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.


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.


  • 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.

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.


  • 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.

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?


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.


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.

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:

2016-05-11 15.42.40

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.


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

Wedel 2005 Morrison sauropod cervicals 1 - Diplodocus

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.

Wedel 2005 Morrison sauropod cervicals 2 - Barosaurus and centra shapes

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.

Wedel 2005 Morrison sauropod cervicals 3 - Brachiosaurus and Apatosaurus

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.

Wedel 2005 Morrison sauropod cervicals 4 - Camarasaurus and Haplocanthosaurus

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…