I think it’s pretty cool stuff, and I’m sure I’ll have more to say about it in the future. They might too–you should be reading their blogs, Dracovenator and Jurassic Journeys, anyway.
We’re still not done with Brontomerus, by the way. If nothing else, there’s the long-overdue post on how sauropod ilia change (or rather fail to change) through ontogeny. But that’s something we’ll have to get back to next week. Stay tuned.
February 12, 2011
When you last saw this rhea neck, I was squeezing a thin, unpleasant fluid out of its esophagus. Previous rhea dissection posts are here and here; you may also be interested in my ratite clearing house post.
We did that dissection back in 2006. Since then I finished my dissertation, got a tenure-track job, and moved twice. The rhea neck followed me, living in a succession of freezers until last spring.
Last spring I thawed it out, straightened it (it had been coiled up in a gallon ziploc), refroze it, and had it cut in half sagittally with a bandsaw. I did all of this for a project that is not yet ready to see the light of day, but there’s a ton of cool morphology here that I am at liberty to discuss, so let’s get on with it.
Throughout the post, click on the images for full resolution, unlabeled versions.
In the image above, you’ll notice that the saw cut was just slightly to the left of the midline, so that almost the entire spinal cord was left in the right half of the neck (the one toward the top of the image; the left half, below, is upside down, i.e. ventral is towards the top of the picture). The spinal cord is the prominent yell0w-white stripe running down the middle of the hemisectioned neck. It’s a useful landmark because it stands out so well. Dorsal to it are the neural arches, spines*, and zygapophyses of the vertebrae, and epaxial muscles; ventral to it are the vertebral centra and the hypaxial muscles.
* If you want to call them that–some of them are barely there!
Here’s the large supraspinous ligament (lig. elasticum interspinale), which is similar to the nuchal ligament of mammals but independently derived. Compare to the nuchal ligament of a horse (image borrowed from here):
Note how the actual profile of the neck is vastly different from what you’d suspect based on the skeleton alone. This is one of the reasons that necks lie. For more on the supraspinous ligament in rheas and its implications for sauropods, see Tsuihiji (2004) and Schwarz et al. (2007).
Birds also have very large interspinous ligaments (lig. elasticum interlaminare), each of which connects the neural spines of two adjacent vertebrae. In the above photo, the blunt probe is passing under (= lateral to) the unpaired, midline interspinous ligament. Rheas are unusual among birds in having such a large supraspinous ligament, and you can see that this interspinous ligament is almost as big. If you tear down the neck of a chicken or turkey, you will find huge interspinous ligaments, and the supraspinous ligament will be tiny if you can identify it at all.
Here’s something I don’t think we’ve ever shown before here on SV-POW!: a photograph of an actual pneumatic diverticulum. That’s the dark hole in the middle of the photo. You can see that we’re in the left half of the neck, lateral to the spinal cord, almost to the postzygapophysis, the articular surface of which is more lateral still (“below” or “deep to” the surface you see exposed in this cut). Usually at each intervertebral joint there is a connection between the lateral pneumatic diverticula that run up the side of the cervical column and pass through the cervical rib loops and the supramedullary diverticula that lie dorsal to the spinal cord inside the neural canal. That connecting diverticulum is the one exposed here.
NB: diverticulum is singular, diverticula is plural. There are no diverticulae or, heaven forbid, diverticuli, although these terms sometimes crop up in the technical literature, erroneously. (I hesitate to point this out, not because it’s not important, but because I’ll be lucky if I didn’t screw up a Latin term elsewhere in the post!)
Here are pneumatic diverticula in a transverse CT section of an ostrich neck (Wedel 2007b: fig. 6; compare to Wedel 2003: fig. 2, another slice from the same neck). In this view, bone is white, muscles and other soft tissues are gray, and air spaces are black. A, lateral diverticula running alongside the vertebral centra. B, air spaces inside the bone. C, supramedullary airways above the spinal cord. This section is close to the posterior end of a vertebra; the flat-bottomed wing-like processes sticking out to either side are the anterior portions of the postzygapophyses. If the slice was a few mm more posterior, we would see the prezygapophyses of the preceding vertebra in contact with them. Also, the vertical bars of bone connecting the centrum to the postzygs would pinch out, and we’d see the diverticula connecting the lateral (A) and supramedullary (C) airways–that’s the diverticulum revealed in the photo two images up.
