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Another raw photo from the road.

The Morrison fossils from the Oklahoma panhandle were dug up and prepped out by  WPA workers in the 1930s, and their preparation toolkit consisted of hammers, chisels, pen-knives, and sandpaper. (Feel free to take a minute if you need to get your nausea under control.) And whereas most Morrison fossils are much darker than the surrounding matrix, in the Oklahoma panhandle the bone and matrix are about the same color. Sometimes the prep guys didn’t know they’d gone too deep until they sanded into the trabecular bone. Or in this case, into the air spaces in the condyle of this anterior dorsal of Apatosaurus.

Still, we have lots of anterior dorsals of Apatosaurus, and very few we can see inside, and they’re too darned big to scan, so this gives us useful information that a more perfect specimen would not. So I salute you, underemployed dude from eighty-odd years ago. Thanks for showing me something cool.

No time for anything new, so here’s a post built from parts of other, older posts.

The fourth sacral centrum of Haplocanthosaurus CM 879, in left and right lateral view. This is part of the original color version of Wedel (2009: figure 8), from this page. (Yes, I know I need to get around to posting the full-color versions of those figures. It’s on my To Do list.)

Note the big invasive fossa on the right side of the centrum. The left side is waisted (narrower at the middle than the ends) like most vertebrae of most animals, but has no distinct fossa on lateral face of the centrum. What’s up with that? Here’s an explanation from an old post (about another sauropod) that still fits:

Now, this asymmetry is also weird, but it’s expected weirdness. Pneumaticity seems to just be inherently variable, whether we’re talking about human sinuses or the facial air sacs of whales or the vertebrae of chickens. It appears that the form of pneumatic features is entirely determined by local tissue interactions, with little or no genetic control of the specific form. Think of it this way: genes prescribe certain developmental events, and those events bring tissues into contact–such as pneumatic epithelium and bone. The morphology of the bone arises out of that interaction, and each interaction of bone and pneumatic epithelium has the potential to produce something new. In this case, the diverticula on the left side of the vertebral column come from the lungs or air sacs on the left, and those on the right side come from the lungs or airs sacs on the right, so it’s really two sets of diverticula contacting the bone independently. The wonder, then, is not that pneumatic bones are so variable, but that we see any regularities at all.

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.

Those ostrich necks I went to Oro Grande to get last Thursday? Vanessa and I started dissecting them last Friday. The necks came to us pre-cut into segments with two to three vertebrae per segment. The transverse cuts were made without regard for joints so we got a bunch of cross sections at varying points through the vertebrae. This was fortuitous; we got to see a bunch of cool stuff at the cut faces, and those cut faces gave us convenient avenues for picking up structures and dissecting them out further.

In particular, the pneumatic diverticula in the neck of this ostrich were really prominent and not hard at all to see and to follow. The photo above shows most of the external diverticula; click through for the full-resolution, unlabeled version. The only ones that aren’t shown or labeled are the diverticula around the esophagus and trachea (which had already been stripped off the neck segments, so those diverticula were simply gone), those around carotid arteries, which are probably buried in the gloop toward the bottom of the photo, and the intermuscular diverticula, of which we found a few in parting out the dorsal and lateral neck muscles.

There is one final group of diverticula that are shown in the photo but not labeled: the interosseous diverticula that fill the air spaces inside the bone.

We have tons of cool photos from this dissection, so expect more posts on this stuff in the future.

For previous posts showing diverticula in bird neck dissections, see:

Things to Make and Do, part 7: fun with rhea necks

Things to Make and Do, part 7b: more fun with rhea necks (admittedly, not the most creative title ever)

We’re starting the new year with a new feature, in which we answer questions that have come our way. We never had a policy about not answering questions, it’s just that previous ones have tended to arrive in the comments section and have been dealt with there. But suddenly in the last few days I’ve gotten two questions from extrabloggular sources, and rather than hide the replies I thought I’d make them available to all.

One of my cohort at Berkeley texted me the other day with the following questions:

OK, phylobuddy: can you suck the marrow from a chicken bone? If they have hollow bones, where’s the marrow?!? Google is getting me nowhere.

Short answer: yes, one can get marrow from chicken bones, from those bones that contain marrow rather than air. In most fully mature chickens, the pneumatic bones include the braincase, the cervical, dorsal, and most or all synsacral vertebrae, some of the dorsal ribs, the central portion of the sternum, the coracoids, and the humeri (if you’re not a regular and some of these terms are unfamiliar, check out these handy guides [1, 2] to the vertebrate skeleton). That leaves marrow in everything else, although the only bones with large marrow cavities–as opposed to tiny trabecular spaces, which also house marrow–are the radii, ulnae, femora, tibiotarsi, and tarsometatarsi. So if you want to actually see large amounts of chicken marrow, or suck the marrow out of chicken bones, you’re basically stuck with the big distal bones of the wing, the thigh, and the drumstick (tibiotarsus). If you are boiling chicken bones to get stock for soups or stews, might as well throw them all in; even the pneumatic bones will still have bits of adhering meat, cartilage, and ligaments that will give up molecules and flavor to the stock.

