We’ve noted that the Taylor et al. SVPCA abstract and talk slides are up now up as part of the SVPCA 2015 PeerJ Collection, so anyone who’s interested has probably taken a look already to see what it was about. (As an aside, I am delighted to see that two more abstracts have been added to the collection since I wrote about it.)

It was my privilege to present a talk on our hypothesis that the distinctive and bizarre toblerone-shaped necks of apatosaurs were an adaptation for intraspecific combat. This talk was based on an in-progress manuscript that Matt is lead-authoring. Also on board is the third SV-POW!sketeer, the silent partner, Darren Naish; and artist/ethologist Brian Engh.

Here is our case, briefly summarised from five key slides. First, let’s take a look at what is distinctive in the morphology of apatosaur cervicals:

Here I’m using Brontosaurus, which is among the more extreme apatosaurs, but the same features are seen developed to nearly the same extent in Apatosaurus louisae, the best-known apatosaur, and to some extent in all apatosaurs.

Now we’ll look at the four key features separately.

First, the cervicals ribs of sauropods (and other saurischians, including birds) anchored the longus colli ventralis and flexor colli lateralis muscles — ventral muscles whose job is to pull the neck downwards. By shifting the attachments points of these muscles downwards, apatosaurs enabled them to work with improved mechanical advantage — that is, to bring more force to bear.

Second, by redirecting the diapophyses and parapophyses ventrally, and making them much more robust than in other sauropods, apatosaurs structured their neck skeletons to better resist ventral impacts.

Third, because the low-hanging cervical ribs created an inverted “V” shape below the centrum, they formed a protective cradle for the vulnerable soft-tissue that is otherwise exposed on the ventral aspect of the neck: trachea, oesophagus, major blood vessels. In apatosaurus, all of these would have been safely wrapped in layers of connective tissue and bubble-wrap-like pneumatic diverticula. The presence of diverticula ventral to the vertebral centrum is not speculative – most neosauropods have fossae on the ventral surfaces of their cervical centra, and apatosaurines tend to have foramina that connect to internal chambers as well (see Lovelace et al. 2007: fig. 4, which is reproduced in this post).

Fourth, most if not all apatosaurs have distinctive ventrally directed club-like processes on the front of their cervical ribs. (It’s hard to tell with Apatosaurus ajax, because the best cervical vertebra of that species is so very reconstructed.) How did these appear in life? It’s difficult to be sure. They might have appeared as a low boss; or, as with rhinoceros horns, they might even have carried keratinous spikes.

Putting it all together, we have an animal whose neck can be brought downwards with great force; whose neck was mechanically capable of resisting impacts on its ventral aspect; whose vulnerable ventral-side soft-tissue was well protected; and which probably had prominent clubs or spikes all along the ventral aspect of the neck. And all of this was accomplished at the cost of making the neck a lot heavier than it would have been otherwise. Off the cuff, it seems likely that the cervical series alone would have massed twice as much in apatosaurines as in diplodocines of the same neck length.

Doubling the mass of the neck is a very peculiar thing for a sauropod lineage to do – by the Late Jurassic, sauropods were the leading edge of an evolutionary trend to lengthen and lighten the neck that had been running for almost 100 million years, through basal ornithodirans, basal dinosauromorphs, basal saurischians, basal sauropodomorphs, and basal sauropods. Whatever the selective pressures that led apatosaurines to evolve such robust and heavy necks, they must have been compelling.

The possibility that apatosaurs were pushing or crashing their necks ventrally in some form of combat accounts for all of the weird morphology documented above, and we know that sexual selection is powerful force that underlies a lot of bizarre structures in extant animals, and probably in extinct ornithodirans as well (see Hone et al. 2012, Hone and Naish 2013).

What form of combat, exactly? There are various possibilities, which we’ll discuss another time. But I’ll leave you with Brian Engh’s beautiful illustration of one possible form of combat: a powerful impact of one neck brought down onto the dorsal aspect of another.

We’re aware that this proposal is necessarily somewhat speculative. But we’re just not able to see any other explanation for the distinctive apatosaur neck. Even if we’re wrong about the ventrolateral processes on the cervical ribs supporting bosses or spikes, the first three points remain true, and given how they fly in the face of sauropods’ long history of making their necks lighter, they fairly cry out for explanation. If anyone has other proposals, we’ll be happy to hear them.

References

• Hone, D. W., Naish, D., & Cuthill, I. C. (2012). Does mutual sexual selection explain the evolution of head crests in pterosaurs and dinosaurs?. Lethaia 45(2):139-156.
• Hone, D. W. E., & Naish, D. (2013). The ‘species recognition hypothesis’ does not explain the presence and evolution of exaggerated structures in non‐avialan dinosaurs. Journal of Zoology 290(3):172-180.
• Lovelace, D. M., Hartman, S. A., & Wahl, W. R. (2007). Morphology of a specimen of Supersaurus (Dinosauria, Sauropoda) from the Morrison Formation of Wyoming, and a re-evaluation of diplodocid phylogeny. Arquivos do Museu Nacional, Rio de Janeiro 65(4):527-544.

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.

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*.

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:

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.

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!)

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.

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!

I’d hoped that we’d see a flood of BRONTOSMASH-themed artwork, but that’s not quite happened. We’ve seen a trickle, though, and that’s still exciting. Here are the ones I know about. If anyone knows of more, please let me know and I will update this post.

First, in a comment on the post with my own awful attempts, Darius posted this sketch of a BROTOSMASH-themed intimidation display:

And in close-up:

Very elegant, and it’s nice to see an extension of our original hypothesis into other behaviours.

The next thing I saw was Mark Witton’s beautiful piece, described on his own site (in a post which coined the term BRONTOSMASH):

And in close-up:

I love the sense of bulk here — something of the elephant-seal extant analogue comes through — and the subdued colour scheme. Also, the Knight-style inclusion in the background of the individual in the swamp. (No, sauropods were not swamp-bound; but no doubt, like elephants, they spent at least some time in water.)