Here’s another cool section showing a diverticulum and some muscles. Note the short interspinous muscles, which connect the neural spines of adjacent vertebrae. The probe indicates another open diverticulum, and the very tip of the probe is under one of the very thin layers of epithelium that line the diverticula. You can see that this diverticulum lies on the dorsal surface of the vertebra, posterior to the prezygapophysis and anterior to the neural spine. This supravertebral diverticulum is near and dear to my heart, because I have published an image of its traces before.
Lots going on in this photo (remember that you can click for an unlabeled version). This is a middle cervical vertebra of an emu, in anterodorsal view, with anterior towards the bottom of the picture. Bonus geek points if you recognized it as the basis for Text-fig. 9 in Wedel (2007a). I published this photo in that paper because it so nicely illustrates how variable the skeletal traces of pneumaticity can be, even from left to right in a single bone. On the right side of the photo (left side of the vertebra), the bone resorption adjacent to the supravertebral diverticulum produced a pneuamtic fossa, but one without distinct bony margins or a pneumatic foramen. On the other side, the fossa contains a pneumatic foramen which communicates with the internal air spaces, but the fossa is otherwise identical. Fossae like the one on the right are a real pain in the fossil record, because they might be pneumatic, but then again they might not be; such shallow, indistinct fossae can house other soft tissues, including cartilage and fat. This is what I was talking about when I wrote (Wedel 2009: p. 624):
If progressively more basal taxa are examined in the quest to find the origin of PSP [postcranial skeletal pneumaticity], the problem is not that evidence of PSP disappears entirely. It is that the shallow, unbounded fossae of basal dinosaurs are no longer diagnostic for pneumaticity.
For more on that problem, see Wedel (2007a) and the post, “X-Men Origins: Pneumaticity”.
The other labelled bits in the above photo are all muscle attachment points, and you may find Wedel and Sanders (2002), especially Fig. 2, a useful reference for the rest of the post. The dorsal tubercles, or epipophyses, are rugosities dorsal to the postzygapophyses that anchor most of the long, multi-segment epaxial muscles, which in birds are the M. longus colli dorsalis, which originates on the anterior faces of the neural spines, and M. ascendens cervicalis, which originates on the cervical rib loops. The crista transvers0-obliqua is a low, bony crest connecting each dorsal tubercle to the neural spine; it corresponds to the spino-postzygapophyseal lamina (SPOL) of sauropods (see Tutorial 4: Laminae!), and anchors the Mm. intercristales, a group of short muscles that span the cristae of adjacent vertebrae, like the Mm. interspinales only more lateral.
The carotid tubercles serve as points of origin for the M. longus colli ventralis, one of the largest and longest of the multi-segment hypaxial muscles; they have no obvious homolog or analog in sauropods. The lack of this feature might indicate that the hypaxial muscles were less of a big deal in sauropods, for whom lifting the neck was presumably a bigger problem than lowering it. Alternatively, the M. longus colli ventralis of sauropods might have attached to the medial sides of the parapophyses and the capitula of the cervical ribs, which tended to be larger and more ventrally-directed than in basal sauropodomorphs and theropods.
The unlabeled red arrows mark the lateral tubercles and crests of the cervical rib loop, to which we will return momentarily.
Here you can see a big bundle of long epaxial muscles, including both the M. longus colli dorsalis and M. ascendens cervicalis, inserting on the left dorsal tubercle of the vertebra on the right. Note that the cut here is quite a bit lateral of the midline, and actually goes through the lateral wall of the neural canal in the vertebra on the right (that vert is the fifth back from the front of the section of neck featured in this post, which is incomplete). That is why you see the big, multi-segment muscles here, and not the shorter, single-segment muscles, which lie closer to the midline.
