In the first installment in this series (link), we looked at a couple of weird sauropod vertebrae with neurocentral joints that were situated either entirely dorsal or ventral to the neural canals. This post has more examples of what I am calling “offset” neurocentral synchondroses.

I decided it made more sense to refer to the synchondrosis as being offset, instead of referring to the neural canal as offset. Because the neural canal in all of these vertebrae is right where it pretty much always is, just dorsal to the articular surfaces of the centrum. In an adult, fused vertebra, there’d be no sign that anything unusual had ever happened. So I think it makes more sense to talk about the neurocentral joint having migrated dorsally or ventrally relative to the canal, rather than vice versa. If you know differently, or if these weirdos have been addressed before elsewhere and I’ve just missed it, please let me know in the comments!

Here’s a plate from Marsh (1896) showing caudal vertebrae of Camarasaurus (“Morosaurus” in O.C. Marsh parlance), which echo the Alamosaurus caudal from the first post in having the neurocentral joint almost entirely ventral to the neural canal. The neural arch here doesn’t just arch over the canal dorsally, it also cuts under it ventrally, at least in part.

This plate is also nice because it shows the relationships among the arch, centrum, and caudal ribs before they fuse. Here’s the caption, from Marsh (1896):

Here’s the preceding plate, Plate 33, with illustrations of an unfused Camarasaurus sacrum.

And its caption:

This plate not only shows how the sacral ribs fuse to the arch and spine medially, and to each other laterally (forming the sacrocostal yoke), it also shows a last sacral that is very similar to the aforementioned caudals in the position of the neurocentral joint. But interestingly that neurocentral joint offset only seems to be present in the last caudal – the lower figure shows widely-separated neurocentral joint surfaces in the more anterior centra, indicating that the neural arches (not figured in this dorsal view) did not wrap around the neural canal to approach the midline. (I think we’re looking at S2 through S5 here, and missing a dorso-sacral.)

So now I’m freaked out, wondering if this neural arch wrap-around in the caudals is common to most sauropods and I just haven’t looked at enough juvenile caudals to have spotted it before. As always, feel free to ablate my ignorance in the comments, particularly if you know of more published examples. I’m a collector.

The neural canal of that last sacral also has a very interesting cross-sectional shape, like a numeral 8. I have some thoughts on that, but they’ll keep for a future post in this series.


When I was nine, a copy of Don Glut’s The New Dinosaur Dictionary turned up in my local Waldenbooks. It wasn’t my first dinosaur book, by far – I’d been a dinosaurophile since the age of three. But The New Dinosaur Dictionary was different.

Up to that point, I had subsisted on a heavy diet of kids’ dino books and the occasional article in National Geographic and Ranger Rick. The kids’ books were aimed at kids and the magazine articles were pitched at an engagingly popular level. I didn’t understand every word, but they were clearly written for curious layfolk, not specialists.

A typical spread from The New Dinosaur Dictionary (Glut, 1982). The armored sauropod blew my young mind.

The New Dinosaur Dictionary was something else entirely. It had photos of actual dinosaur bones and illustrations of skeletons with cryptic captions like, “Skeleton of Daspletosaurus torosus. (After Russell)”. Okay, clearly this Russell cove was out there drawing dinosaur skeletons and this book had reproduced some of them. But nobody I knew talked like that, and the books I had access to up to that point held no comparable language.

The New Dinosaur Dictionary (Glut, 1982: p. 271)

Then there was stuff like this: “The so-called Von Hughenden sauropod restored as a brachiosaurid by Mark Hallett”. A chain of fascinating and pleasurable ideas detonated in my brain. “The so-called” – say what now? Nobody even knew what to call this thing? Somehow I had inadvertently sailed right to the edge of human knowledge of dinosaurs, and was peering out into taxa incognita. “Restored as a brachiosaurid” – so this was just one of several possible ways that the animal might have looked. Even the scientists weren’t sure. This was a far cry from the bland assurances and blithely patronizing tones of all my previous dinosaur books.

“By Mark Hallett.” I didn’t know who this Hallett guy was, but his art was all over the book, along with William Stout and some guy named Robert T. Bakker and a host of others who were exploding my conception of what paleo art could even be. Anyway, this Mark Hallett was someone to watch, not only because he got mentioned by name a lot, but because his art had a crisp quality that teetered on some hypercanny ridge between photorealism and scribbling. His sketches looked like they might just walk off the page.

In case that line about scribbling sounds dismissive: I have always preferred sketches by my favorite artists to their finished products. The polished works are frequently inhumanly good. They seem to have descended in a state of completed perfection from some divine realm, unattainable by mere mortals. Whereas sketches give us a look under the hood, and show how a good artist can conjure light, shadow, form, weight, and texture from a few pencil strokes. Put it this way: I am anatomist by temperament first, and by training and occupation second. Of course I want to see how things are put together.