And finally (for now, at least) we have Matthew Inabinett’s piece, simply titled BRONTOSMASH:

I love the use of traditional materials here — yes, it still happens! — and I like the addition of the dorsal midline spike row to give us a full on TOBLERONE OF DOOM. (Also: the heads just look right. I wish I could do that. Maybe one day.)

Update (Monday 26 October)

Here is Oliver Demuth’s sketch, as pointed out by him in a comment.

Thanks, Oliver! Nice to see the ventral-on-dorsal combat style getting some love.

So that’s where we are, folks. Did I miss any? Is anyone working on new pieces on this theme? Post ’em in the comments!

Just a quick heads up for any SV-POW! readers within convenient striking distance of Norman, Oklahoma, this Wednesday, March 16. Like all of the lectures in the “Dinosaurs Past & Present” series at OMNH, this one is free to the public. I hope to see some of you there!

When I was back in Oklahoma in March, I met with Anne Weil to see some of the new Apatosaurus material she’s getting out of her Homestead Quarry. It’s nice material, but that’s a post for another day. Anne said something that really resonated with me, which was, “I love it when you guys post about vertebral morphology, because it helps me learn this stuff.” Okay, Anne, message received. This will begin to make things right.

I spent a week at BYU back in 2005, collecting data for my dissertation. One of the first things I had to do was teach myself how to identify the vertebrae of different sauropods, because BYU has just about all of the common Morrison taxa. These are the notes I made back then.

I always planned to do something with them – clean them up, get them into a more usable form. There are a lot of scribbly asides that are probably hard for others to read, and it would be more useful if I put the easily confused taxa next to each other – Barosaurus next to Brachiosaurus, for example. And I didn’t go into serial changes at all.

Still, hopefully someone will find these useful. If there are things I missed or got wrong, the comment thread is open. And if you want all four spreads in one convenient package, here’s a PDF: Wedel 2005 notes on Morrison sauropod cervicals

Mike and I leave for the Sauropocalypse tomorrow. I’m hoping to post at least a few pretty pictures from the road, as I did for the Mid-Mesozoic Field Conference two years ago. Stand by…

Things remain frantic on the Sauropocalypse tour. Today, we were back at the BYU Museum of Paleontology, working on four or five separate projects. Here’s Matt, photographing broken bone of the iconic Supersaurus cervical BYU 9024, while a pallet of Big Pink Apatosaur cervicals wait for attention in the background:

You’ve seen this bone before – I first posted on it 8 years ago this month, and it turned up again here and here. It is still the longest known vertebra of any animal that has ever lived.

And here’s Mike, getting Jensen’s sculpture of the same vertebra down from storage to compare it to the original:

In Jensen’s (1985) original description of this vertebra – which he at first referred to Ultrasauros – the only relevant illustration he included was one of the model, so it was good to see this bit of history in the flesh (Jensen did include photos of the actual bone in later papers). We’ll show the two vertebrae, real and sculpted, side by side in a future post.

References

• Jensen, J. A. 1985. Three new sauropod dinosaurs from the Upper Jurassic of Colorado. Great Basin Naturalist 45, 697-709.

Quick heads up: Mark Hallett and I are both at the Society of Vertebrate Paleontology meeting in Salt Lake City. Tomorrow afternoon (Friday, October 28) at 4:15 PM we’ll be signing copies of our book, The Sauropod Dinosaurs: Life in the Age of Giants. If you’d like to get a copy of the book, or to have your already-purchased copy signed, please come to the Johns Hopkins University Press booth in the exhibitor/poster area tomorrow afternoon. We’re both generally happy to sign books whenever and wherever, but if you’d like to catch us both at the same time, this is a good opportunity. We’re hoping to do another joint book signing in Los Angeles before long – more info on that when we get it arranged.

In the meantime, or if you’re not at SVP, or if you just like cool things, check out this rad claymation video of fighting apatosaurs, by YouTube user Fred the Dinosaurman. I love this. My favorite thing is that if you’re familiar with the previously-produced, static visual images of neck-fighting apatosaurs (links collected here), you’ll see a lot of those specific poses and moments recreated as transient poses in the video. This was published back in June, but I’d missed it – many thanks to Brian Engh for the heads up.

Fig. 14. Vertebrae of Pleurocoelus and other juvenile sauropods. in right lateral view. A-C. Cervical vertebrae. A. Pleurocoelus nanus (USNM 5678, redrawn fromLull1911b: pl. 15). B. Apatosaurus sp. (OMNH 1251, redrawn from Carpenter &McIntosh 1994: fig. 17.1). C. Camarasaurus sp. (CM 578, redrawn from Carpenter & McIntosh 1994: fig. 17.1). D-G. Dorsal vertebrae. D. Pleurocoelus nanus (USNM 4968, re- drawn from Lull 1911b: pl. 15). E. Eucamerotus foxi (BMNH R2524, redrawn from Blows 1995: fig. 2). F. Dorsal vertebra referred to Pleurocoelus sp. (UMNH VP900, redrawn from DeCourten 1991: fig. 6). G. Apatosaurus sp. (OMNH 1217, redrawn from Carpenter & McIntosh 1994: fig. 17.2). H-I. Sacral vertebrae. H. Pleurocoelus nanus (USNM 4946, redrawn from Lull 1911b: pl. 15). I. Camarasaurus sp. (CM 578, redrawn from Carpenter & McIntosh 1994: fig. 17.2). In general, vertebrae of juvenile sauropods are characterized by large pneumatic fossae, so this feature is not autapomorphic for Pleurocoelus and is not diagnostic at the genus, or even family, level. Scale bars are 10 cm. (Wedel et al. 2000b: fig. 14)

The question of whether sauropod cervicals got longer through ontogeny came up in the comment thread on Mike’s “How horrifying was the neck of Barosaurus?” post, and rather than bury this as a comment, I’m promoting it to a post of its own.