Here are some more muscle attachment points in a bird vertebra (a turkey this time, courtesy of Mike). The lateral crests and tubercles (tubecula ansae and cristae laterales, if you’re keeping track of the Latin) are the same bony features indicated by the red arrows in the photo of the emu vertebra up above. They anchor both the long M. ascendens cervicalis, which inserts on the dorsal tubercles of more anterior vertebrae, and the short Mm. intertransversarii, which span the cervical rib loops of adjacent vertebrae. Sauropods usually have at least small rugosities on their diapophyses and the tubercula of their cervical ribs (which articulate with the diapophyses) that probably anchored homologous muscles.
Here’s a dorsal tubercle above the postzyg on the neural arch of a juvenile Apatosaurus (cervical 6 of CM 555, shown in right lateral view). Notice that the spinopostzygapophyseal lamina (SPOL) and postzygodiapophyseal lamina (PODL) actually converge on the dorsal tubercle rather than on the postzyg. This is pretty common, and makes good mechanical sense.
Dorsal tubercles again, this time on the world’s most wonderful fossil, cervical 8 of the HM SII specimen of Giraffatitan brancai, in the collections of the Humbolt museum in Berlin. While you’re here, check out the pneumato-riffic sculpting on the lateral faces of the neural arch and spine, and the very rugose texture on the tip of the neural spine, SPOLs, and dorsal tubercles. In fact, compare the numerous pocket-like external fossae on this vertebra with the internal air cells exposed in the cross-sectioned rhea neck. I have argued here before that sauropod cervical vertebrae are pretty similar to those of birds; the main differences are that the cervical rib loops are proportionally much smaller in sauropods, and sauropod vertebrae mostly wore their pneumaticity on the outside.
Farther anteriorly in the neck–the three vertebrae pictured here are the third, fourth, and fifth (from right to left) in this partial neck–and somewhat closer to the midline. Now you can see some short epaxial muscles, probably Mm. intercristales and Mm. interspinales (the two groups grade into each other and are often not distinct), spanning adjacent vertebrae. As in several previous photos, the supravertebral diverticulum is visible, as well as the communicating diverticulum that connects the lateral diverticula to the supramedullary airways. I forgot to label them, but ventral to the centra you can see long, light-colored streaks running through the hypaxial muscles. These are the tendons of the M. longus colli ventralis, and in some of the previous photos you can see them running all the way to their origination points on the carotid tubercles. These extend posteriorly from the short cervical ribs of birds, and are homologous with the long cervical ribs of sauropods.
That’s all I have for this time. If you’d like to see all of this stuff for yourself, turkey necks are cheap and big enough to be easy to work with. Geese are good, too. You can see all the same bits in a chicken or a duck, it’s just harder because everything is smaller (if you’re a real glutton for punishment, try a Cornish game hen).
When I first started working on sauropods, their cervical vertebrae made no sense to me. They were just piles of seemingly random osteology. The first time I dissected a bird neck was an epiphany; ever since then, it is hard for me to look at sauropod vertebrae and not see them clad in the diverticula and muscles that shaped their morphology. Go have fun.
- Schwarz, D., Frey, E., and Meyer, C.A. 2007. Pneumaticity and soft−tissue reconstructions in the neck of diplodocid anddicraeosaurid sauropods. Acta Palaeontologica Polonica 52(1):167–188.
- Tsuihiji, T. 2004. The ligament system in the neck of Rhea americana and its implications for the bifurcated neural spines of sauropod dinosaurs. Journal of Vertebrate Paleontology 24: 165–172.
- Wedel, M.J. 2003a. Vertebral pneumaticity, air sacs, and the physiology of sauropod dinosaurs. Paleobiology 29:243-255.
- Wedel, M.J. 2007a. What pneumaticity tells us about ‘prosauropods’, and vice versa. Special Papers in Palaeontology 77:207-222.
- Wedel, M.J. 2007b. Aligerando a los gigantes (Lightening the giants). ¡Fundamental! 12:1-84. [in Spanish, with English translation]
- Wedel, M.J. 2009. Evidence for bird-like air sacs in saurischian dinosaurs. Journal of Experimental Zoology 311A(8):611-628.
- 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.
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!
- Powell, Jaime E. 2003. Revision of South American Titanosaurid dinosaurs: palaeobiological, palaeobiogeographical and phylogenetic aspects. Records of the Queen Victoria Museum 111: 1-94.
