Update

This is an actual page from the late, lamented Weekly World News, from December 14, 1999. I always thought it was pretty darned funny that they had the alien remains discovered in the “belly” of an animal known only from neck vertebrae. Now, subjecting a tabloid story to technical scrutiny really is like dancing about architecture, but…it just tickles me. As does the entire story. I haven’t been able to get hold of Dr. Posvby to confirm his findings, but it’s been over a decade and he still hasn’t published, so I’m not holding my breath.

Incidentally, the WWN archives are available on Google Books: go here to read about Bat Boy siring a 3-headed alien Elvis baby on a female Sasquatch. Or something to that effect.

In an email, Vladimir Socha drew my attention to the fact that Tom Holtz’s dinosaur encyclopaedia estimates the maximum height of Sauroposeidon as 20 meters plus, and asked whether that was really possible.  Here’s what Tom actually wrote: “Sauroposeidon was one of the largest of all dinosaurs.  At perhaps 98 to 107 feet (30 to 32.5 meters) long and weighing 70 to 80 tons […] Sauroposeidon would have been the tallest of all dinosaurs. […] If it could crane its neck up, it might have been able to hold its head 66 to 69 feet (20 to 21 meters) high or more” (Holtz and Rey 2007:207).  Vladimir was understandably skeptical.  But can it be true?

Wedel and Cifelli (2005: fig. 15) shows Matt’s best skeletal reconstruction of Sauroposeidon, with Boring Old Brachiosaurus and a human for scale:

Sauroposeidon with Boring Old Brachiosaurus and human for scale and 20 m height indicated. Lightly modified from Wedel and Cifelli (2005: fig. 15)

Sauroposeidon with Boring Old Brachiosaurus and human for scale and 20 m height indicated. Lightly modified from Wedel and Cifelli (2005: fig. 15)

Amazingly, those dummies didn’t include an actual scalebar; but apparently the human figure is 1.8 m tall, so by measuring pixels and cross-scaling, I determined that in this image, Sauroposeidon is a mere 13.43 m tall.  I took the liberty of adding in a marker for the 20 m height proposed by Holtz, and as things stand you’d have to say that it doesn’t look probable.

But let’s see what we can do.  We’ll begin with the classic brachiosaur skeleton of Paul (1988), which shows the well represented species Brachioaurus brancai:

Brachiosaurus brancai skeletal reconstruction in left lateral view. From Paul (1988:fig. 1)

Brachiosaurus brancai skeletal reconstruction in left lateral view. From Paul (1988:fig. 1)

(Some other time, we should take a moment to discuss the differences between this and the Wedel brachiosaur reconstruction; but it will not be this day.)

This reconstruction is in a nice erect-necked posture which, in light of our own recent paper, is probably not too extreme.  Since all the neural arches and processes are missing from the only known posterior cervicals of this species, we don’t know how much flexibility they allowed, and so in light of how the rest of the animal is built (high shoulders and all) it seems reasonable to allow a lot of extension at the base of the neck.  So let’s assume that the pose offered by Paul is correct.  By measuring my scan of that figure, and I see that the 2.13 m humerus is 306 pixels long.  The entire reconstruction, from tip of cranial crest down to forefoot, is 1999 pixels tall, which is 1999/306 = 6.53 times as long as the humerus, which scales to 6.53*2.13 = 13.91 m — a little taller than Sauroposeidon (not Brachiosaurus) in Matt’s reconstruction, which seems about right if we imgine Matt’s Brachiosaurus raising its neck into a Paul-compliant posture.

Now Paul’s reconstruction is based on the Berlin mounted skeleton HMN S II.  Cervical 8 is very well preserved in that animal, and has a centrum length of 98 cm (Janensch 1950a:44).  By contrast, the centrum of C8 of Sauroposeidon OMNH 53062 (the only known specimen) is 125 cm long (Wedel et al. 2000a: 110). So if Sauroposeidon is merely an elongated Brachiosaurus brancai, then it’s 125/98 = 1.28 times as long and tall, which would be 17.74 m.

But wait: it seems that Sauroposeidon is to Brachiosaurus brancai as Barosaurus is to Diplodocus — similar overall but more elongate.  And it turns out that Barosaurus has at least 16, maybe 17 cervicals (McIntosh 2005:45) compared with Diplodocus‘s 15.  So maybe Sauroposeidon also added cervicals from the brachiosaur base-state — in fact, that would hardly be surprising given that Brachiosaurus brancai has so few cervicals for a long-neck: 13, compared with 15 in most diplodocids, 16 or 17 in Barosaurus, and 19 in Mamenchisaurus.  If you reconstruct Sauroposeidon with two more C8-like cervicals in the middle of the neck, that adds 2*125 = 250 cm, which would give us a total height of 17.74+2.5 = 20.24 m.