The short answer is, yeah, in most sauropods, and maybe all, the cervical vertebrae did lengthen over ontogeny. This is obvious from looking at the vertebrae of very young (dog-sized) sauropods and comparing them to those of adults. If you want it quantified for two well-known taxa, fortunately that work was published 16 years ago – I ran the numbers for Apatosaurus and Camarasaurus to see if it was plausible for Sauroposeidon to be synonymous with Pleurocoelus, which was a real concern back in the late ’90s (the answer is a resounding ‘no’). From Wedel et al. (2000b: pp. 368-369):

Despite the inadequacies of the type material of Pleurocoelus, and the uncertainties involved with referred material, the genus can be distinguished from Brachiosaurus and Sauroposeidon, even considering ontogenetic variation. The cervical vertebrae of Pleurocoelus are uniformly short, with a maximum EI of only 2.4 in all of the Arundel material (Table 4). For a juvenile cervical of these proportions to develop into an elongate cervical comparable to those of Sauroposeidon, the length of the centrum would have to increase by more than 100% relative to its diameter. Comparisons to taxa whose ontogenetic development can be estimated suggest much more modest increases in length.

Carpenter & McIntosh (1994) described cervical vertebrae from juvenile individuals of Apatosaurus and Camarasaurus. Measurements and proportions of cervical vertebrae from adults and juveniles of each genus are given in Table 4. The vertebrae from juvenile specimens of Apatosaurus have an average EI 2.0. Vertebrae from adult specimens of Apatosaurus excelsus and A. louisae show an average EI of 2.7, with an upper limit of 3.3. If the juvenile vertebrae are typical for Apatosaurus, they suggest that Apatosaurus vertebrae lengthened by 35 to 65% relative to centrum diameter in the course of development.

The vertebrae from juvenile specimens of Camarasaurus have an average EI of 1.8 and a maximum of 2.3. The relatively long-necked Camarasaurus lewisi is represented by a single skeleton, whereas the shorter-necked C. grandis, C. lentus, and C. supremus are each represented by several specimens (McIntosh, Miller, et al. 1996), and it is likely that the juvenile individuals of Camarasaurus belong to one of the latter species. In AMNH 5761, referred to C. supremus, the average EI of the cervical vertebrae is 2.4, with a maximum of 3.5. These ratios represent an increase in length relative to diameter of 30 to 50% over the juvenile Camarasaurus.

If the ontogenetic changes in EI observed in Apatosaurus and Camarasaurus are typical for sauropods, then it is very unlikely that Pleurocoelus could have achieved the distinctive vertebral proportions of either Brachiosaurus or Sauroposeidon.

C6 of Apatosaurus CM 555 – despite having an unfused neural arch and cervical ribs, the centrum proportions are about the same as in an adult.

A few things about this:

1. From what I’ve seen, the elongation of the individual vertebrae over ontogeny seems to be complete by the time sauropods are 1/2 to 2/3 of adult size. I get this from looking at mid-sized subadults like CM 555 and the hordes of similar individuals at BYU, the Museum of Western Colorado, and other places. So to get to the question posed in the comment thread on Mike’s giant Baro post – from what I’ve seen (anecdata), a giant, Supersaurus-class Barosaurus would not necessarily have a proportionally longer neck than AMNH 6341. It might have a proportionally longer neck, I just haven’t seen anything yet that strongly suggests that. More work needed.
2. Juvenile sauropod cervicals are not only shorter than those of adults, they also have less complex pneumatic morphology. That was the point of the figure at the top of the post. But that very simple generalization is about all we know so far – this is an area that could use a LOT more work.
3. I’ve complained before about papers mostly being remember for one thing, even if they say many things. This is the canonical example – no-one ever seems to remember the vertebrae-elongating-over-ontogeny stuff from Wedel et al. (2000b). Maybe that’s an argument for breaking up long, kitchen-sink papers into two or more separate publications?

Reference

Wedel, M.J., Cifelli, R.L., and Sanders, R.K. 2000b. Osteology, paleobiology, and relationships of the sauropod dinosaur Sauroposeidon. Acta Palaeontologica Polonica 45:343-388.

Turns out that if Mike and I don’t post about sauropods for a while, people start doing it for us! This very interesting project by Tom Johnson of Loveland, Colorado, first came to my attention when Tom emailed Mark Hallett about it and Mark kindly passed it on to me. I got in touch with Tom and asked if he’d be interested in writing it up for SV-POW!, and here it is. Many thanks to Tom for his willingness to share his work with us. Enjoy! – Matt Wedel

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The sauropod formerly known as Apatosaurus in the American Museum of Natural History was the first permanently mounted sauropod dinosaur in the world, and for many years, the most famous (Brinkman 2010). The greater part of the skeleton consists of the specimen AMNH 460 from the Nine Mile Crossing Quarry north of Como Bluff, Wyoming, supplemented with bones from other AMNH specimens from Como Bluff, Bone Cabin Quarry, and with plaster casts of the forelimbs of the holotype specimen of Brontosaurus excelsus (YPM 1980) at the Yale Peabody Museum.

A herd of Brontosaurus skeleton models parading before four box covers issued between the 1950s and 1990s.

Like many aging boomer dinophiles, my dinosaur epiphany was the result of books, movies, and toys available in the 1950s, but especially a series of plastic model dinosaur skeletons that appeared around 1958. The Brontosaurus was my personal favorite, and, like the Tyrannosaurus and Stegosaurus models in the series, was very obviously based on the AMNH mount. The models were reissued at least three times over the years and can still be found either “mint in box” or more often in various stages of completion.