SV-POW! is, as I’m sure you know, devoted to sauropod vertebrae. But occasionally we look at other stuff… and you might have noticed that, in recent months, we’ve been looking at, well, an awful lot of other stuff. I’m going to continue that theme here and talk about salamanders. Yeah: not sauropods, not sauropodomorphs, not saurischians, and not even dinosaurs or archosaurs. But salamanders. Don’t worry, all will become clear. This all started back in May 2010 when I blogged about amphiumas over at Tet Zoo. Amphiumas are very unusual, long-bodied aquatic salamanders.
As it happens, amphiuma vertebrae are particularly interesting if you work on saurischians because (drum-roll)… they have laminae. The term lamina is not restricted to structures present only in pneumatic saurischians: I would argue that it should be used for any sheet-like bony process on a vertebra, and I hope everyone agrees with me. Laminae are not common outside of Saurischia, but are present here and there: they’re present in stem-archosaurs (like Erythrosuchus), various crurotarsan archosaurs (including aetosaurs), some neosuchian crocodilians, and silesaurids (Desojo et al. 2002, Parker 2003, Nesbitt 2005, Wedel 2007, Butler et al. 2009). Even weirder, they’re present in Aneides lugubris, the Arboreal salamander of California and Baja California (Wedel 2007). But that’s about it.
Why would a salamander ‘want’ vertebral laminae? The laminae of the Arboreal salamander are presumed to be related to the extensive accessory ossification present in the skeleton of this animal, itself a consequence of adaptation to a peculiar climbing lifestyle. In other words, it’s hypothesised that the function (if I may be so bold as to use that word…) of the salamander’s laminae is nothing like that of the archosaurs that have them.
And now we know that A. lugubris isn’t the only salamander with laminae: amphiumas have them too. They’re clearly figured in the amphiuma literature (Gardner 2003), but (so far as I know) no-one has previously drawn attention to them when discussing archosaur laminae.
Gardner (2003) figured schematic amphiuma dorsal vertebrae that were based on a combination of features present in two of the three living amphiuma species (namely, Amphiuma means and A. tridactylum). On the lateral sides of the centra are structures that – if seen in an archosaur – would almost certainly be identified as anterior and posterior centrodiapophyseal laminae (using, as always, the nomenclature proposed by Wilson (1999)) [see the digram above, from Gardner (2003)]. There are also structures on the dorsal surfaces of the postzygapophyses that look something like laminae: they extend from the posterolateral parts of the neural arch and run across the tops of the postzygapophyses, hence recalling spinopostzygapophyseal laminae. Actually, I’ve just realised that similar structures are also sometimes present in anurans (frogs and toads) where they’ve been called paraneural crests or paraneural processes. These structures do have a ‘known’ function: in amphiumas they’re associated with complex dorsalis trunci epaxial muscles. Unlike the spinopostzygapophyseal laminae of saurischians, the structures in the amphibians are low ridges that don’t contact the neural spines, so it could be argued that they aren’t so lamina-like after all.
But what about the structures on the sides of the centra? Why are laminae present in a group of long-bodied aquatic salamanders? Why are laminae present at all? This question has been asked a few times here on SV-POW! (here, for example), and there are two primary hypotheses. One is that the laminae keep the various air sacs separate from each other, perhaps because they persist while much of the bone around them is resorbed during ontogeny, while the other is that they somehow provide mechanical support and are aligned along lines of stress (for more on this subject see the piece on finite element analysis).
The pneumaticity explanation can’t work for amphiumas (given that they’re apneumatic): does the ‘mechanical support’ one apply instead? We don’t know anything about stress distribution in amphiuma vertebrae – in fact, I don’t think we know anything about the mechanics of amphiumas at all – but it’s possible that the laminae might play this role, especially given that amphiumas have to bend, twist and push with their bodies while excavating burrows.