So I don’t think Tom Holtz’s estimate is completely unrealistic, even for the one Sauroposeidon specimen we have now — and remember that the chances of that individual being the biggest that species got are vanishingly small.

On the other hand, maybe Sauropodseidon‘s neck was the only part of it that was elongated in comparison to Brachiosaurus brancai — maybe its forelimbs were no longer than those of its cousin, so that only the neck elongation contributed to greater height.  And maybe it had no additional cervicals, so its neck was “only” 1.28 times as long as that of Brachiosaurus brancai — 1.28*8.5 = 10.88 m, which is 2.38 m longer; so the total height would be 13.91+2.38 = 16.29 m (assuming the additional neck length was vertical).  And maybe the neck couldn’t get very close to vertical, so that the true height was lower still.

All of this just goes to show the perils of reconstructing an animal based only on a sequence of four cervicals.  (Reconstructing on the basis of a single partial mid-to-posterior dorsal, on the other hand, is a much more exact science.)

Finally: Matt’s reconstruction of Sauroposeidon is really rather conservative, and looks very much like a scaled-up vanilla brachiosaur.  Just to see how it looks, I’ve made a reconstruction of the putative (and very possible) elongated, attenuated version of Sauroposeidon, showing the legs and cervicals 28% longer than in B. brancai, and with two additional cervicals.  I made this by subjecting Greg Paul’s 1988 brachiosaur to all sorts of horrible and half-arsed distortions, so apologies to Greg.  But remember, folks: this is just as likely correct as Matt’s version!

A different view of Sauroposeidon, based on elongation of the cervicals and legs of Brachiosaurus brancai and the insertion of two additional cervicals. Heavily and carelessly modified from Paul (1988: fig. 1)

A different view of Sauroposeidon, based on elongation of the cervicals and legs of Brachiosaurus brancai and the insertion of two additional cervicals. Heavily and carelessly modified from Paul (1988: fig. 1)

References

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Internal structure of a cervical vertebra of Sauroposeidon, OMNH 53062. A, parts of two vertebrae from the middle of the neck. The field crew that dug up the bones cut though one of them to divide the specimen into manageable pieces. B, cross section of C6 at the level of the break, traced from a CT image and photographs of the broken end. The left side of the specimen was facing up in the field and the bone on that side is badly weathered. Over most of the broken surface the internal structure is covered by plaster or too damaged to trace, but it is cleanly exposed on the upper right side (outlined). C, the internal structure of that part of the vertebra, traced from a photograph. The arrows indicate the thickness of the bone at several points, as measured with a pair of digital calipers. The camellae are filled with sandstone.

Image and caption recycled from fig. 14 here. Hat tip to Mike from Ottawa for the wonderful title.

Addendum (from Mike)

What Matt’s failed to mention is that this section of prezygapophyseal ramus is one of the elements for which he calculated the Air-Space Proportion (ASP) in his chapter in “The Sauropods”. As shown in his table 7.2, this calculation yielded 0.89.  Just think about that for a moment.  89% of the bone was air.  Yikes.

It’s interesting that this was the only prezygpapophyseal ramus in the survey, and that it had a way higher value that any of the other elements considered, which topped out at 0.77, i.e., more than twice as much bone as this specimen.  So maybe all prezyg rami are ridiculously pneumatic? So far (as far as I know) no-one’s measured the ASP of another ramus, so the answer remains, for now, ridiculously unknown to our planet.

Special bonus weirdness

Basal sauropodomorph wizard Adam Yates has posted an entry on his blog showing more sauropod vertebrae/ceratopsian frill convergence, as follow-up to our own recent post. Too weird.

Since my last post was rather heavier on the sushi than on the sauropod vertebrae, I offer this special bonus post. One of the frustrating things about the otherwise marvelous Sauroposeidon monograph (Wedel et al. 2000b) is that the figures are so small. Sadly this is also true of all the other publications that illustrate its remains, and so the published literature has no nice, detailed images.

No longer!  I’ve scored a rare paper copy of Matt’s undergraduate thesis (Wedel 1997) which contained basically all the material that eventually became that monograph, and which in addition has much larger versions of the figures.  So without further ado, I give you figure 5 of that paper:

Sauroposeidon cervical vertebrae. A, C5-C8; B, C6

Part A is similar to Wedel et al. (2000b:fig 6), and part B to Wedel et al. (2000b:fig. 7A), but this older version is rather nicer, and from a waaay better scan than is available for Wedel et al. (2000b).

And that’s all I have to say about that.

References.

This figure is stolen from Wedel et al. (2000:fig. 5). A shows the first 11 cervical vertebrae* of Sauroposeidon in articulation. B shows how the holotype specimen, OMNH 53062, must have disarticulated, and C shows it as it was found. Shaded vertebrae and bits of vertebrae were not found. The thickness of the cervical ribs is greatly exaggerated for clarity.