Apatosaurus lousiae 1/12 scale skeleton, modelled by Phil Platt, assembled and photographed by Brant Bassam. Image courtesy of BrantWorks.com.

The crème de la crème today, of course, is the 1:12 scale Apatosaurus skeleton model by Phil Platt, available from Gaston Design in Fruita, Colorado. A particularly nice example is the one completed and mounted by Brant Bassam of BrantWorks. The Platt skeleton is a replica in the true sense of the word. The plastic models are pretty crude in comparison, as cool as they appeared to us as kids.

I was interested in skeletal illustrations I have seen of Tyrannosaurus rex, which compare the completeness of various specimens by showing the actual bones included by coloring them red. A 2005 study of Apatosaurus by Upchurch et. al. examined eleven of the most complete Apatosaurus individuals, and I was interested to see the actual bones known for each specimen. Using published descriptions, red markers, and copies of a skeletal silhouette of Apatosaurus (permission obtained from the artist), I prepared a comparison of the most completely known Apatosaurus specimens. It was clear, of course, that Apatosaurus louisae (CM 3018) is the most complete specimen of the Apatosaurus/Brontosaurus group. But it also was apparent that old AMNH 460 included a substantial portion of the skeleton, even if it is a composite.

I grabbed some additional markers and, using the illustration of the mount in William Diller Matthew’s popular book Dinosaurs (Matthew 1915, fig. 20, which I trust is in the public domain by now), I color-coded the bones according to the composition as listed in Matthew’s (1905) article:

• AMNH 460, Nine-Mile Crossing Quarry: 5th, 6th, 8th to 13th cervical vertebrae; 1st to 9th dorsal; 3rd to 19th caudal; all ribs; both coracoids; “parts of” sacrum and ilia; both ischia and pubes; left femur and astragalus; and “part of” the left fibula. RED
• AMNH 222, Como Bluff: right scapula, 10th dorsal vertebra, right femur and tibia. GREEN
  (Visitors to AMNH: you can see the rest of AMNH 222 under the feet of the hunched-over Allosaurus)
• AMNH 339, Bone Cabin Quarry: 20th to 40th caudal vertebrae. LIGHT BLUE
• AMNH 592, Bone Cabin Quarry: metatarsals of the right hind foot. VIOLET
• YPM 1980, Como Bluff: left scapula, forelimb long bones (casts). YELLOW
• The remaining parts of the skeleton are either modeled in plaster or are unspecified (“a few toe bones”). BLACK

It occurred to me that I might have sufficient spare parts of old ITC and Glencoe Brontosaurus models to create a three-dimensional version. I did, and painting prior to assembly definitely made the job easier.

There are obviously limitations to using Matthew’s (1915) reconstruction (e.g., only 13 cervicals) and the model (12 cervicals). It is also not clear from Matthew’s description how much of the sacrum and ilia were restored. Nevertheless, the painted model does provide a colorful, if crude, visualization of the composition of the composite.

Here are some more photos of the finished product:

A view from the front of the model, compared with a historical AMNH photo of the forelimbs and pelvic girdle.

Long considered a specimen of Brontosaurus excelsus or Apatosaurus excelsus, AMNH 460 was referred to Apatosaurus ajax by Upchurch et. al. in 2005. In the most comprehensive analysis of diplodocid phylogeny to date, Tschopp et. al. (2015) found AMNH 460 to be an “indeterminate apatosaurine” pending a “detailed analysis of the specimen.” What to call it? Oh, yeah, that’s been covered in another post!

This is a nostalgia shot for the old brontophiles. Notice that the Triceratops is entering the lake for a swim!

Tom Johnson with the mounted skeleton of Amphicyon, a Miocene “bear-dog”,
in the Raymond Alf Museum of Paleontology in Claremont, California.

References

• Brinkman , Paul D. (2010). The Second Jurassic Dinosaur Rush, University of Chicago Press, 2010.
• Matthew, William Diller, (1905). “The Mounted Skeleton of Brontosaurus,” The American Museum Journal, Vol. V, No. 2, April.
• Matthew, W.D. (1915). Dinosaurs, With Special Reference to the American Museum Collections, American Museum of Natural History, New York.
• Tschopp, Emanuel, Octávio Mateus, and Roger Benson. (2015). “A Specimen-Level Phylogenetic Analysis and Taxonomic Revision of Diplodocidae (Dinosauria, Sauropoda).” Ed. Andrew Farke. PeerJ 3 (2015): e857.
• Upchurch, P., Tomida, Y., Barrett, P.M. (2005). “A new specimen of Apatosaurus ajax (Sauropoda: Diplodocidae) from the Morrison Formation (Upper Jurassic) of Wyoming, USA”. National Science Museum Monographs (Tokyo) 26 (118): 1–156.

Here’s a dorsal vertebra of Camarasaurus in anterior view (from Ostrom & McIntosh 1966, modified by Wilson & Sereno 1998). It is one of the most disturbing things I have ever seen in a sauropod. It makes my skin crawl.

Here’s why: the centrum and the thing we habitually call the ‘neural arch’ aren’t fully fused, and as this modified version makes clear, the ‘neural arch’ is neither neural nor an arch. Instead of being bounded ventrally by the centrum and dorsally and laterally by the neural arch, the neural canal lies entirely below the synchondrosis between the not-really-an-arch and the centrum.

Why?! WHY WOULD YOU DO THAT, CAMARASAURUS? This is not ‘Nam. This is basic vertebral architecture. There are rules.

Look at c6 of Apatosaurus CM 555 here, behaving as all good vertebrae ought to. Neural arch be archin’, as the kids say.

And if you are seeking solace in the thought that maybe the artist just drew that Cam dorsal incorrectly, forget it. I’ve been to Yale and examined the original specimen. I’ve seen things, man!