In conclusion, we just don’t really know what’s going on here. In fact, all we can really do at the moment is wave our arms around a bit and say “Hey, amphiumas have vertebral laminae, too”, and that’s pretty much all I’m doing here. It’s also possible that the structures I’m talking about in amphiumas are very different in detail from the vertebral laminae present in archosaurs: I’ve never even seen a single amphiuma skeletal element and am basing all of this on photos and diagrams in the literature. Nevertheless, it’s something definitely worth bringing attention to. As usual, we stand poised at the abyss, straining our eyes to see into the infinite darkness ahead.
Butler, R. J., Barrett, P. M. & Gower, D. J. 2009. Postcranial skeletal pneumaticity and air-sacs in the earliest pterosaurs. Biology Letters 5, 557-60.
Desojo, J. B., Arcucci, A. B. & Marsicano, C. A. 2002. Reassessment of Cuyosuchus huenei, a Middle–Late Triassic archosauriform from the Cuyo Basin, west-central Argentina. Bulletin of the New Mexico Museum of Natural History and Science 21, 143–148.
Gardner, J. D. 2003. The fossil salamander Proamphiuma cretacea Estes (Caudata; Amphiumidae) and relationships within the Amphiumidae. Journal of Vertebrate Paleontology 23, 769-782.
Nesbitt, S. J. 2005. Osteology of the Middle Triassic pseudosuchian archosaur Arizonasaurus babbitti. Historical Biology 17, 19–47.
Parker, W. G. 2003. Description of a new specimen of Desmatosuchus haplocerus from the Late Triassic of northern Arizona. Unpublished MS thesis, Northern Arizona University, Flagstaff, AZ, 312 pp.
Wedel, M. J. 2007. What pneumaticity tells us about ‘prosauropods’, and vice versa. Special Papers in Palaeontology 77, 207-222.
Wilson, J. A. 1999. A nomenclature for vertebral laminae in sauropods and other saurischian dinosaurs. Journal of Vertebrate Paleontology 19, 639-653.
March 23, 2010
For various arcane reasons, the SV-POW!sketeers are all neck-deep in work, so the blog may actually become somewhat more of the APOD-style picture-n-paragraph thing we originally envisioned, and less of the TetZoo-style monograph-of-the-week thing it’s often leaned toward, at least for a while.
I like it when people decorate their papers with megapixels of vertebral goodness, so here are some caudal vertebrae of the African diplodocine Tornieria, from Remes (2006:fig. 5). Click through to see the figure at its massive native resolution. And check out that pneumaticity! Really, the only question about this image is whether you can settle for just using it as your desktop background, or if you need to print out a wall-sized poster for your bedroom. So the next time you see Kristian Remes, buy him a beer for doing solid work here, on the Humbolt sauropod remount, and on pretty much everything else (including this).
Remes, K. 2006. Revision of the Tendaguru sauropod Tornieria africana (Fraas) and its relevance for sauropod paleobiogeography. Journal of Vertebrate Paleontology 26 (3): 651–669.
January 18, 2010
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.
- 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.
January 4, 2010
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!
- 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.
December 7, 2009
Broadly speaking, pneumatic sauropod vertebrae come in two flavors. In more primitive, camerate vertebrae, modeled here by Haplocanthosaurus, the centrum is a round-ended I-beam and the neural arch is composed of intersecting flat plates of bone called laminae (lam above; fos = fossa, nc = neural canal, ncs = neurocentral suture; Ye Olde Tyme vert pic from Hatcher 1903).
In more derived, camellate vertebrae, the centrum and neural arch are both honeycombed with many small air spaces. This inflated-looking morphology is very similar to that seen in birds, like the turkey we recently discussed. The fossae and foramina on the outside tend to be smaller and more numerous than in camerate vertebrae, as shown here in a titanosauriform axis from India (Figure 3 from Wilson and Mohabey 2006). The green arrows show that the fossae visible on the external surface are excavations or depressions into the honeycombed internal structure of the bone.
External fossae on bones can house many different soft tissues, including muscles, pads of fat or cartilage, and pneumatic diverticula (O’Connor 2006). Pneumatic fossae are often strongly lipped and internally subdivided and may contain pneumatic foramina, which makes them easier to diagnose (but they may also be simple, smooth, and “blind”, which makes them harder to diagnose as pneumatic). But in all of these cases we are usually talking about the same thing: a visible excavation into a corpus of bony tissue, which may have marrow spaces inside if it is apneumatic, or air spaces inside if it is pneumatic (the corpus of bone, not the dent). That’s probably how most of us think about fossae, and it would hardly need to be explained…except that sometimes, something much weirder happens.