The long answer is that the expression “hollow bones” has caused no end of confusion, because there are at least two ways to interpret hollow: filled with air, or not filled with bone (the former is a subset of the latter). If you mean “not filled with bone”, then the bones of almost all amniotes* are hollow, and the spaces inside are occupied by marrow (most commonly) or air. If filled with air, the bones are referred to as pneumatic, and an accessible introduction to them is here.

* At least; I know less about amphibians and fish, although at least one osteoglossomorph (IIRC) pneumatizes its vertebrae from its swim bladder!

The reasons it gets confusing are twofold. First, sometimes authors describe bones as hollow and mean only that they have chambers inside, but later readers see ‘hollow’ and infer ‘pneumatic’. Not all hollow bones are pneumatic; in fact, the vast majority of them are not, including the long bones of your arms and legs. The criteria for inferring pneumaticity from dry bones are more strict, and are explored in this paper and this one. Anyway, this point is just confusion caused by an ambiguous term.

The second case is more interesting, because it involves real unknowns. In the fossil record we can almost always tell if a bone is hollow, sensu lato, but sometimes it is not possible to say for certain whether the hollow space(s) inside were filled with marrow or air. Particularly vexing and intriguing examples include the humerus of Eotyrannus and the iliac chambers of some sauropods, which are discussed in this paper. My guess is that the iliac chambers of sauropods are genuinely pneumatic, because they only occur in sauropods that already have sacral pneumaticity, and we know from broken ilia of more basal sauropods and sauropodomorphs that large marrow-filled chambers are not present in those taxa. Conversely, I suspect that the humerus of Eotyrannus was apneumatic (marrow-filled), given that humeral pneumaticity is otherwise unknown in non-avian theropods, although the pneumatic furcula of Buitreraptor at least shows that the necessary clavicular air sac was present in some.

Next question! This one came to me on Facebook, from ReBecca Hunt-Foster, whom you may know from her awesome Dinochick Blogs. You should also envy her and hubby John Foster for getting the most awesome wedding present of all time: a 1/12 scale skeleton of Apatosaurus sculpted by Phil Platt, which you can read about here. That’s cool enough that I am stealing it for this otherwise picture-challenged post.

ANYWAY, ReBecca wrote on my FB wall today to ask:

Random question: Have you seen many tooth marks on sauro cervical verts? I am debating on whether something I have is a dessication crack or really some tooth marks. Thanks :)

In all the 15 years that I have spent looking at sauropod remains in the bowels of many, many museums, I have never seen a single tooth mark on a sauropod vertebra.

[Update the next day: Er, except for the bitten Apatosaurus tail on display in the AMNH! Many thanks to reptilianmonster and steve cohen for reminding me about this in the comments. I'm going to go hide for a while now.]

Now, that doesn’t mean that they aren’t there. Truth be told, I’ve never looked for them, and my usual mental search pattern for pneumatic traces (large, irregular) would probably exclude tooth scratches (small, linear) as noise. But I’ve certainly never seen any vertebrae with easily recognizable signs of predation or scavenging or with obvious bites removed.

People also sometimes ask me what kinds of healed traumas I’ve seen in pneumatic sauropods bones. That’s easy: apart from vertebral fusions, most of which probably have nothing to do with trauma, I’ve seen zip. Nada. Null set. The wingspan of the average tadpole. I’ve seen some pretty cool pneumatic bones from extant birds that were broken and later healed, including a eagle femur in the UCMP comparative collection that is now shaped like the letter Z, but nothing in sauropods.

I can think of three possible reasons for this, which sort of flow into each other. The first is that apart from the very solid and blocky centra of apneumatic vertebrae, sauropod verts were pretty fragile, and prone to getting distorted and busted up even when they started out intact, and those verts that started out broken just had a tougher time with the taphonomic lottery.

The second is that pneumatic sauropod bones would been nothing to most predators other than a mouthful of relatively dry bone shards, so either carnivores left them alone, or if they were osteovores like T. rex, they ate the shards and whatever is left over is unrecognizable. I have seen, and mostly ignored, plenty of vert-shrapnel in quarries and in collections, and maybe sharper eyes than mine could have discerned evidence of predation from those bits. To me it mostly looked like trampling, hydraulic transport, erosion, and other mundane ways to explode a vertebra.