Camarasaurus isn’t the only pervert around here. Check this out:

Unfused neural arch of a caudal vertebra of a juvenile Alamosaurus from Big Bend. And I mean, this is a neural arch. This may be the most neural of all neural arches, in that it contains the entire neural canal. It’s more of a neural…ring, I guess. That’s right, this Alamosaurus caudal is batting for the opposite team from the Cam dorsal above. And it’s a team that neither you nor I play on, because we have well-behaved normal-ass vertebrae with neural arches that actually arch, and then stop, like God and Richard Owen intended.

Scientifically, my question about these vertebrae is: well, that is, I mean to say, what!? I think they have damaged me in some fundamental way.

If you have anything more intelligent to add (or even less intelligent – consider the gauntlet thrown down!), the comment thread is open.

References

• Ostrom, John H., and John S. McIntosh. 1966. Marsh’s Dinosaurs. Yale University Press, New Haven and London. 388 pages including 65 absurdly beautiful plates.
• Wilson, J. A. and Paul C. Sereno. 1998. Early evolution and higher-level phylogeny of sauropod dinosaurs. Society of Vertebrate Paleontology, Memoir 5: 1-68.

Or, how a single lateral fossa becomes two foramina: through a finely graded series of intermediate forms. Darwin would approve. The ‘oblique lamina’ that separates the paired lateral foramina in C6 starts is absent in C2, but C3 through C5 show how it grows outward from the median septum. How do I know it grows outward, instead of being left behind during the pneumatization of the more posterior cervicals? Because with very few exceptions, all neosauropod cervicals start out with a single lateral fossa on each side, as illustrated in this post. But many of them end up with two or more foramina. Diplodocus is a nice example of this (from Hatcher 1901: plate 3):

I should clarify that the vertebrae above show that character transformation in this individual, at this point in its ontogeny. The vertebrae of CM 555 are about two-thirds the size of those of CM 3018, the holotype of A. louisae. In CM 3018, even C4 and C5 have completely divided lateral fossae, corresponding to the condition in C6 of CM 555.

As Mike and I discussed in our 2013 neural spine bifurcation paper, isolated sauropod cervicals require cautious interpretation because the morphology of the vertebrae changes so much along the series. The simple morphology of anterior cervicals reflects both earlier ontogenetic stages and more primitive character states. As Mike says, in sauropod necks, serial position recapitulates both ontogeny and phylogeny. So if you have a complete series, you can do something pretty cool: see the intermediate stages by which simple structures become complex.

If you’re thinking this might have something to do with my impending SVPCA poster, you’re right. Here’s the abstract.

For more on serially increasing complexity in sauropodomorph cervicals, see this post.

This was an interesting exercise. It was my first time generating a poster to be delivered at a conference since 2006. Scientific communication has evolved a lot in the intervening decade, which spans a full half of my research career to date. So I had a chance to take the principles that I say that I admire and try to put them into practice.

It helped that I wasn’t working alone. Jann and Brian both provided strong, simple images to help tell the story, and Mike and I were batting ideas back and forth, deciding on what we could safely leave out of our posters. Abstracts were the first to go, literature cited and acknowledgments were next. We both had the ambition of cutting the text down to just figure captions. Mike nailed that goal, but my poster ended up being slightly more narrative. I’m cool with that – it’s hardly text-heavy, especially compared with most of my efforts from back when. Check out the text-zilla I presented at SVP back in 2006, which is available on FigShare here. I am happier to see, looking back, that I’d done an almost purely image-and-caption poster, with no abstract and no lit cited, as early as 1999, with Kent Sanders as coauthor and primary art-generator – that one is also on FigShare.

I took 8.5×11 color printouts of both my poster and Mike’s, and we ended up passing out most of them to people as we had conversations about our work. That turned out to be extremely useful – I had a 30-minute conversation about my poster at a coffee break the day before the posters even went up, precisely because I had a copy of it to hand to someone else. Like Mike, I found that presenting a poster resulted in more and better conversations than giving a talk. And it was the most personally relaxing SVPCA I’ve ever been to, because I wasn’t staying up late every night finishing or practicing my talk.

I have a lot of stuff to say about the conference, the field trip, the citability of abstracts and posters (TL;DR: I’m for it), and so on, but unfortunately no time right now. I’m just popping in to get this posted while it’s still fresh. Like Mike’s poster, this one is now published alongside my team’s abstract on PeerJ PrePrints.

I will hopefully have much more to say about the content in the future. This is a project that Jann, Brian, and I first dreamed up over a decade ago, when we were grad students at Berkeley. Mike provided the impetus for us to get it moving again, and kindly stepped aside when I basically hijacked his related but somewhat different take on ontogeny and serial homology. When my fall teaching is over, I’m hoping that the four of us can take all of this, along with additional examples found by Mike that didn’t make it into this presentation, and shape it into a manuscript. I’ll keep you posted on that. In the meantime, the comment field is open. For some related, previously-published posts, see this one for the baby sauropod verts, this one for CM 555, and this one for Plateosaurus.

Flying over Baffin Island on the way home.

And finally, since I didn’t put them into the poster itself, below are the full bibliographic references. Although we didn’t mention it in the poster, the shell apex theory for inferring the larval habits of snails was first articulated by G. Thorson in 1950, which is referenced in full here.