Consider this cervical of Brachiosaurus (this is BYU 12866, from Dry Mesa, Colorado). Brachiosaurus and Giraffatitan have an in-between form of vertebral architecture that my colleagues and I have called semicamellate (Wedel et al. 2000); the centrum does have large simple chambers (camerae), but smaller, thin-walled camellae are also variably present, especially along the midline of the vertebra and in the ends of the centrum. As in Haplocanthosaurus, the neural arch is composed of intersecting plates of bone; unlike Haplocanthosaurus, these laminae are not flat or smooth but are instead highly sculpted with lots of small fossae. Janensch (1950) called these “Aussenkaverne”, or accessory outside cavities, because and they are smaller and more variable than the large fossae and foramina that invade the centrum.
And that’s the weird thing. As the red arrows in the above image show, the “Aussenkaverne” are not excavations or depressions into anything, except the space on the other side of the lamina (which in life would have been occupied by another diverticulum). The neural arches of Brachiosaurus and Giraffatitan are not excavated by fossae, they’re embossed, like corporate business cards and fancy napkins.
What’s up with that!? We tend to think of pneumaticity as reducing the mass of the affected elements, but the shortest distance between two vertebral landmarks is a smooth lamina. These embossed laminae actually require slightly more bony material than smooth ones would.
As you can see above, the outer edges of the laminae are thick but the bone everywhere else is very thin. Maybe, like the median septa in pneumatic sauropod vertebrae, the thin bone everywhere except the edges of the laminae was just not loaded very much or very often, and was therefore free to get pushed around by the diverticula on either side, in the sense of being continually and quasi-randomly remodeled into shapes that don’t strike us as being very mechanically efficient. But also like the median septa, the thin parts of the laminae are only rarely perforated (but it does happen), for possible (read: arm-wavy) reasons discussed in the recent FEA post. And maybe the amount of extra bone involved in making embossed laminae versus smooth ones was negligible even by the very light standards of sauropod vertebrae.
Another question: since these thin sheets of bone were sandwiched in between two sets of diverticula, why are the “unfossae” always embossed into them, in the medial or inferior direction? Why don’t any of them pop out laterally or dorsally, looking like domes or bubbles instead of holes, like Mount Fist-of-God from Larry Niven’s Ringworld? Did the developmental program get accustomed to making fossae that went down and into a corpus of bone, and just kept on with business as usual even when there was no corpus of bone to excavate into? I’m only half joking.
I don’t have good answers for any of these questions. I scanned this vert a decade ago and I only noticed how weird the “unfossae” were a few months ago. I’m putting all this here because “Hey, look at this weird thing that I can only wave my arms about” is not a great basis for a peer-reviewed paper, and because I’d like your thoughts on what might be going on.
In Other News
The Discovery Channel’s Clash of the Dinosaurs premiered last night. I would have given you a heads up, except that I didn’t get one myself. I only discovered it was on because of a Facebook posting (thanks, folks!).
COTD is intended to be the replacement, a decade on, for Walking With Dinosaurs. I’m happy to report that one of the featured critters is Sauroposeidon. I grabbed a couple of frames from the clips posted here.
I haven’t seen the series yet, because I don’t have cable. But I’m hoping to catch it at a friend’s place next Sunday night, Dec. 13, when the entire series will be shown again. With any luck, I’ll have more news next week.
Finally, I got to do an interview at Paw-Talk, a forum for all things animal. I’m very happy with how it turned out, so thanks to Ava for making it happen. While you’re over there, have a look around, there’s plenty of good stuff. Brian Switek, whom you hopefully know from this and this, is a contributor; check out his latest here.
- Hatcher, J.B. 1903. Osteology of Haplocanthosaurus, with a description of a new species, and remarks on the probable habits of the Sauropoda, and the age and origin of Atlantosaurus beds. Memoirs of the Carnegie Museum 2:1–72.
