Well, I’m a moron again. In the new preprint that I just published, I briefly discussed the six species of sauropod for which complete necks are known — Camarasaurus lentus (but it’s a juvenile), Apatosaurus louisae (but the last three and maybe C5 are badly damaged), Mamenchisaurus hochuanensis (but all the vertebrae are broken and distorted), Shunosaurus lii, Mamenchisaurus youngi and Spinophorosaurus nigerensis.

I did have the wit to say, in the Author Comment:

Although I am submitting this article for formal peer-review at the same time as publishing it as a preprint, I also solicit comments from readers. In particular I am very keen to know if I have missed any complete sauropod necks that have been described in the literature. In the final version of the manuscript, I will acknowledge those who have offered helpful comments.

Happily, several people have taken me up on this (see the comments on the preprint), but one suggestion in particular was a real D’oh! moment for me. Oliver Demuth reminded me about Kaatedocus — a sauropod that we SV-POW!sketeers love so much that it has its own category on our site and we’ve held it up as an example of how to illustrate a sauropod specimen. More than that: we have included several illustrations of its vertebrae in one of our own papers.

Aaanyway … the purpose of this post is just to get all the beautiful Kaatedocus multiview images up in one convenient place. They were freely available as supplementary information to the paper, but now seem to have vanished from the publisher’s web-site. I kept copies, and now present them in the conveniently viewable JPEG format (rather the download-only TIFF format of the originals) and with each image labelled with its position in the column.

Please note, these images are the work of Tschopp and Mateus (2012) — they’re not mine!

Atlas and axis (C1-2)

Atlas and axis (C1-2)

























C15 (and the rest of the skeleton) is missing, which makes this a very nearly, but not quite, complete sauropod neck.


  • Tschopp, Emanuel, and Octávio Mateus. 2012. The skull and neck of a new flagellicaudatan sauropod from the Morrison Formation and its implication for the evolution and ontogeny of diplodocid dinosaurs. Journal of Systematic Palaeontology 11(7):853-888. doi:10.1080/14772019.2012.746589

Here’s the last post (at least for now) in the Fighting Apatosaur Art series — and we’re back to Brian Engh, who we started with.

Early in the process of putting together artwork to illustrate our apatosaur neck combat hypothesis, Brian tried out a whole bunch of outlandish concepts. Here are two that he showed us, but which were too speculative to push forward with. First, necks as big, floppy display structures:


As a piece of art, I really like this one: the boldness, the vivid contrasts, the alien quality of the animals. But as a palaeobiological hypothesis, it doesn’t really work: so much of the neck morphology in apatosaurs is to do with absorbing ventral forces that soft-tissue display structures down there don’t make a whole lot of sense.

Here’s the other one — which Brian titles “Apatosaur inflato-porcupine fish neck-bag”.


I particularly like the way the theropod being rolled around on the ground and repeatedly spiked. It’s no more than it deserves.

Does the idea of an inflatable neck make sense? I wouldn’t be at all surprised if there were sauropods that did something like this — plenty of extant animals inflate parts of their body for display purposes, after all — but I don’t think it would have been apatosaurs. Again, the characteristic features of the neck don’t seem well matched to this scenario.

Well, that’s all the apatosaur neck-combat art we have. If there’s to be a part 7 in this series, it will be made of artwork that you, dear readers, have contributed. Fire away!

If we accept that the distinctive ventral projections of the gigantic and ventrally displaced cervical ribs of apatosaurs were likely the base of some form of soft-tissue rugosity — such as keratinous horns like those of rhinos — then does it follow that those necks were used in combat as we suggested?

Maybe, maybe not. As scientists, we are always open to other hypotheses. We’re looking for the simplest, most parsimonious model — the one which best explains the facts.

That’s why we like Mark Witton’s “neck-velcro wall-climbing” hypothesis, as shown in this actual scientific life restoration.


As Mark explained to me, apatosaurs may have used their neck-hooks for more than passive clinging. They may also have been used for inching up the rock-face: first one side of the neck advancing and then the next, in the manner of the “pterygoid walking” that snakes use to progressively swallow large prey.

This is why it’s important to present early-stage work at conferences (and as preprints). Otherwise, you may never hear about important alternative hypotheses like this until after the paper is out and it’s too late to include them.

I mentioned last time that, as I sat next to Bob Nicholls in an SVPCA session, I started sketching an apatosaur combat in the hope that my horrible drawing would provoke Bob to do a good one. That worked admirably, which means there is no good reason for me to subject you to my own sketch.

So here it is.


I think the main lessons to draw from this piece are:

  1. I can’t draw heads.
  2. I can’t draw limbs.
  3. I can’t draw torsos.
  4. I may be just about capable of drawing tails.

In defence of this picture, it does have something of a How And Why Wonder Book of Dinosaurs quality to it, which people of a certain age may find nostalgic. (See also: How fat was Brontosaurus?)

During a break, I asked for Bob’s advice on how I can do better. I know I’ll never be an artist, but it’s fun to sketch (especially during mammal talks) and I’d like to improve a little. The main point Bob made was to think about where the light is coming from. Be consistent about that, and you get an immediate improvement in realism.

So here’s what I sketched the next day, with that in mind:


So what have we learned this time?