Literature Cited

• Bair, A.R. 2007. A model of wear in curved mammal teeth: controls on occlusal morphology and the evolution of hypsodonty in lagomorphs. Paleobiology 33(1):53-75.
• Gilmore, C.W. 1936. Osteology of Apatosaurus with special reference to specimens in the Carnegie Museum. Memoirs of the Carnegie Museum 11: 175-300.
• Hatcher, J.B. 1901. Diplodocus (Marsh): its osteology, taxonomy, and probable habits, with a restoration of the skeleton. Memoirs of the Carnegie Museum 1:1-63.
• Kraatz, B.P., Meng, J., Weksler, M. and Li, C. 2010. Evolutionary patterns in the dentition of Duplicidentata (Mammalia) and a novel trend in the molarization of premolars. PloS one, 5(9), p.e12838.
• Sych, L. 1975. Lagomorpha from the Oligocene of Mongolia. Palaeontogia Polonica 33:183-200.
• Thorson, G. 1950. Reproductive and larval ecology of marine bottom invertebrates. Biological Reviews 25(1):1-45.
• Wedel, M.J., and Taylor, M.P. 2013. Neural spine bifurcation in sauropod dinosaurs of the Morrison Formation: ontogenetic and phylogenetic implications. Palarch’s Journal of Vertebrate Palaeontology 10(1):1-34. ISSN 1567-2158.
• Wedel, M.J., Cifelli, R.L., and Sanders, R.K. 2000b. Osteology, paleobiology, and relationships of the sauropod dinosaur Sauroposeidon. Acta Palaeontologica Polonica 45:343-388.

Back in the spring of 1998, Kent Sanders and I started CT scanning sauropod vertebrae. We started just to get a baseline for the Sauroposeidon project, but in time the data we collected formed the basis for my MS thesis, and for a good chunk of my dissertation as well. Mostly what we had available to scan was Morrison material. Between imperfect preservation, inexpert prep (by WPA guys back in the ’30s), and several moves over the decades, most of the verts from the Oklahoma Morrison have their neural spines and cervical ribs broken off. One of the first things I had to figure out was how to tell broken vertebrae of Camarasaurus from those of Apatosaurus (at the time; Brontosaurus is back in contention now). Here’s a thing I made up to help me sort out cervical centra of Camarasaurus and whatever the Oklahoma apatosaurine turns out to be. It’s a recent production, but it embodies stuff from my notebooks from 20 years ago. Should be useful for other times and places in the Morrison as well, given the broad spatiotemporal overlap of Camarasaurus and the various apatosaurines.

For a related thing in the same vein, see Tutorial 30: how to identify Morrison sauropod cervicals.

More elephant seals soon, I promise.

UPDATE 20 Feb 2018

Ken Carpenter sent this by email, with a request that I post it as a comment. Since it includes an image, I’m appending to the post, because it makes an important point that I neglected to mention.

Ken: Sorry, Matt. Not so easy. The last cervical of Camarasaurus from the Cleveland Lloyd Quarry is more apatosaurine-like than Camarasaurus-like based on your posting. Note the position of both zygapohyses with both ends of the centrum.

My response: Yes, good catch. I meant to say in the post that my distinguishing characters break down at the cervico-dorsal transition. Even so, in this Cleveland Lloyd vert the postzyg is still forward of a line drawn directly up from the cotyle. I’ve never seen that in an apatosaurine–going into the dorsal series, the postzygs tend to be centered over a line projected up from the rim of the cotyle. (If anyone knows of counterexamples, speak up!)

For distinguishing cervico-dorsals, apatosaurines tend to have much taller neural spines than Camarasaurus, and this carries on through the rest of the dorsal series. In apatosaurine dorsals, the height of the spine above the transverse processes always equals or exceeds the height of the arch below the transverse processes. In Camarasaurus, the height of the dorsal neural spines is always less than or equal to the height of the arch. The shapes of the spines are fairly different, too. Maybe that will be the subject of a future post.

Last Wednesday, May 9, Brian Engh and I bombed out to Utah for a few days of paleo adventures. Here are some highlights from our trip.

We started at a Triassic tracksite on Thursday. But I’m not going to post any pictures of the tracks – those will be coming to a Brian Engh joint near you in the future. Instead, I’m going to talk about this little male collared lizard whose territory included the tracksite. He was fearless – didn’t want to run off and leave us yahoos wandering around his patch of desert unsupervised. Brian tickled his chin at one point.

Getting this close to him is how I got shots like this one:

Click through to the big version, it’s worth it.

One more shot of a couple of cool desert dwellers. I was so fixated on the lizard that I didn’t realize until later that Brian was in the frame, taking a much-needed hydration break.

On Friday we had a temporary breaking of the fellowship. I went to Fruita, Colorado, to visit the Dinosaur Journey museum. You’ve seen photos from DJ here before, from the 2014 Mid-Mesozoic Field Conference and the 2016 Sauropocalypse. Here’s an apatosaur pubis with some obvious bite marks on the distal end. This is on display next to a similarly-bitten ischium, which is shown in the MMFC14 post linked above.

Here’s a big apatosaur cervical, in antero-ventral view, with a dorsal rib draped over its left side. The cervical ribs are not fused in this specimen, so it was probably still growing. Here’s a labeled version:

The short centrum and nearly-vertical transverse processes indicate that this is a pretty posterior cervical, possibly a C13 or thereabouts. This specimen was over the fence in the exhibit area and I couldn’t throw a scale bar at it, but I’d describe it as “honkin'”. Like most of the apatosaur material at DJ, this vert is from the Mygatt-Moore Quarry.

Of course the real reason I was at Dinosaur Journey was to see the Snowmass Haplocanthosaurus that John Foster and I described back in 2014. You may remember that its caudal vertebrae have wacky neural canals. You may also have noticed a recent uptick in the number of posts around here about wacky neural canals. The game is afoot.

But as cool as they were, the Triassic tracks, the collared lizard, and even the Snowmass Haplo were only targets of opportunity. Brian and I had gone to Utah for this:

That photo was taken by Paige Wiren of Salt Lake City, on the day that she discovered that bone eroding out of a riverbank, just as you see it.