- Janensch, W. 1950. Die Wirbelsaule von Brachiosaurus brancai. Palaeontographica (Suppl. 7) 3:27-93.
- O’Connor, P.M. 2006. Postcranial pneumaticity: an evaluation of soft-tissue influences on the postcranial skeleton and the reconstruction of pulmonary anatomy in archosaurs. Journal of Morphology 267:1199-1226.
- Wedel, Mathew J., Richard L. Cifelli and R. Kent Sanders. 2000. Osteology, paleobiology, and relationships of the sauropod dinosaur Sauroposeidon. Acta Palaeontologica Polonica 45(4): 343-388.
- Wilson, J. A. and Mohabey, D. M. 2006. A titanosauriform axis from the Lameta Formation (Upper Cretaceous: Maastrichtian) of central India. Journal of Vertebrate Paleontology 26:471–479.
November 26, 2009
Trying two new things this morning: grilling a turkey, and live-blogging on SV-POW!
I like to grill. Steak, chicken, kebabs, yams, pineapple, bananas–as long as it’s an edible solid, I’m up for it. But I’ve never grilled a turkey before. Neighbor, colleague, fellow paleontologist and grillmeister Brian Kraatz sent me his recipe, which is also posted on Facebook for the edification of the masses. See Brian’s excellent writeup for the whole process, I’m just going to hit the photogenic parts here. Oh, and usually I tweak any photos I post within an inch of their lives, but I don’t have time for that this morning, so you’re getting as close to a live, unedited feed as I can manage. Stay tuned for updates.
Enough of that. Let’s rock!
The process starts more than a day in advance, with the brine. Salt water, fruit, onions, garlic, spices, and some apple juice.
The turkey needs to be entirely immersed in the brine for at least 24 hours. Doing this in a solid container would require an extra big container and too much liquid to cover the bird. I follow Brian’s method of brining in a triple-layer of trash bags. You can see a turkey roaster peeking out underneath the trash bags. Helps with the carrying.
Put the turkey in the trash bags first, then pour in the brine. Unless you like huge messes.
The genius of the trash bag method on display. You can squeeze out all the air so that the volume of the bag is equal to just the turkey and the brine.
Into the fridge for a day.
First thing this morning: out come the giblets, and save the goodies from the brine. We’ll get back to the neck later.
The bird awaits.
Crucial step: putting in a drip pan. Keeps the coals off to the side for indirect heat, and catches the grease so you don’t burn down the neighborhood.
Putting in the herb butter. I used three short sticks of butter mixed with sage, lemon pepper, and Mrs. Dash. Working the skin away from the meat and then filling the space with butter was extremely nasty. This must be what diverticula feel like.
A chimney is helpful to get the coals going.
To eat is human; to grill is divine.
Smoke bombs: mesquite chips soaked in water, wrapped up in balls of tinfoil, with holes poked on top to let the smoke out.
Fruit and spices into the body cavity.
At this point, I was fairly certain that today would be the greatest day of my life. The turkey is centered over the drip pan, stuffed with goodness, subcutaneously loaded with herb butter, draped with bacon. You can see one of the smoke bombs sitting right on top of the coals.
Know what you’re getting into. This 15 lb bird just barely cleared the lid of my grill.
A little over an hour in. I installed foil heat shields to keep the wings and thighs from cooking too fast. It’s all about the indirect heat. Some of the bacon comes off now, as a mid-morning treat.
Okay, the bird is about halfway done, and I have to whip up some sustainer coals and another batch of smoke bombs. Further updates as and when. Happy Thanksgiving!
I was hoping to get some more pictures posted before we ate, but you know how it is in the kitchen on Thanksgiving Day (or, if you’re not an American, maybe you don’t know, so I’ll tell you: dogs and cats living together, we’re talking total chaos).
The turkey just before I pulled it off the grill. The heat shields turned out to be clutch, I would have completely destroyed the limbs without them. That’s going to be SOP from now on.
Ah yes, the bird, she turned out even more succulent than I hadda expected. Check out the pink shade of the meat just below the skin. I recognize that, from good barbeque, but I’ve never produced it before.