  1. I didn’t consciously do this, but I ended up with a composition kind of similar to what Bob came up with, but worse.
  2. In my desire to achieve the intertwined-necks pose, I made the necks too long and thin.
  3. I still can’t draw heads.
  4. Let’s just forget about the hindlimb of the one on the left.
  5. Uh, and let’s forget the torsos, too.
  6. But at least the light is coming from top right!

In short, as Stephen Sondheim put it, art isn’t easy. I wish I had more time to put into it.

The real moral of this story is: if I had a crack at drawing fighting apatosaurs, you definitely can. Let us know if you do — leave a comment. We’ll gather people’s contributions in a future post.

(See also the previous Fighting Apatosaur Art posts: Brian Engh #1, Brian Engh #2, Bob Nicholls. More to come!)

On the morning of Tuesday 1st December, on SVPCA day 1, I gave my talk about apatosaur neck combat. In one of the afternoon sessions, I sat next to Bob Nicholls, and found myself thinking how awesome it would be if he sketched some apato-combat.

But I didn’t want to come right out and say “Hey, Bob, how ’bout you spontaneously illustrate our palaeobiological hypothesis?” So instead I used a tactic that Fiona often uses when she wants me to do something: she starts to do it herself, badly, and waits for me to take over. (This is often how I find myself cooking in the evenings.) In the same spirit, sat next to Bob, I started a horrible sketch of wrestling apatosaurs. Sure enough, Bob, saw what I was doing, internally decided it ought to be done properly, and produced this:

WrestleBronto2a (c)Nicholls2015

What I love most about this (beside the casual way he knocked it out in fifteen minutes) is the sense of heft about the apatosaurs. These are big, solid animals. Someone’s gonna get hurt.


Last time, we looked at some of Brian Engh’s preliminary sketches of pieces to illustrate our fighting-apatosaur hypothesis. But there’s more: some way into the process, Brian also came up with this very rough sketch, illustrating a different style of combat:


All the pictures in the previous post show various forms of ventral-to-ventral combat, but we’ve also been thinking about possibilities, and an important one is ventral-to-dorsal.

That could work in at least two ways. We can imagine a wresting match, where each animal tries to get its neck above its opponent’s, and to force it to the ground. There is precedent for this in the behaviour of various extant animals. (Or perhaps I should call it postcedent, since apatosaurs came first.)

But other extant animals have a much more violent combat style, based on striking blows rather than exerting steady force. Notably, giraffes do this, using their long necks as levers to crash their uncharismatic, highly fused mammalians heads into each other.

Could apatosaurs have done this? Not exactly: their heads were far too small to be effective clubs, and far too fragile to survive being used in this way. But the necks themselves would have been formidable weapons: we’re confident that apatosaurs striking blows would have done so with their necks, bringing them powerfully downwards on their adversaries.

Brian liked this idea enough to work the rough sketch above up into a completed drawing, which we also plan to include in the paper (and which, by the way, I unreservedly love):


So what style of combat did apatosaurs use? Ventral-on-ventral shoving? Wrestling to the ground? Striking downwards blows with the neck?

My best guess (and it’s only a guess, necessarily) is that among the half-dozen or so recognised species of apatosaurine, all these styles were likely in use. And this may explain the variation in cervical morphology that we see between species (though of course ontogeny and sexual dimorphism may also be at work).

In short, I think all of these scenarios are credible — and therefore perfectly legitimate subjects for palaeo-art *hint hint*.

In putting together our thoughts on how apatosaurs used their necks, we were motivated by genuine curiosity — which in Matt’s and my case, at least, goes back many years. (We briefly discussed the problem, if only to throw our hands up in despair, in our 2013 neck-anatomy paper.) We didn’t land on the combat hypothesis because it’s cool, but because it’s where the evidence points.

That said, it is cool.

Brian Engh is on the authorship for this paper largely because of his insights into extant animal behaviour. But there’s no denying that it’s a real bonus that he’s also an awesome artist. He’s been putting together sketches to illustrate our hypothesis for some time, partly with the goal of figuring out which compositions to work up into finished pieces. Here, with Brian’s permission, are some of those preliminary sketches.

First, a really nice sketch showing a ventral-to-ventral shoving match from down at ground level.


I really like this one, and would have been happy for it to be one of the anointed ones. I like the sense of huge beasts towering over the viewer. That said, I always love pencil sketches, often more than I do finished pieces, so I’m not too unhappy that the world gets to see this one in pencil-sketch form.

Next up, sketched more roughly, is a concept for a different form of combat in a different aspect. Here, we see two animals side by side, wrestling with both necks and tails.


I like the dynamism of this one, and especially that the one on the right is in the process of being pushed over. But there’s nothing in apatosaur tail morphology that particularly says “combat”, so I guess I’m not too unhappy that this one didn’t make the cut.

The third sketch shows two individuals rearing into into ventral-to-ventral push.


Matt and Brian liked this one the most, so it got worked up into a finished and coloured piece which will be one of the figures in the paper when we get around to submitting it. Here is the current version — as I understand it, Brian plans to revise it further before it’s done.


The craftsmanship here is superb, but I can’t help regretting that the dinosaurs are rearing less than in the sketch. I feel it’s lost some of the power of the concept sketch.

What you’re seeing here, folks, is a bona fide instance of co-authors disagreeing. Happens all the time, but you usually don’t see it, because it’s all resolved by the time the paper is submitted. Brian is the artist, and ultimately it’s for him to decide what to depict and how; but I’ll always be glad that we still have the pencil-sketch as well as the finished version.



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