November 21, 2016
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.
A few things about this:
- 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.
- 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.
- 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?
October 27, 2016
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.
September 18, 2016
I have before me the reviews for a submission of mine, and the handling editor has provided an additional stipulation:
Authority and date should be provided for each species-level taxon at first mention. Please ensure that the nominal authority is also included in the reference list.
In other words, the first time I mention Diplodocus, I should say “Diplodocus Marsh 1878″; and I should add the corresponding reference to my bibliography.
What do we think about this?
I used to do this religiously in my early papers, just because it was the done thing. But then I started to think about it. To my mind, it used to make a certain amount of sense 30 years ago. But surely in 2016, if anyone wants to know about the taxonomic history of Diplodocus, they’re going to go straight to Wikipedia?
I’m also not sure what the value is in providing the minimal taxonomic-authority information rather then, say, morphological information. Anyone who wants to know what Diplodocus is would be much better to go to Hatcher 1901, so wouldn’t we serve readers better if we referred to “Diplodocus (Hatcher 1901)”
Now that I come to think of it, I included “Giving the taxonomic authority after first use of each formal name” in my list of
Idiot things that we we do in our papers out of sheer habit three and a half years ago.
Should I just shrug and do this pointless busywork to satisfy the handling editor? Or should I simply refuse to waste my time adding information that will be of no use to anyone?
- Hatcher, Jonathan B. 1901. Diplodocus (Marsh): its osteology, taxonomy and probable habits, with a restoration of the skeleton. Memoirs of the Carnegie Museum 1:1-63 and plates I-XIII.
- Marsh, O. C. 1878. Principal characters of American Jurassic dinosaurs, Part I. American Journal of Science, series 3 16:411-416.
September 16, 2016
Suppose that I and Matt were right in our SVPCA talk this year, and the
“Supersaurus” cervical BYU 9024 really is the C9 of a gigantic Barosaurus. As we noted in our abstract, its total length of 1370 mm is exactly twice that of the C9 in AMNH 6341, which suggests its neck was twice as long over all — not 8.5 m but 17 m.
How horrifying is that?
I realised one good way to picture it is next to the entire mounted skeleton of Giraffatitan at the Museum für Naturkunde Berlin. That skeleton is 13.27 m tall. At 17 m, the giant barosaur neck would be 28% longer than the total height Giraffatitan.
Yes, this looks ridiculous. But it’s what the numbers tell us. Measure the skeleton’s height and the neck length off the image yourself if you don’t believe me.
(Note, too, that the size of the C9 in that big neck is about right, compared with a previous scaled image that Matt prepared, showing the “Supersaurus” vertebra in isolation alongside the Chicago Brachiosaurus.)
September 14, 2016
Long-time SV-POW! readers will remember that three years ago, full of enthusiasm after speaking about Barosaurus at the Edinburgh SVPCA, Matt and I got that talk written up in double-quick time and had it published as a PeerJ Preprint in less than three weeks. Very quickly, the preprint attracted substantive, helpful reviews: three within the first 24 hours, and several more in the next few days.
This was great: it gave us the opportunity to handle those review comments and get the manuscript turned around into an already-reviewed formal journal submission in less then a month from the original talk.
So of course what we did instead was: nothing. For three years.
I can’t excuse that. I can’t even explain it. It’s not as though we’ve spent those three years churning out a torrent of other awesome papers. We’ve both just been … a bit lame.
Anyway, here’s a story that will be hauntingly familiar. A month ago, full of enthusiasm after speaking about Barosaurus at the Liverpool SVPCA, Matt and I found ourselves keen to write up that talk in double-quick time. It’s an exciting tale of new specimens, reinterpretation of an important old specimen, and a neck eight times as long as that 0f a world-record giraffe.
But it would be crazy to write the new Barosaurus paper without first having dealt with the old Barosaurus paper. So now, finally, three years on, we’ve done that. Version 2 of the preprint is now available (Taylor and Wedel 2016), incorporating all the fine suggestions of the people who reviewed the first version — and with a slightly spiffed-up title. What’s more, the new version has also been submitted for formal peer-review. (In retrospect, I can’t think why we didn’t do that when we put the first preprint up.)