*We assume that Sauroposeidon had 13 cervicals like Brachiosaurus. It is not beyond the bounds of possibility that it had more, but it is unlikely that it had fewer. Sauroposeidon seems to be all about crazy neck elongation, and it doesn’t make sense to make some vertebrae longer while losing others.

Some facts:

  • In life, the long cervical ribs formed overlapping bundles, just like the long neck tendons of birds, and that is how the preserved cervical ribs are arrayed–in vertically stacked bundles.
  • Each cervical rib is about 4 cm in diameter where it attaches to its vertebra, and tapers to a point about 3 meters away. The last meter or so of each rib goes from being the diameter of a pencil to the diameter of a mechanical pencil lead. They just sort of peter out into nothingness.
  • The fact that even the pencil-lead-sized wisps of the cervical ribs are still in articulation suggests pretty strongly that the neck was buried with the muscles intact.
  • If the neck had simply been broken transversely (like a guillotine cut), the two most anterior vertebrae in the preserved block of four should have the cervical ribs of even more anterior vertebrae beneath them, and the cervical ribs from the two most posterior vertebrae would not stick out the back of the preserved block.
  • The facts that the cervical ribs from the missing anterior vertebrae are also missing, and that the cervical ribs from the preserved vertebrae trail behind the articulated block, suggest that the neck was pulled apart lengthwise, as shown in B.
  • None of the vertebrae have any teeth marks or any sign of mechanical damage, other than the missing neural spine from the third preserved vertebra. The front third of the first preserved vertebra was eroded away before the vertebrae were discovered in the field.
  • Assuming that Sauroposeidon was built like Brachiosaurus, it must have had a body mass somewhere between 40 and 60 tons. Even if it was built more like Mamenchisaurushellacious neck tacked on fairly dinky body–it was still probably a 20-ton critter.
  • After 14 years of subsequent erosion and fieldwork, no other sauropod bones have been discovered at the site.

Some questions:

  • How did the neck get separated from the body? The body was presumably too big to move, and the neck is too well preserved to have been moved very far.
  • What pulled the neck apart?
  • How did the neck come apart without disturbing those little pencil-lead cervical rib ends?

I don’t know the answers to those questions, by the way. And I’m open to suggestions.

Here’s my best guess. I think the body stayed put, and the neck floated away. Not far–a few hundred feet would be enough to put the body outside the outcrop area at the holotype site, but not so far that the neck would be all beat up. I think it floated rather than being dragged (by an Acrocanthosaurus, for example) because the vertebrae are all in such good shape and none of them have any tooth marks. I think it floated in calm water because the preservation is so good. I think the neck muscles rotted enough to let the force of the current rip part of the neck away from the base, just like you can pull a cooked chicken neck apart lengthwise without messing up the articulations among the vertebrae in the chunk that breaks free.

All of that will suffice to get the neck separated from the body. What really bugs me is the separation of the anterior part of the neck from the preserved block of vertebrae. It is tempting to think that the anterior part never came off, and that those vertebrae simply eroded away before they were found, like the front third of the most anterior preserved vert. But that can’t be; if those vertebrae were in articulation and just eroded away, we should still have their cervical ribs below the first two preserved verts.

Who knows, maybe the scenario I outlined above is good enough to explain both breaks. For some reason it is just easier to image most of the neck coming off the carcass than to imagine one part of the neck coming off the other part of the neck. But maybe the anteriormost vertebrae were ripped off and floated away first, and then the preserved block came free and floated off on its own later. (The head probably exploded, as these things were wont to do.)

It is worth noting that there are probably only a handful of people alive who have any first-hand experience with how multi-ton animal carcasses are dispersed, and zero people alive who have ever seen a dead sauropod rot. So, like too much in paleontology, what seems plausible or reasonable to me may not line up with objective reality.

BTW, this post fulfills a promise I made in a comment thread here. If we promise a post, we deliver. (We just don’t specify a due date.)

Comments, suggestions, hypotheses, rants, and crank fringe theories welcome.

References:

The image I put together explaining the new discovery. Modified from Staples et al. (2019: fig. 6).

Today sees the publication of a new paper, “Cutaneous branch of the obturator nerve extending to the medial ankle and foot: a report of two cadaveric cases,” by Brittany Staples, Edward Ennedy, Tae Kim, Steven Nguyen, Andrew Shore, Thomas Vu, Jonathan Labovitz, and yours truly. I’m excited for two reasons: first, the paper reports some genuinely new human gross anatomy, which happens surprisingly often but still isn’t an everyday occurrence, and second, the first six authors are my former students. This isn’t my discovery, it’s theirs. But I’m still going to yap about it.