Here’s Paige with the element, which proved to be the left femur of an apatosaurine sauropod. It’s face down in these photos, so we’re looking at the medial side. The articular head is missing from the proximal end – it should be facing toward Paige’s right knee in the above photo – but other than that and a few negligible nicks and dings, the femur was complete and in really good shape.

Paige did the right thing when she found the femur: she contacted a paleontologist. Specifically, she asked a friend, who in turn put her in touch with Carrie Levitt-Bussian, the paleontology Collections Manager at the Natural History Museum of Utah. Based on Paige’s photos and maps, Carrie was able to identify the element as a dinosaur femur, probably sauropod, within the territory of the BLM Hanksville Field Office. John Foster, the Director of the Museum of Moab, has a permit to legally collect vertebrate fossils from that area, and he works on sauropods, so Carrie put Paige in touch with John and with ReBecca Hunt-Foster, the district paleontologist for the BLM’s Canyon Country District in Utah.

Now, I know there’s a lot of heated rhetoric surrounding the Bureau of Land Management, but whatever your political bent, remember this: those are our public lands. Therefore the fossils out there are the collective property of all of us, and we should all be upset if they get poached or vandalized. Yes, that is a big problem – the Brontomerus type quarry was partially poached before the bones we have now were recovered, and vandalism at public fossil sites in Utah made the national news while we were out there.

So that’s what we went to do: salvage this bone for science and education before it could be lost to erosion or asshats. Brian and I were out there to assist John, ReBecca, and Paige, who got to see her find come out of the ground and even got her hands dirty making the plaster jacket. Brian and John headed out to the site Friday morning and met up with Paige there, and ReBecca and I caravanned out later in the day, after I got back from Fruita.

But I’m getting ahead of myself a bit. We didn’t have to jacket the whole thing. It had naturally broken into three pieces, with thin clay infills at the breaks. So we just slid the proximal and middle thirds away as we uncovered them, and hit any loose-looking pieces with consolidant. The distal third was in more questionable shape, so we did make a partial jacket to hold it together.

We also got to camp out in gorgeous country, with spectacular (and welcome) clouds during the day and incredible starry skies at night.

We floated the femur out of the site using the Fosters’ canoe at the end of the day on Saturday, and loaded up to head back to Moab on Sunday. At one point the road was empty and the sky was not, so I stood on the center line and took some photos. This one is looking ahead, toward I-70 and Green River.

And this one is looking behind, back toward Hanksville.

Here are John and Brian with the femur chunks in one of the back rooms of the Museum of Moab. The femur looks oddly small here, but assembled it was 155 cm (5’1″) long and would have been 160 (5’3″) or more with the proximal head. Smaller than CM 3018 and most of the big mounted apatosaurs, but nothing to sneeze at.

What happens to it next? It will be cleaned, prepped, and reassembled by the volunteers and exhibit staff at the Museum of Moab, and eventually it will go on public display. Thousands of people will get to see and learn from this specimen because Paige Wiren made the right call. Go thou and do likewise.

That was the end of the road for the femur (for now), but not for Brian and me. We had business in Cedar City and St. George, so we hit the road Sunday afternoon. Waves of rainclouds were rolling east across Utah while we were rolling west, with breaks for sunlight in between. I miiiight have had to swerve a couple of times when all the scenery distracted me from driving, and I definitely made an obnoxious number of stops to take pictures.

I don’t remember which scenic overlook this was, but it was a pretty darned good view. This is another one that will reward embiggening – check out those mesas marching off into the distance.

In Cedar City we were guests of Andrew R.C. Milner, Site Paleontologist and Curator at the St. George Dinosaur Discovery Site at Johnson Farm (SGDS). We spent most of Monday at SGDS, getting our minds comprehensively blown by the amazing trace and body fossils on display. It was my first time visiting that museum, but it sure as heck won’t be the last.

I didn’t take nearly enough photos in St. George – too busy helping Brian do some filming for a future project – but I did get this gem. This is a Eubrontes track, from a Dilophosaurus-sized theropod. This is a positive track, a cast of the dinosaur’s foot made by sandy sediment that filled the natural mold formed when the dino stepped into mud. The high clay content of the mud recorded the morphology of the foot in fine detail, including the bumps of individual scales on the foot pads. The vertical streaks were cut into the side of the track by similar scales as the animal’s foot pushed into the mud.

The full story of the Johnson Farm tracks and trackmakers is beautifully told in the book Tracks in Deep Time: The St. George Dinosaur Discovery Site at Johnson Farm, by Jerry Harris and Andrew Milner. I hadn’t read it before, so I picked up a copy in the gift shop and I’ve been devouring it. As a professional scientist, educator, and book author myself, I’m jealous of what Jerry and Andrew produced – both the text and the abundant full-color illustrations are wonderfully clear, and the book is well-produced and very affordable.

From St. George we hit the road home, and rolled into Claremont just before midnight on Monday. It was a whirlwind tour – 1800 miles, three museums, and two fossil sites in six days – and my brain is still fizzing with all of the things we got to see and do.

One of the many pros of having a professional artist as a friend is that minimal hospitality, like letting him crash on my couch, is sometimes rewarded with original art. Brian was already gone when I got up Tuesday morning, but this was waiting for me on the dining room table. (Want your own? Help Brian make more monsters here.)

I owe plenty of thanks myself: to the Foster and Milner families for their near-maximal hospitality, to Julia McHugh of Dinosaur Journey for assistance in collections, to Diana Azevedo, Jalessa Spor, Jerry Harris, and the rest of the SGDS staff for being such gracious hosts, to Brian for being such a great friend and traveling companion, and most of all to Paige Wiren for finding the apato femur and helping us save it for science. You’re all top-notch human beings and I hope our paths cross again soon.

We all know that apatosaurines have big honkin’ cervical ribs (well, most of us know that). But did they also have unusually large neural spines?