The third is that in addition to a preservation bias against half-destroyed verts, there is probably also a collection bias against them. I’m probably not the only one would pass up a few shards of excellence to dig out the complete fibula sitting next to them in the quarry, and I love this stuff. That said, we did get a LOT of blasted vert bits out of the Wolf Creek quarry in the Cloverly, so if you want to pore over sauropod shards looking for tooth marks, visit the OMNH.

And, if you do know of tooth marks on sauropod vertebrae, please let us know in the comments. And consider publishing them, given the apparent vacuum of such things.

Wedel’s Theorem:

freezer full of interesting dead animals + great anatomy student who actually wants to get up on Saturday morning and dissect = happiness

The rhea has been the gift that keeps on giving. Saturday was my fourth session with some part of this bird, going back to 2006 (previous posts are here, here, and here). The first two sessions were just about reducing the bird to its component parts, and the last session was all about midline structures.

The goal for the neck is to dissect down to the vertebrae and document everything along the way–muscles, tendons, fascia, blood vessels, and especially diverticula. In the past I have been pessimistic about the chances of seeing diverticula without having them injected with latex or resin or something. But this bird is changing my mind, as we saw in a previous post and as you can see below.

The goal for Vanessa is to grok all of this anatomy, and hopefully make some publishable observations along the way. She has a chance to do something that I think is rather rare for a sauropod paleobiologist, which is to get a firm, dissection-based grounding in bird and croc anatomy before she first sets foot in a museum collection to play with sauropod bones.

That sounds awesome, and probably will be awesome, but before there can be any awesomeness, the fascia has to be picked off the neck. And by ‘picked’ I mean ‘actually cut away, millimeter by arduous millimeter’. It wasn’t that bad everywhere–the fascia over the long dorsal muscles came off very easily. But the lateral neck muscles were actually originating, in part, from the inner surface of the fascia. That’s not unheard of, it happens in the human forearm and leg all the time, but I’ve never seen it as consistently as in this rhea. So picking fascia took a loooong time–that’s what Vanessa is doing in the photo at top.

Once the fascia was off, Vanessa started parting out the long tendons of the hypaxial muscles in the left half of the neck. Meanwhile, I started stripping fascia from the right half. I had forgotten that the right half of the neck still had the trachea and esophagus adhered to the side. That probably sounds weird, given that our trachea and esophagus–and those of most mammals–run right down the middle of our necks and aren’t free to move around much. In birds, they’re more free-floating and can drift around between the skin and the vertebral muscles, sometimes even ending up dorsal to the  vertebral column–there’s a great x-ray of a duck in a  2001 paper that shows this, which I’ll have to blog sometime.

Anyway, when I cut the fascia to pull back the trachea and esophagus, I found that they were separated from the underlying tissues by a dense network of pneumatic diverticula winding through the fascia.

I had heard, anecdotally, of networks of diverticula described as looking like bubble wrap. I can now confirm that is true, for at least some networks. What was especially cool about these is that they were occupying space that would be filled with adipose or other loose connective tissue in a mammal, which illustrates the point that pneumatic epithelium seems to replace many kinds of connective tissue, not just bone–something Pat O’Connor has talked about, and which I also briefly discussed in this post.

I should mention that there was no connection between these diverticula and the trachea, as there is between the subcutaneous throat sac and the trachea in the emu (story and pictures here).

While I was geeking out on diverticula, Vanessa was methodically separating the long hypaxial muscles, which looked pretty cool all fanned out.

And that’s all we had time for on Saturday. But we’re cutting again soon, so more pictures should be along shortly.

Busy days. I just published a popular article on skeletal pneumaticity as a web feature at the Australian science magazine Cosmos. It’s entitled, “We are all air-heads: of sinus headaches and strangled birds”, and it includes a few things I don’t think I’ve discussed here, so hopefully even you regulars will find it a worthwhile read. I’d tell you more about it, but that would defeat the point, wouldn’t it? Go on over and check it out.

While you’re there, look at all the cool articles by award-winning science blogger and Cosmos Editorial Assistant Bec Crew, who served as my editor in this venture. I’m grateful to Bec for her help getting the article bashed into shape, her patience with my own article revision incontinence (don’t laugh, some writer you know might suffer from ARI), and most of all her enthusiasm where gory tales of science are concerned. If you’re not familiar with Bec’s work at Save Your Breath for Running Ponies, you’re in for a treat. Set your drink down first so you don’t spew it on the keyboard laughing.

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.

BUT.

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.

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.

References

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.
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