When the obturator nerve won’t stay in its lane

Your skin is innervated by cutaneous nerves, which relay sensations of touch, pressure, vibration, temperature, and pain to your central nervous system, and carry autonomic (involuntary) fibers to your sweat and sebaceous glands and the arrector pili muscles that raise and lower your hairs (as when we get goosebumps). Every inch of your skin lies in the domain of one cutaneous nerve or another. Known boundaries between cutaneous branches of different nerves are approximate, both because they vary from person to person, and because the territories of the nerves themselves interdigitate and overlap at very fine scales. That said, aside from complex areas where the domains of multiple nerves intersect (like the groin), most body regions get their cutaneous innervation from just one nerve.

The obturator nerve arises from the spinal levels of the 2nd-4th lumbar vertebrae (L2-L4), exits the pelvis through the obturator canal behind the superior ramus of the pubis, and innervates the adductor muscles of the medial compartment of the thigh. The cutaneous branch of the obturator nerve typically innervates a variable but limited patch of skin on the inner thigh. Here’s a diagram from Gray’s Anatomy, 40th edition, showing the common cutaneous distribution of the obturator nerve (Standring et al. 2008 fig. 79.17, modified):

In rare cases, however, the obturator nerve doesn’t stay in the thigh. I was teaching in the gross anatomy lab in the fall of 2013 when one of our podiatry students, Brittany Staples, called me over to her table. We were skinning the thigh and leg that day, and in her assigned cadaver, Brittany had found a nerve from the medial thigh running all the way down to the inner side of the ankle and foot.

I didn’t immediately freak out, because everyone has a nerve from the thigh running down to the inner side of the ankle and foot: the saphenous branch of the femoral nerve, which comes out of the anterior thigh (also highlighted in the above image). But when we traced back Brittany’s nerve, it wasn’t coming from the femoral nerve. Instead, it was coming from the anterior division of the obturator nerve, right behind the adductor longus muscle (when people do the splits, this is the muscle that makes a visible ridge from the inner thigh to the groin). We carefully cleaned and photographed the nerve, and then we hit the books. Our first question: was this a known variation, or had Brittany discovered something new in the annals of human anatomy?

Standing on the shoulders of giants

Virtually all introductory anatomy textbooks show the obturator nerve only going to the thigh. But a little digging turned up Bouaziz et al. (2002), which in turn reproduced a figure from Rouvière and Delmas (1973), a French textbook, which showed the obturator nerve passing the knee and innervating part of the calf. That was at least an advance on what we knew starting out. We found a similar written description in Sunderland (1968).

Bouaziz et al. (2002: fig. 1)

Then we discovered Bardeen (1906), a magnificent and magisterial work 130 pages in length. Titled, “Development and variation of the nerves and the musculature of the inferior extremity and of the neighboring regions of the trunk in man”, the paper delivers on its impressive title. Bardeen (1906: 285 and 317) reported than in 22 out of 80 cadavers, the cutaneous branch of the obturator nerve (CBO) reached the knee; in 10 of those cases it could be traced at least to the middle third of the calf; and in one case it reached “nearly to ankle”. Bardeen also commented on the difficulty of tracing out the limits of this tiny nerve (p. 285):

“How constant the cutaneous branch of the obturator may be I have been unable satisfactorily to determine. Students dissecting frequently fail to find it. Owing to the fact that this may often be due to its small size the negative records cannot safely be used in making up statistics.”

All of us on the paper can back up Bardeen’s comments here: by the time they reach the skin, cutaneous nerves might be as big around as a pencil lead, or a strand of dental floss, or a human hair, but they won’t be much bigger. Sometimes they run just under the skin, sometimes down in the subcutaneous fat and fascia (with vanishingly small extensions spidering out to the underside of the skin), always variable in their courses and often devilishly hard to find, preserve, and trace.

If there is a prior report in the literature of a CBO passing the ankle, we haven’t found it, and neither have the numerous podiatric physicians who commented on the manuscript before we submitted, nor the reviewers and editors of the Journal of Foot & Ankle Surgery. I feel pretty safe saying that this is truly new (and if you know otherwise, please let me know in the comments!).

The second case, and the long silence

Every year since 2013, I’ve warned our medical and podiatric students to be on the lookout for anomalously long branches of the obturator nerve. The very next year, a group of summer anatomy students found a second example (they’re authors 2-6 on the paper). Since then, nada, in over 200 more bodies as of this summer. Either we got crazy lucky to find two examples in back to back years, or long CBOs are more common than we think, just really hard to find and identify. More on that in a minute.

