This is corn on the cob:

Corn on the cob, in cross section. Stolen from http://www.istockphoto.com/file_thumbview_approve/214165/2/istockphoto_214165-co rn-cob-cross-section.jpg

This is a shish kebab:

Shish kebab. Stolen from http://www.mediterraneancafe-flatiron.com/images/shish.jpg

Most tetrapods are like shish kebabs: a whole lot of meat stuck on a proportionally tiny skeleton.  If you don’t believe me, you can look at the human and cow neck torso cross-sections in Matt’s last post, or check out this ostrich-neck cross-section from his 2003 Paleobiology paper:

Ostrich neck in cross section, CT scan. From Wedel (2003a: fig. 2)

Remember that this is a freakin’ ostrich — of all extant animals, one of the ones with a most extreme long, skinny neck.  And yet, if sauropods were muscled like ostriches, then their necks would have looked like this in cross section:

Putative shish kebab-style sauropod neck in cross section. Ostrich soft-tissue from Wedel (2003a: fig. 2), Diplodocus vertebra cross-section from Paul (1997: fig. 4) scaled to match size of ostrich vertebra

And soft-tissue reconstructions would have to look like this:

Diplodocus with its neck as fat as an ostrich's. Modified from Paul (1998: fig. 1F)

Which, happily, no-one is suggesting.  Instead, published reconstructions of sauropod neck soft-tissue are startlingly emaciated.  As exhibit A, I call this pair of Greg Paul cross-sections:

Diplodocus and Brachiosaurus neck cross-sections, showing very light musculature. From Paul (1997: fig. 4)

(Yes, the Diplodocus on the left is the one I used in the photoshopped ostrich cross-section above.  It’s instructive to compare Paul’s original with the What If It Was Like A Big Ostrich version.)

Paul’s reconstructions seem to be widely considered too lightly muscled.  But even the very careful and rigorous more recent reconstructions of Daniela Schwarz and her colleague show a neck much, much thinner than that of the ostrich:

Diplodocus neck cross-sections. From Schwarz et al. (2007: fig. 7a)

Although Schwarz has put a lot more soft tissue onto the neck vertebrae than Paul did, it is still a tiny proportion of what we see in extant animals — even the ostrich, remember, which has a super-thin neck compared with pretty much anything else alive today.  If sauropod necks were muscled as heavily as those of, say, cows, then the soft tissue would pretty much reach down to the ground.  But they weren’t: they were more like corn on the cob, with a broad core of skeleton and relatively little in the way of delicious edibles festooned about it.

So why is this?  Why does everyone agree that sauropod necks were much less heavily muscled than those of any extant animal?

It’s a simple matter of scaling.  A really big ostrich might have a neck 1 m long.  (Actually, ostriches don’t get that big, but let’s pretend they do because it makes the maths easier).  If the x meter-long neck of a sauropod was just a scaled-up ostrich neck, then it would be x times longer, x times taller and x times wider, for a total of x^3 times as voluminous and therefore x^3 times as heavy.  But the cross-sectional area of the tension members that support it is only x times taller and x times wider, for a total of x^2 times the strength.  In total, then, the neck’s mass/strength is x^3/x^2 = x times as great as in the ostrich.  (The sauropod neck’s mass also acts further out from the fulcrum by an additional factor of x, but that is cancelled by the fact that the tension in the neck also acts x times higher above the fulcrum.)

It seems intuitively obvious (which is is code for “I have no way to prove”) that you can’t reasonably expect the neck muscles of a giant ostrich to work ten times as hard as they do in their lesser cousins, which is what you’d need to do for the 10 m neck of, say, Sauroposeidon.  So simple isometric scaling won’t get the job done, and you need to restructure the neck.

But how?  Surely just reducing all the muscle around the vertebrae can’t help?  No indeed — but that is not really what sauropods were doing.  If you look at the typical sauropod-neck life restoration, you’ll see that the proportional thickness of the neck is actually not too dissimilar to that of an ostrich — rather thicker, in fact.  If you scaled an ostrich neck up to sauropod size and compared it with a real sauropod neck, you would find not that the soft tissue was too fat, but that the vertebrae were too thin.

And so we come to it at last: rather than thinking of sauropods as having reduced the amount of soft-tissue hanging on the cervical vertebrae, we do better to think of them as having kept a roughly similar soft-tissue profile to that of an an ostrich, but enlarging the vertebrae within the soft-tissue envelope.  Of course if you just blindly made the vertebrae taller and wider, they would become heavier in proportion, which would defeat the whole purpose of the exercise — but as everyone who reads this blog surely knows by now, sauropod cervicals were extensively lightened by pneumaticity.  By bringing air into the center of the neck, they were effectively able to displace bone, muscle and ligament away from the centre, so that they acted with greater mechanical advantage: higher epaxial tension members, lower hypaxial compression members, and more laterally positioned paraxials.