That’s it for the cooking part of today’s program. As for the ultimate fate of the bird…we ate a stupifying amount of it. I sent even more home with our guests. And the other half–yes, half–of this thunder beast is sitting in the fridge. Hello-o leftovers!
And hello-o science!
I was going to post some more pictures of the neck, but I didn’t get around to eating it, so…another time, perhaps. In lieu, here’s Mike’s turkey vertebra in left lateral view (see the original in all its supersized glory here). Note the pneumatic foramen in the lateral wall of the centrum, just behind the cervical rib loop. This is actually kind of a lucky catch; a lot of times with chickens and turkeys, the pneumatic foramina are so far up in the cervical rib loop that they can’t be seen in lateral view.
It used to freak me out a little bit that birds often don’t have their pneumatic foramina in the middle of the lateral wall of the centrum, like sauropods. But a possible explanation occurred to me just this morning as I was planning this post. I think that birds have their pneumatic foramina right where you’d expect them, based on sauropods. I’ll explain why.
The first part of the explanation is that instead of wearing their pneumatic cavities on the outside, like this Giraffatitan cervical, bird vertebrae tend to be inflated from within, with just a few tiny foramina outside. The second part is that birds have HUGE cervical rib loops compared to sauropods. If the sauropod vert shown above had its rib on, the resulting loop would be fairly dainty, the osteological equivalent of a bracelet. The cervical rib loops of birds are more like tubes, they’re so antero-posteriorly elongated.
So take the brachiosaur cervical shown above and shrink all of the external pneumatic spaces by several inches. The cavities on the arch and spine would close up entirely, and the complex of fossae and foramina on the lateral side of the centrum would be reduced to a small hole right behind the cervical rib. Then stretch out the cervical rib loop in the fore-aft direction and voila, you’d have something like a turkey cervical, with a little tiny pneumatic foramen tucked up inside the cervical rib loop.
This doesn’t explain why bird verts are inflated from within instead of being eroded from without, or why sauropods had such dinky cervical rib loops (mechanical what, now?), or why pneumatic diverticula tend to make the biggest holes in the front half of the centrum, adjacent to the cervical ribs. I just think that maybe bird and sauropod pneumaticity are not as different as they appear at first glance. Your thoughts are welcome.
November 24, 2009
I drew a couple of these a while back, and I’m posting them now both to fire discussion and because I’m too lazy to write anything new.
Here’s the neck of Apatosaurus, my own reconstruction based on Gilmore (1936), showing the possible paths and dimensions of continuous airways (diverticula) outside the vertebrae.
Here’s figure 4 from Lovelace et al. (2007), which first got me thinking about pneumatic traces on the ventral surfaces of the centra and what they might imply. You can see pneumatic spaces between the parapophyses in Supersaurus (A) and Apatosaurus (C) but not in Barosaurus (B).
This is another of my moldy oldies, again based on one of Gilmore’s pretty pictures, showing how I think the soft tissues were probably arranged. The muscles are basically the technicolor version of Wedel and Sanders (2002). Two points:
- How bulky you make the neck depends mainly on how much muscle you think was present (which of course depends on how heavy you think the neck was…). Here I was just trying to get the relationships right without worrying about bulk, but it’s worth considering.
- The volume of air inside the vertebra was dinky compared to the probable volume of air outside. In Apatosaurus, either of the canals formed by the transverse foramina has almost twice the cross-sectional area of the centrum.
A fair amount of this has been superseded with better data and prettier pictures by Schwarz et al. (2007), so don’t neglect that work in any ensuing discussion (it’s free, fer cryin’ out loud). And have a happy Thanksgiving!
- Gilmore, C.W. 1936. Osteology of Apatosaurus with special reference to specimens in the Carnegie Museum. Memoirs of the Carnegie Museum 11: 175-300.
- Lovelace, D.M., Hartman, S.A., and 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.
- Schwarz, D., Frey, E., and Meyer, C.A. 2007. Pneumaticity and soft−tissue reconstructions in the neck of diplodocid and dicraeosaurid sauropods. Acta Palaeontologica Polonica 52 (1): 167–188.
- 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.
Mike asked me to add the labeled version of Nima’s brachiosaur parade, so here you go. Click to embiggen.