A quick aside: we didn’t deliberately hold up the paper while we were looking for more examples, we’ve all just been busy. Brittany and the other student authors were occupied with passing med school and their board exams, surviving clinical rotations, and applying to residency programs. I’m happy to say “were occupied” with all those things because they’re all graduated now, and in residency training. Anyway, that’s why the paper had a 5-year gestation: med school doesn’t leave a lot of time for research and writing. Kudos to Brittany for giving all of us regular kicks to keep things moving along. In every sense, the paper would not exist without her skill and dedication.

So what’s going on here?

There are two sides to this: what happened to produce the variants we found in 2013 and 2014, and why variants like that escaped detection for so long, and I’ll tackle them in that order.

We found both of the long CBOs in the territory normally occupied by the saphenous branch of the femoral nerve. The saphenous nerve is so named because it runs along the great saphenous vein, the major superficial vein of the medial leg and thigh. Sometimes the saphenous nerve has only a single main trunk, but more commonly it splits into two parallel branches, one on either side of the saphenous vein, as illustrated here by Wilmot and Evans (2013: fig. 3):

In both of our cases, the saphenous branch of the femoral nerve was present, but it only had one branch, in front of the big vein, and the long CBO ran behind the vein, in the place usually occupied by the posterior branch of the saphenous nerve. In effect, the posterior part of the saphenous branch of the femoral nerve had been replaced by a sort of saphenous branch of the obturator nerve. This has interesting implications.

Suppose you were a surgeon, harvesting the distal portion of the saphenous vein for a coronary artery bypass graft, and you saw two nerves accompanying the vein, one in front and one behind. You would probably assume that both branches arose from the femoral nerve, because that is what happens most commonly. But if the posterior branch actually came from the obturator nerve, you’d have no way of knowing that, without tracing the nerve back to its origin in the inner thigh. The watchwords in surgery these days are “minimally invasive” and “patient outcomes” — smaller openings in the body mean less pain, fewer complications, faster recoveries, and happier patients. So surgeons aren’t going to flay patients open from ankle to groin just to chase down a nerve that might be coming from the normal place after all.

If you only get to look inside the box, these two things look the same.

We suspect that long CBOs may be fairly common, just hard to recognize, because who is going to find them? Medical students dissecting human cadavers have the opportunity to trace long cutaneous nerves back to their origins, but since it’s the students’ first time cutting, they usually haven’t yet developed the experience to recognize weird versions of tiny nerves, nor the skill to preserve them. Surgeons have the experience and the skill, but not the opportunity, because they can’t go around filleting their patients to see where all the nerves come from. So long CBOs probably fall into a perceptual blind spot, in which almost no-one who cuts on human bodies has both the opportunity to find them, and the skill to preserve them — my former students excepted (he said with no small helping of pride).

That’s pretty darned interesting, and it makes me wonder what other perceptual blind spots are out there, in both anatomy and paleontology. I know of at least one: the true nature and extent of the fluid-filled interstitial tissues that pervade our bodies (and those of all other vertebrates at least) were not fully appreciated until just last year, because the first step in the production of most histological slides is to dehydrate the tissues, which collapses the fluid-filled spaces and makes the interstitium look like regular connective tissue (Benias et al. 2018). That is a spooky kind of observer effect, and it makes me wonder what else we’re missing because of the ways we choose — or are constrained — to look.

What next?

What’s the fallout from this study? For me, two things. First — obviously — we’re going to keep looking for more examples of long CBOs, and for other similar cases in which one nerve may have been replaced by its neighbor. This is more than trivia. Knowing which nerves to expect and where to find them is important, not only for surgeons but also for anaesthetists and pain management physicians doing nerve blocks. The decks may be stacked against med students for some of these discoveries, but clearly “difficult” does not mean “impossible” or I’d have nothing to write about. Lightning has already struck twice, so I’ll keep flying this particular kite.

Second, this case, a few other odd things we’ve found in the lab over the years, and other recently-reported discoveries in human anatomy have caused me to wonder: could we formulate predictive maxims to help guide future discoveries in human anatomy, or in anatomy full stop? I think so, and provided my abstract is accepted, I’ll be presenting on that topic at SVPCA in a couple of months. More on that in due time.

Finally — and this cannot be overstated — without the keen eyes, skilled hands, sharp minds, and hard work of the student authors, there would be no discovery and no paper. So congratulations to Brittany, Edward, Tae, Steven, Andrew, and Thomas. Or as I’m happy to address them now, Drs. Staples, Ennedy, Kim, Nguyen, Shore, and Vu. Y’all done good. Keep it up.