The New Dinosaur Dictionary (Glut, 1982: p. 75)

Anyway, The New Dinosaur Dictionary was something completely new in my experience. It wasn’t aimed at kids and written as if by kids, like lots of kids’ books. It wasn’t even written by adults talking down (deliberately or inadvertently) to kids, or trying to reach a wide audience that might include kids. It was written by an adult, aiming at other adults. And it was admitting in plain language that we didn’t know everything yet, that there were lots of animals trembling on the outer threshold of scientific knowledge. I didn’t understand half of it – I was down in an ontogenetic trench, looking up as these packets of information exploded like fireworks over my head.

In Seeing In the Dark, the best book about why you should go out stargazing for yourself, Timothy Ferris writes about growing up on Florida’s Space Coast in the early 1960s, and watching the first generation of artificial satellites pass overhead:

I felt like an ancient lungfish contemplating the land from the sea. We could get up there.

That’s precisely the effect that The New Dinosaur Dictionary had on me: I could get up there. Maybe not immediately. But there were steps, bodies of knowledge that could be mastered piecemeal, and most of all, mysteries to be resolved. The book itself was like a sketch, showing how from isolated and broken bones and incomplete skeletons, scientists and artists reconstructed the world of the past, one hypothesis at a time. Now I take it for granted, because I’ve been behind the curtain for a couple of decades. But to my 9-year-old self, it was revolutionary.

This has all come roaring back because of something that came in the mail this week. Or rather, something that had been waiting in the mailroom for a while, that I finally picked up this week: a package from Mark Hallett, enclosing a copy of his 2018 dinosaur calendar. And also this:


An original sketch, which he gave to me as a Christmas present. The published version appears on one of the final pages of our book, where we discuss the boundaries between the known – the emerging synthesis of sauropod biology that we hoped to bring to a broader audience by writing the book in the first place – and the unknown – the enduring mysteries that Mark and I think will drive research in sauropod paleobiology for the next few decades. Presented without a caption or commentary, the sketch embodies sauropods as we see them: emerging from uncertainty and ignorance one hard-won line at a time, with ever-increasing solidity.

Thank you, Mark, sincerely. That sketch, what it evokes, both for me now and for my inner 9-year-old – you couldn’t have chosen a better gift. And I couldn’t be happier. Except perhaps to someday learn that our book exploded in the mind of a curious kid the way that The New Dinosaur Dictionary did for me 34 years ago, a time that now seems as distant and romantic as the primeval forests of the Mesozoic.

Computer programmer, essayist and venture capitalist Paul Graham writes:

In most fields, prototypes have traditionally been made out of different materials. Typefaces to be cut in metal were initially designed with a brush on paper. Statues to be cast in bronze were modelled in wax. Patterns to be embroidered on tapestries were drawn on paper with ink wash. Buildings to be constructed from stone were tested on a smaller scale in wood.

What made oil paint so exciting, when it first became popular in the fifteenth century, was that you could actually make the finished work from the prototype. You could make a preliminary drawing if you wanted to, but you weren’t held to it; you could work out all the details, and even make major changes, as you finished the painting.

You can do this in software too. A prototype doesn’t have to be just a model; you can refine it into the finished product. I think you should always do this when you can. It lets you take advantage of new insights you have along the way. But perhaps even more important, it’s good for morale.

– Paul Graham, “Design and Research

Mike and I have long been drawn by the idea that blog posts, like conference talks and posters, could be first drafts of research papers. In practice, we haven’t generated many successful examples. We basically wrote our 2013 neural spine bifurcation paper as a series of blog posts in 2012. And Mike’s 2014 neck cartilage paper grew out of this 2013 blog post, although since he accidentally ended up writing 11 pages I suppose the blog post was more of a seed than a draft.

I should also note that we are far from the first people to do the blog-posts-into-papers routine. The first example I know of in paleo was Darren’s Tet Zoo v1 post on azhdarchid paleobiology, which formed part of the skeleton of Witton and Naish (2008).

Nevertheless, the prospect of blogging as a way to generate research papers remains compelling.

And as long as I’m on about blogging and papers: sometimes people ask if blogging doesn’t get in the way of writing papers. I can’t speak for anyone else, but for me it goes in the opposite direction: I blog most when I am most engaged and most productive, and drops in blogging generally coincide with drops in research productivity. I think that’s because when I’m rolling on a research project, I am constantly finding or noticing little bits that are cool and new, but which aren’t germane to what I’m working on at the moment. I can’t let those findings interfere with my momentum, but I don’t want to throw them away, either. So I blog them. Also the blog gives me a place to burn off energy at the end of the day, when I can still produce words but don’t have the discipline to write technical prose.