The question occurred to me the other day when I was driving home from work. I was thinking about C10 of CM 3018, the holotype of Apatosaurus louisae, and I thought, “Man, that is a lot of neural spine right there.”

Why was I thinking about C10, particularly? I traced and also stacked Gilmore’s (1936) drawing for my 2002 paper with Kent Sanders, and recycled the trace for my 2007 prosauropod paper, and recycled the stack-o-C10s for my 2013 PeerJ paper with Mike. So for better or worse C10 is my mental shorthand for A. louisae, the same way that their respective C8s seem to capture the essence of Giraffatitan and Sauroposeidon.

I decided that the quick-and-dirty solution was to compare the vertebrae of A. louisae with those of Diplodocus carnegii, the default reference diplodocid, and see how they stacked up. With the cotyles scaled to the same vertical diameters, this is what we get for C9 and C10 of CM 3018 (lighter gray, background, traced from Gilmore 1936) vs CM 84/94 (darker gray, foreground, traced from Hatcher 1901):

The A. louisae verts are a hair taller, proportionally, than those of D. carnegii, but not by much. The difference is trivial compared to the differences in centrum length and cervical rib size.

So where did I get this apparently erroneous impression that Apatosaurus had giant neural spines? Maybe it’s not that the neural spines of apatosaurines in particular are so large, but rather than diplodocids of all types have large neural spines compared to non-diplodocids. Here are the same vertebrae compared for D. carnegii (dark gray, background) and Camarasaurus supremus (black, foreground, traced from Osborn and Mook 1921):

I deliberately picked the longest C9 in the AMNH collection, and the least-distorted C10. The first surprise for me was how well this C. supremus C9 hangs with D. carnegii in terms of proportions. That is one looooong Cam vert. In any other sauropod, it would probably be beautiful. But because it’s Camarasaurus it attained its length in the most lumpen possible way, with the diapophysis way up front, the neural spine apex way at the back, and in the middle just…more vertebra. Like a stretch limo made from a Ford Pinto, or Mike’s horrifying BOBA-horse.

Inevitable and entirely justified Cam-bashing aside, it’s striking how much smaller the whole neural arch-and-spine complex is in C. supremus than in D. carnegii. And remember that D. carnegii is itself a bit smaller than Apatosaurus, spine-wise. Is this maybe a diplodocoid-vs-macronarian thing, at least in the Morrison? Here’s the C10 stack with Brachiosaurus included, represented by BYU 12867 (which I think is probably a C10 based on both centrum proportions and neural spine shape – see Wedel et al. 2000b for details), and with labels added because it’s getting a little nuts:

I like this; it shows a lot. Here are some things to note:

• The diplodocids don’t just have taller neural spines, their pre- and postzygapophyses are also higher than in the macronarians. That’s gotta mean something, right? All else being equal, putting the zygs farther from the intervertebral joints would reduce the flexibility of the neck. Maybe diplodocoids could get away with it because they had more cervicals, or maybe their necks were stiffened for some reason.
• The zygs being set forward of their respective centrum ends in the macronarians really comes through here.
• The Brachiosaurus vert isn’t that different from a stretched (and de-uglified) Cam vert, with a slightly higher neural spine to help support the longer neck. (Maybe this is why Cam inspires such visceral revulsion: it reads as a failed brachiosaur.)
• This emphasizes the outlier status of Apatosaurus in the cervical rib department. It bears repeating: the cervical ribs of Camarasaurus are certainly wide, but they’re not nearly as massive or ventrally expanded as in apatosaurines.

So far, pretty interesting. I’d like to add Barosaurus and Haplocanthosaurus to round out the “big six” Morrison sauropods. I known Haplo has big, tall, almost apatosaurine neural spines (as shown above, with arrows highlighting the epipophyses), but for Baro I’d have to actually do the comparison to see where it falls out.

The idea of bringing in Barosaurus also forces the question, previously glossed over, of how legit it is to compare C10s of all these animals when their cervical counts differed. C. supremus is thought to have had 12 vertebrae in its neck, Brachiosaurus 13 (based on Giraffatitan), A. louisae and D. carnegii 15, and Barosaurus probably 16. It would be more informative to graph neural spine height divided by cotyle diameter along the column for all of these critters, plus Kaatedocus and Galeamopus. But that’s a lot of actual work, and as much fun as it sounds (really, I’d rather be doing that), I have summer teaching to prep for and field gear to wrangle. So I’ll have to revisit this stuff another time.

References

• Gilmore Charles W. 1936. Osteology of Apatosaurus, with special reference to specimens in the Carnegie Museum. Memoirs of the Carnegie Museum 11:175–300 and plates XXI–XXXIV.
• Hatcher, John Bell. 1901. Diplodocus (Marsh): its osteology, taxonomy, and probable habits, with a restoration of the skeleton. Memoirs of the Carnegie Museum 1:1-63.
• Osborn, Henry Fairfield, and Charles C. Mook. 1921. Camarasaurus, Amphicoelias and other sauropods of Cope. Memoirs of the American Museum of Natural History, n.s. 3:247-387, and plates LX-LXXXV.
• Taylor, Michael P., and Mathew J. Wedel. 2013. Why sauropods had long necks; and why giraffes have short necks. PeerJ 1:e36. 41 pages, 11 figures, 3 tables. doi:10.7717/peerj.36
• Wedel, M.J., and Sanders, R.K. 2002. Osteological correlates of cervical musculature in Aves and Sauropoda (Dinosauria: Saurischia), with comments on the cervical ribs of Apatosaurus. PaleoBios 22(3):1-6.
• Wedel, M.J., Cifelli, R.L., and Sanders, R.K. 2000b.  Osteology, paleobiology, and relationships of the sauropod dinosaurSauroposeidon.  Acta Palaeontologica Polonica 45(4): 343-388.