References

  • Bardeen, C.R. 1906. Development and variation of the nerves and the musculature of the inferior extremity and of the neighboring regions of the trunk in man. Developmental Dynamics 6(1):259-390.
  • Benias, P.C., Wells, R.G., Sackey-Aboagye, B., Klavan, H., Reidy, J., Buonocore, D., Miranda, M., Kornacki, S., Wayne, M., Carr-Locke, D.L. and Theise, N.D. 2018. Structure and distribution of an unrecognized interstitium in human tissues. Scientific Reports, 8:4947.
  • Bouaziz, H., Vial, F., Jochum, D., Macalou, D., Heck, M., Meuret, P., Braun, M., and Laxenaire, M.C. 2002. An evaluation of the cutaneous distribution after obturator nerve block. Anesthesia & Analgesia 94(2):445-449.
  • Rouvière, H., and Delmas, A. 1973. Anatomie humaine, descriptive, topographique et fonctionnelle: tome 3—membres-système nerveux central, ed 11, Masson, Paris.
  • Standring, S. (ed.) 2008. Gray’s Anatomy: The Anatomical Basis of Clinical Practice, 41st ed, Elsevier Health Sciences, London.
  • Staples, B., Ennedy, E., Kim, T., Nguyen, S., Shore, A., Vu, T., Labovitz, J., and Wedel, M. 2019. Cutaneous branch of the obturator nerve extending to the medial ankle and foot: a report of two cadaveric cases. Journal of Foot & Ankle Surgery, advance online publication.
  • Sunderland, S. 1968. Nerves and Nerve Injuries. Churchill Livingstone, Edinburgh.
  • Wilmot, V.V., and Evans, D.J.R. 2013. Categorizing the distribution of the saphenous nerve in relation to the great saphenous vein. Clinical Anatomy 26(4):531-536.

I was delighted today to see a tweet from dinodadreviews:

(Here is it, archived, in case it goes away for any reason):

Another kid’s book featuring @MikeTaylor’s baby, Xenoposeidon! Seen in this “#Alphasaurs” book as its old brachiosaurus interpretation, I love the “X-ray” flap showing the approximate location of its one known bone! 🦕

This is a nice, elegant bit of artwork, based of course on the old brachiosaurid interpretation of Xenoposeidon — which has been superseded by the new rebbachisaurid interpretation, but the author and designer weren’t to know that.

My only reservation, really, is that the pronunciation isn’t quite right. There’s no real excuse for that as I gave it right in the paper: it should be “ZEE-no-puh-SYE-d’n”. Oh well.

The inspiration for the book illustration will have been this image:

which we used in Post 4 of the original Xenoposeidon week, and also in my old, pre-SV-POW! web-page about it. That in turn came from this one:

which I made as a joke and described as “the first scientifically rigorous skeletal reconstruction of Xenoposeidon. As the Day-four post says, “I thought it would be funny to do this for an animal known only from a single bone, showing the bone floating in the middle of a big black silhouette. Har har.” It’s funny, now, twelve years later, to the see the descendent of that image in a kids’ book.

Finally, these Xenoposeidon “reconstructions” were based on the solid work that Matt had done on a Brachiosaurus reconstruction (actually Giraffatitan, but back then we thought the latter was a species of the former) to be used in the papers about Sauroposeidon:

Matt wrote a short paper for Prehistoric Times about his work on this reconstuction. It’s only one page: go and read it.

dinodadreviews’ tweet was the first I’ve heard of the Alphasaurs book, but following the #Alphasaurs hashtag took me to a tweet by the book’s designer, which in turn took me to the book’s Amazon page. And there, I was surprised but pleased to see the Xenoposeidon gets the star billing in the Booklist review:

“X marks the spot” for Xenoposeidon. In this alpha-bestiary, the X denoting the only bone found for this long-necked dinosaur—from which its entire structure has been extrapolated—is cut into a flap that, when lifted, reveals Xenoposeidon’s very, very long tail. This dinosaur, like the other 25 who walk, swim, fly, and prowl through these foldout pages, is made up of hundreds of the first letter of its name. Check out the red capital As that mark Allosaurus’ fangs, or the vicious-looking Vs of the Velociraptor’s claws, or the way the Ws of Wuerhosaurus form spikes on its dangerous tail. Each of the dinosaurs showcases a different typeface, too (all the typefaces are identified at the book’s end). Meanwhile, fast facts about dinosaurs fill the margins. There is little doubt the strange art will reel them in—and probably keep them reading. A wholly unique mix of typography and dinosaur science. Grades 1-3. — Connie Fletcher

I’m not quite sure how Connie Flecher concluded that the lift-the-flap reveals the tail, but I’m prepared to give her a pass since the had the good judgement to lead with Xeno.

 

 

If you followed along with the last post in this series, you now have some bird vertebrae to play with. Here are some things to do with them.