– – – – – – – – – – – –

The photo at the top of the post is of Giraffatitan dorsal vertebrae in a case at the MfN Berlin, from Mike’s and my visit with the DfG 533 group back in late 2008. I picked that photo so I could make the following dumb off-topic observation: with its upturned transverse processes, the dorsal on the right looks like it’s being all faux melodramatic, a la:

This post started out as a comment on this thread, kicked off by Dale McInnes, in which Mike Habib got into a discussion with Mike Taylor about the max size of sauropods. Stand by for some arm-waving. All the photos of outdoor models were taken at Dino-Park Münchehagen back in late 2008.

I think it’s all too easy to confuse how big things do get from how big they could get, assuming different selection pressures and ecological opportunities. I’m sure someone could write a very compelling paper about how elephants are as big as they could possibly be, or Komodo dragons, if we didn’t have indricotheres and Megalania to show that the upper limit is elsewhere. This is basically what Economos (1981) did for indricotheres, either forgetting about sauropods or assuming they were all aquatic.

Truly, a mammal of excellence and distinction. With Mike and some dumb rhino for scale.

In fact, I’ll go further: a lot of pop discussions of sauropod size assume that sauropods got big because of external factors (oxygen levels, etc.) but were ultimately limited by internal factors, like bone and cartilage strength or cardiovascular issues. I think the opposite is more likely: sauropods got big because of a happy, never-repeated confluence of internal factors (the Sander/et al. [2008, 2011, 2013] hypothesis, which I think is extremely robust), and their size was limited by external, ecological factors.

Take a full-size Argentinosaurus or Bruhathkayosaurus – even modest estimates put them at around 10x the mass of the largest contemporary predators. Full-grown adults were probably truly predator-immune, barring disease or senescence. So any resources devoted to pushing the size disparity higher, instead of invested in making more eggs, would basically be wasted.

If there was reproductive competition among the super-giants, could the 100-tonners have been out-reproduced by the 70-tonners, which put those extra 30 tonnes into making babies? Or would the 100-tonners make so many more eggs than the 70-tonners (over some span of years) that they’d still come out on top? I admit, I don’t know enough reproductive biology to answer that. (If you do, speak up in the comments!) But if – if – 70-tonners could out-reproduce 100-tonners, that by itself might have been enough to put a cap on the size of the largest sauropods.

Another possibility is that max-size adult sauropods were neither common nor the target of selection. In most populations most of the time, the largest individuals might have been reproductively active but skeletally-immature and still-growing subadults (keep in mind that category would encompass most mounted sauropod skeletons, including the mounted brachiosaurs in Chicago and Berlin). If such individuals were the primary targets of selection, and they were selected for a balance of reproductive output and growth, then the few max-size adults might represent the relatively rare instances in which the developmental program “overshot” the selection target.

Dave Hone and Andy Farke and I mentioned this briefly in our 2016 paper, and it’s come up here on the blog several times before, but I still have a hard time wrapping my head around what that would mean. Maybe the max-size adults don’t represent the selective optimum, but rather beneficial traits carried to extreme ends by runaway development. It seems at least conceivable that the bodies of such animals might have been heavily loaded with morphological excrescences – like 15- to 17-meter necks – that were well past the selective optimum. As long as those features weren’t inherently fatal, they could possibly have been pretty darned inefficient, riding around on big predator-immune platforms that could walk for hundreds of kilometers and survive on garbage.

What does that swerve into weird-but-by-now-well-trod ground have to do with the limits on sauropod size? This: if max-size adults were not heavy selection targets, either because the focus of selection was on younger, reproductively-active subadults, or because they’d gotten so big that the only selection pressure that could really affect them was a continent-wide famine – or both – then they might not have gotten as big as they could have (i.e., never hit any internally-imposed, anatomical or biomechanical limits) because nothing external was pushing them to get any bigger than they already were.

Or maybe that’s just a big pile of arm-wavy BS. Let’s try tearing it down, and find out. The comment thread is open.


Here’s BYU 12866, a mid-cervical of a neosauropod from Dry Mesa Quarry. It’s cataloged as Brachiosaurus, an identification I’ve never found any compelling reason to doubt. It’s definitely brachiosaurid, and for now Brachiosaurus is the only game in town for the Late Jurassic of North America. I expect that will change when more and better material comes to light, based on the different coracoid shapes of the Brachiosaurus holotype and the “Ultrasauros” scapulocoracoid.

I reckon it’s probably a C5 or so, based on its proportions and comparisons with Giraffatitan (for example).

As you can see, it’s a bit distorted, sheared over with the dorsal side to the right and the ventral side to the left.

I don’t think there’s any major anterior/posterior shearing – the zygs are set forward of their respective centrum ends by about the same amount in this specimen as in Giraffatitan.