It’s a rather brilliant system — using the same volume of bone to achieve greater strength by displacing it outwards and filling the center with air (and, in doing so, also displacing soft tissue outwards).  And it will be hauntingly familiar to anyone who loves birds, because it is of course exactly what birds (and pterosaurus) have done in their long bones: the hollow humeri of flying vertebrates famously allow them to attain greater strength — specifically, resistance to bending — for the same volume and mass of bone.  It’s a neat trick when done with long bones, but it takes a truly awesome taxon to do it with the neck.

So maybe sauropods were not corn on the cob after all.  Maybe they were Hostess Twinkies.

Hostess Twinkie. Not truly pneumatic, as the internal cavity is filled with adipose tissue rather than air, but do you have any idea how difficult it is to find good images of hollow junk food? Stolen from http://dixiedining.files.wordpress.com/2008/07/twinkie_070918_ms1.jpg

And now for something completely different

Now that I’ve finished my Ph.D at the University of Portsmouth, what am I going to do with the rest of my scientific life?  I’ve always said that I have no intention of going into palaeo full time: my knowledge is far too narrow for that, so that even if paid jobs were not in insanely short supply, I wouldn’t stand much chance of getting one.  And in any case, I’d hate to get into the all-too-common situation of being up against a friend for a position we both wanted. Throw in the fact that I really enjoy my computer-programming day-job and it seems pretty clear that what I need is an unpaid affiliation that lets me get on with lovely research.

Well: I am absolutely delighted to announce that, as of last month, I am an Honorary Research Associate in the Department of Earth Sciences at UCL.  It’s not just that UCL is such a well-respected institution — see that Wikipedia article for some details — more importantly, it’s where Paul Upchurch hangs out, as Senior Lecturer in Palaeobiology.  Sauropod fans will be familiar with Paul’s characteristically detailed and careful work, from his pioneering work on sauropod phylogeny (Upchurch 1995, 1998), through his and John Martin’s indispensible Cetiosaurus makeovers (Upchurch and Martin 2002, 2003) to the state-of-the art review that he lead-authored for Dinosauria II (Upchurch et al. 2004) and the Tokyo Apatosaurus monograph (Upchurch et al. 2005).  What many of you won’t know is what an excellent collaborator he is — quick, conscientious, insightful and diplomatic.  We’ve already collaborated on a few short papers (Upchurch et al. 2009 and a couple of Phylocode companion-volume chapters that are in press), and I hope there will be more in the future.

References

• Paul, Gregory S.  1997.  Dinosaur models: the good, the bad, and using them to estimate the mass of dinosaurs.  pp. 129-154 in: D. L. Wolberg, E. Stump, and G. D. Rosenberg (eds.), DinoFest International: Proceedings of a Symposium Sponsored by Arizona State University.  Academy of Natural Sciences, Philadelphia.
• Paul, Gregory S.  1998.  Terramegathermy and Cope’s Rule in the land of titans.  Modern Geology 23: 179-217.
• Schwarz, Daniela, Eberhard Frey and Christian A. Meyer.  2007. Pneumaticity and soft-tissue reconstructions in the neck of diplodocid and dicraeosaurid sauropods.  Acta Palaeontologica Polonica 52(1): 167-188.
• Upchurch, Paul.  1995.  The evolutionary history of sauropod dinosaurs.  Philosophical Transactions of the Royal Society of London Series B, 349: 365-390.
• Upchurch, Paul.  1998.  The phylogenetic relationships of sauropod dinosaurs.  Zoological Journal of the Linnean Society, 124: 43-103.
• Upchurch, Paul and John Martin.  2002.  The Rutland Cetiosaurus: the anatomy and relationships of a Middle Jurassic British sauropod dinosaur.  Palaeontology, 45(6): 1049-1074.
• Upchurch, Paul and John Martin.  2003.  The anatomy and taxonomy of Cetiosaurus (Saurischia, Suaropoda) from the Middle Jurassic of England.  Journal of Vertebrate Paleontology 23(1): 208-231.
• Upchurch, Paul, Paul M. Barrett and Peter Dodson.  2004.  Sauropoda. pp. 259-322 in D. B. Weishampel, P. Dodson and H. Osmólska (eds.), The Dinosauria, 2nd edition.  University of California Press, Berkeley and Los Angeles.  861 pp.
• Upchurch, Paul, Yukimitsu Tomida, and Paul M. Barrett.  2005.  A new specimen of Apatosaurus ajax (Sauropoda: Diplodocidae) from the Morrison Formation (Upper Jurassic) of Wyoming, USA.  National Science Museum Monographs No. 26.  Tokyo.  ISSN 1342-9574.
• Upchurch, Paul, John Martin, and Michael P. Taylor.  2009.  Case 3472: Cetiosaurus Owen, 1841 (Dinosauria, Sauropoda): proposed conservation of usage by designation of Cetiosaurus oxoniensis Phillips, 1871 as the type species.  Bulletin of Zoological Nomenclature 66(1): 51-55.
• Wedel, Mathew J.  2003.  Vertebral pneumaticity, air sacs, and the physiology of sauropod dinosaurs.  Paleobiology 29(2): 243-255.