1. Learn the parts of the vertebrae, and compare them with those of other animals

Why are we so excited about bird vertebrae around here? Mostly because birds are reasonably long-necked living dinosaurs, and although their vertebrae differ from those of sauropods in relative proportions, all of the same bits are present in roughly the same places. If you know the parts of a bird vertebra and what each one does, you have a solid foundation for inferring the functions of sauropod vertebrae. Here’s a diagram I made for my SVP poster with Kent Sanders way back in 1999. I used an ostrich vertebra here but you should be able to find the same features in a cervical vertebra of just about any bird.

These are both middle cervical vertebrae in right lateral view. A middle cervical vertebra of a big ostrich will be between 3 and 4 inches long (7.5-10 cm), and one from a big brachiosaur like Giraffatitan will be about ten times longer.

I should do a whole post on neck muscles, but for now see this post and this paper.

2. Put the vertebrae in order, and rearticulate them

It is often useful to know where you are in the neck, and the only way to figure that out is to determine the serial position of the vertebrae. Here’s an articulated cervical series of a turkey in left lateral view, from Harvey et al. (1968: pl. 65):

Harvey’s “dorsal spine” is the neural spine or spinous process, and his “ventral spine” is the carotid process. The “alar process” is a sort of bridge of bone connecting the pre- and postzygapophyses; you can see a complete version in C3 in the photo below, and a partial version in C4.

Speaking of that photo, here’s my best attempt at rearticulating the vertebrae from the smoked turkey neck I showed in the previous post, with all of the vertebrae in left dorsolateral view.

These things don’t come with labels and it can take a bit of trial and error to get them all correctly in line. C2 is easy, with its odd articular surface for the atlas and narrow centrum with a ventral keel. Past that, C3 and C4 are usually pretty blocky, the mid-cervicals are long and lean, and then the posterior cervicals really bulk out. Because this neck section had been cut before I got it, some of the vertebrae look a little weird. Somehow I’m missing the front half of C6. The back half of C14 is also gone, presumably still stuck to the bird it went with, and C7 and C12 are both sectioned (this will come in handy later). I’m not 100% certain that I have C9 and C10 in the right order. One handy rule: although the length and neural spine height change in different ways along the column, the vertebrae almost always get wider monotonically from front to back.

And here’s the duck cervical series, in right lateral view. You can see that although the specific form of each vertebra is different from the equivalent vert in a turkey, the same general rules apply regarding change along the column.

Pro tip: I said above that these things don’t come with labels, but you can fix that. Once you have the vertebrae in a satisfactory order, paint a little dot of white-out or gesso on each one, and use a fine-point Sharpie or art pen to write the serial position (bone is porous and the white foundation will keep the ink from possibly making a mess). You may also want to put the vertebrae on a string or a wire to keep them in the correct order, but even so, it’s useful to have the serial position written on each vertebra in case you need to unstring them later.

3. Look at the air spaces

One nice thing about birds is that all of the species that are readily commercially available have pneumatic traces on and in their vertebrae, which are broadly comparable to the pneumatic vertebrae of sauropods.

The dorsal vertebrae of birds are even more obviously similar to those of sauropods than are the cervicals. These dorsal vertebrae of a duck (in left lateral view) show a nice variety of pneumatic features: lateral fossae on the centrum (what in sauropods used to be called “pleurocoels”), both with and without foramina, and complexes of fossae and foramina on the neural arches. Several of the vertebrae have small foramina on the centra that I assume are neurovascular. One of the challenges in working with the skeletal material of small birds is that it becomes very difficult to distinguish small pneumatic foramina and spaces from vascular traces. Although these duck vertebrae have small foramina inside some of the lateral fossae, the centra are mostly filled with trabecular, marrow-filled bone. In this, they are pretty similar to the dorsal vertebrae of Haplocanthosaurus, which have fossae on the neural arches and the upper parts of the centra, but for which the ventral half of each centrum is a brick of non-pneumatic bone. For more on distinguishing pneumatic and vascular traces in vertebrae, see O’Connor (2006) and Wedel (2007).

This turkey cervical, in left posterolateral view, shows some pneumatic features to nice advantage. The lateral pneumatic foramina in bird cervicals are often tucked up inside the cervical rib loops where they can be hard to see and even harder to photograph, but this one is out in the open. Also, the cervicals of this particular turkey have a lot of foramina inside the neural canal. In life these foramina are associated with the supramedullary diverticula, a set of air-filled tubes that occupy part of the neural canal in many birds — see Atterholt and Wedel (2018) for more on this unusual anatomical system. The development of foramina inside the neural canal seems to be pretty variable among individuals. In ostriches I’ve seen individuals in which almost every cervical has foramina inside the canal, and many others with no foramina. For turkeys it’s even more lopsided in my experience; this is the first turkey in which I’ve found really clear pneumatic foramina inside the neural canals. This illustrates one of the most important aspects of pneumaticity: pneumatic foramina and cavities in bones show that air-filled diverticula were present, but the absence of those holes and spaces does not mean that diverticula were absent. Mike and I coined the term “cryptic diverticula” for those that leave no diagnostic traces on the skeleton — for more on that, see the discussion section in Wedel and Taylor (2013b).