Kent Sanders and I CT scanned this vert back in the day and those scans made it into several papers, including Wedel et al. (2000b) on Sauroposeidon and Wedel (2005) on sauropod pneumaticity and mass estimates.

I have the original, uncropped, full-res photos, and I’ll probably get them posted at some point (faster if people bug me to do so, so speak up in the comments if you want them). But for now I’m sticking to getting stuff posted quickly, easily, and regularly, and I found these as-is on my hard drive, so here we are.



Vicki and London and I were in downtown Los Angeles for a friend’s wedding on Dec. 30, and afterward we visited The Last Bookstore. Embarrassingly, even though I’m LA-adjacent, I had not been before. I believe the mounted woolly mammoth visible in the far corner was one of the last ones to be shot in the LA Basin.

The Last Bookstore is an awesome place, with two floors of new and used books, records, comics, and related esoterica. Made me nostalgic for Logos in downtown Santa Cruz, which sadly closed shop this past summer.

The visit was a momentous occasion for me. Although my book with Mark Hallett has been out for almost a year and a half now, and many copies have passed through my hands at book signings, I’d never run into one out in the wild.

I quickly and quietly did a guerilla signing, and left the book on the shelf. And I intend to keep doing them, as often as I run into unsigned copies. As a public service message, if you ever find a copy of the book out in the world that looks like it’s been signed by me, it’s probably legit (send me a pic or post in the comments if you have doubts). Since I’m inflicting these on an unsuspecting public, if you get stuck with a signed copy but would prefer otherwise, let me know and I’ll swap a fresh book for your vandalized one.

I also did okay finding books for myself. I got two: On a Piece of Chalk, by Thomas Henry Huxley, and The Anatomy of the Salamander, by Eric T.B. Francis.

On a Piece of Chalk is a legendary bit of natural history. In 1868, T.H. Huxley gave a public lecture with that title to the working folk of Norwich, during a meeting of the British Association for the Advancement of Science. A piece of chalk was both his physical tool and his subject, which he used to illustrate, literally and figuratively, the evidence for uniformitarian stratigraphy and biological evolution. Huxley’s talk has been printed twice: later in 1868 by Macmillan’s Magazine of London, and in a nice hardback in 1967 by Charles Scribner’s Sons of New York. I found a copy of the latter for five bucks, which I note is the going rate for used copies on Amazon. But I can’t actually find any evidence that my copy has been used. It appears to be utterly pristine, and I suspect it may be New Old Stock.

If you don’t own a copy of this wonderful book, you should drop what you’re doing, acquire one, and read it. If you’re reading this blog, you probably know Huxley’s punchline. But the way Huxley draws the reader in, illustrates his points with clear and compelling examples, and builds his argument steadily outward, from a piece of chalk to the vertiginous spectacle of deep time, is masterful in both concept and execution.

I know less about the salamander anatomy book, but I snagged it anyway. It’s a reprint of an original 1934 text, published in 2002 by the Society for the Study of Amphibians and Reptiles. Reproduction quality is excellent, especially of the numerous and minutely detailed plates. I picked up the book for two reasons: one, because I’m getting progressively more interested in the peripheral nervous systems of nonhuman tetrapods, and two, because I have a peculiar fetish for good illustrations of the recurrent largyneal nerve, especially in short-necked animals (for example). I did not come away disappointed.

The moral of the story? Stay alert for good natural history writing. I find that older natural history books turn up in used bookstores pretty regularly, and it’s possible to grow your library inexpensively if you are patient. And support your local bookstore, while it’s still there to support.

Hey, look, a new sauropod vertebra to kick off the new year!

I’ve blogged a lot about the giant – and tiny – apatosaurines from the Morrison Formation of Oklahoma, and just once on Saurophaganax. But otherwise I don’t think I’ve covered any of the other Oklahoma Morrison dinos. So here’s a start: a pretty decent Camarasaurus dorsal. Broken transverse processes traced from Osborn and Mook (1921). Like all of the Oklahoma Morrison dinos, it’s from the quarries on or near Black Mesa, at the far northwestern corner of the Oklahoma panhandle.

Based on the narrowness of the neural arch and spine, I don’t think this vert can be any farther forward than D6 – anterior Cam dorsals are w-i-d-e. It would be odd for a camarasaur to have a spine split that deeply as far back as D10 or D11 (see Wedel and Taylor 2013). The centrum is very anteriorposteriorly short, which is a posterior dorsal character, but based on Osborn and Mook (1921) the centra can be this short as far forward as D6. So on the balance of the evidence, I think it’s probably a D6 or D7. But that is just an estimate, which might be off by a couple of positions either way.

Tons more that could be said about this specimen, but I’m going to play against type and not write a dissertation for a change. So, here’s OMNH 1811. We’ll probably come back to it at some point.