Finally, it’s worth taking a look at the air spaces inside the vertebrae. Here’s a view into C12 of the turkey cervical series shown above. The saw cut that sectioned this neck happened to go through the front end of this vertebra, and with a little clean-up the honeycomb of internal spaces is beautifully displayed. If you are working with an intact vertebra, the easiest way to see this for yourself is to get some sandpaper and sand off the front end of the vertebra. It only takes a few minutes and you’ll be less likely to damage the vertebrae or your fingers than if you cut the vertebra with a saw. Similar complexes of small pneumatic cavities are present in the vertebrae of some derived diplodocoids, like Barosaurus (see the lateral view in the middle of this figure), and in most titanosauriforms (for example).

I have one more thing for you to look for in your bird vertebrae, and that will be the subject of the next installment in this series. Stay tuned!

References

What it says on the tin. This is a specimen from the UCMP comparative collection.

I was just going to post this photo with zero commentary, but I can’t help myself. Note that on the two vertebrae in the middle, the crista transverso-obliqua (what in non-avian dinos would be the spinopostzygapophyseal lamina or SPOL) rises higher than either the neural spine apex or the epipophyses. That’s crazy. And it demonstrates something we also see in sauropods, which is that laminae are not merely the plates of bone left behind after pneumatization has scooped all of the unnecessary material out of a normal vertebra–sometimes they are additive structures, too.

If all of that sounded like gibberish, I can sympathize. I spent my first few months as a sauropodologist just learning the lingo (another thing I should blog about sometimes). Here’s a labeled version:

As long as I’m yapping, note the light shining through the honeycombed internal structure of these highly pneumatic vertebrae. For more on the ridiculous pneumaticity of pelican bones, see this post and this one. For more on the homology of bird and sauropod vertebrae, see Wedel and Sanders (2002), and for more on laminae as additive versus reductive structures, see the discussion on pages 210-212 of Wedel (2007).

References

 

In a comment on the last post, Mike wrote, “perhaps the pneumaticity was intially a size-related feature that merely failed to get unevolved when rebbachisaurs became smaller”.

Caudal pneumaticity in saltasaurines. Cerda et al. (2012: fig. 1).

Or maybe pneumaticity got even more extreme as rebbachisaurids got smaller, which apparently happened with saltasaurines  (see Cerda et al. 2012 and this post).

I think there is probably no scale at which pneumaticity isn’t useful. Like, we see a saltasaurine the size of a big horse and think, “Why does it need to be so pneumatic?”, as if it isn’t still one or two orders of magnitude more massive than an ostrich or an eagle, both of which are hyperpneumatic even though only one of them flies. Even parakeets and hummingbirds have postcranial pneumaticity.

Micro CT of a female Anna’s hummingbird. The black tube in the middle of the neck is the supramedullary airway. Little black dots in the tiny cervical centra are air spaces.

We’re coming around to the idea that the proper way to state the dinosaur size question is, “Why are mammals so lousy at being big on land?” Similarly, the proper way to state the pneumaticity question is probably not “Why is small sauropod X so pneumatic?”, but rather “Why aren’t some of the bigger sauropods even more pneumatic?”

Another thought: we tend to think of saltsaurines as being crazy pneumatic because they pneumatized their limb girdles and caudal chevrons (see Zurriaguz et al. 2017). Those pneumatic foramina are pretty subtle – maybe their apparent absence in other sauropod clades is just because we haven’t looked hard enough. Lots of things have turned out to be pneumatic that weren’t at first glance – see Yates et al. (2012) on basal sauropodomorphs and Wedel and Taylor (2013b) on sauropod tails, for example.

Back of the skull of a bighorn sheep, showing the air spaces inside one of the broken horncores.

Or, even more excitingly, if the absence is genuine, maybe that tells us something about sauropod biomechanics after all. Maybe if you’re an apatosaurine or a giant brachiosaurid, you actually can’t afford to pneumatize your coracoid, for example. One of my blind spots is a naive faith that any element can be pneumatized without penalty, which I believe mostly on the strength of the pneumatic horncores of bison and bighorn sheep. But AFAIK sauropod girdle elements don’t have big marrow cavities for pneumaticity to expand into. Pneumatization of sauropod limb girdles might have come at a real biomechanical cost, and therefore might have only been available to fairly small animals. (And yeah, Sander et al. 2014 found a pneumatic cavity in an Alamosaurus pubis, but it’s not a very big cavity.)

As I flagged in the title, this is noodling, not a finding, certainly not certainty. Just an airhead thinking about air. The comment thread is open, come join me.

References