A big part of the purpose of this post is to thank Emanuel Tschopp, Mark Robinson, Andy Farke, John Foster and Mickey Mortimer for their reviews back in 2013. I know it’s overdue, but they are at least all acknowledged in the new version of the manuscript.
Now we cross our fingers, and hope that the formally solicited reviews for the new version of the manuscript are as helpful and constructive as the reviews in that first round. Once those reviews are in, we should be able to move quickly and painlessly to a formally published version of this paper. (I know, I know — I shouldn’t offer such a hostage to fortune.)
Meanwhile, I will finally be working on handling the reviews of this other PeerJ submission, which I received back in October last year. Yes, I have been lax; but I am back in the saddle now.
- Taylor, Michael P., and Mathew J. Wedel. 2016. The neck of Barosaurus: longer, wider and weirder than those of Diplodocus and other diplodocines. PeerJ PrePrints 1:e67v2 doi:10.7287/peerj.preprints.67v2
September 8, 2016
If you keep an eye on the wacky world of zoological nomenclature, you’ll know that earlier this year Emanuel Tschopp and Octávio Mateus published a petition to the International Commission on Zoological Nomemclature, asking them to establish Diplodocus carnegii, represented by the ubiquitous and nearly complete skeleton CM 84, as the type species of Diplodocus.
That is because Marsh’s (1878) type species, YPM 1920, is a pair of non-diagnostic mid-caudals which no-one has paid any attention to since 1901:
I have now submitted a formal comment to the ICZN in support of the petition, in which I argue:
In its use as the definitive exemplar of the genus Diplodocus, as the foundation for numerous palaeobiological studies of the genus, and as the specifier for numerous important clades, the species D. carnegii is already effectively functioning as the type species of Diplodocus. Therefore the petition of Tschopp and Mateus (2016) requests only that the commission recognises de jure what is already the case de facto.
Anyone else who has strong feelings either in favour of or against the establishment of D. carnegii as a replacement type species for Diplodocus is welcome to submit their own comment to the ICZN. (I know of at least one person who has submitted a comment opposing the petition.)
The procedure is straightforward: just write your comment and email it to the Commision at firstname.lastname@example.org. (But it’s best also to copy your email to email@example.com, as that seems to be where the ICZN is operating out of now: it took the NHM address four days to reply to my initial inquiry, but the Singaporean address responds quickly.)
- Marsh, O.C. 1878. Principal characters of American Jurassic dinosaurs, Part I. American Journal of Science (series 3) 16:411–416.
- Tschopp, Emanuel, and Octávio Mateus. 2016. Case 3700: Diplodocus Marsh, 1878 (Dinosauria, Sauropoda): proposed designation of D. carnegii Hatcher, 1901 as the type species. Bulletin of Zoological Nomenclature 73(1):17-24.
UPDATE 19 May 2016
I belatedly realized that I caused some confusion in the original version of this post. This will hopefully sort things out:
The ventrolateral processes (1) are nothing new. As Ken Carpenter pointed out in a comment, Hatcher noted them back in 1901 in his monograph on Diplodocus carnegii. These are the features I describe below as being, “huge in Barosaurus, big in Diplodocus, small in Apatosaurus, and nonexistent in Haplocanthosaurus, Camarasaurus, and the brachiosaurids, at least from what I’ve seen.” To clarify: occasionally in camarasaurs and frequently in brachiosaurs you can trace a ridge along the ventrolateral margin of the centrum from the parapophysis to the cotyle. But these ridges are basically just the ‘corners’ of the centrum, leftover by the lateral and ventral waisting of the centrum – they do not project beyond the margin of the cotyle. In contrast, what I’ve been calling the ventrolateral flanges in diplodocids do project beyond the margins of the cotyle – they are additive structures, not just architectural leftovers. They also don’t vary much, other than to be more pronounced in more posterior cervicals.
The irregular ventral ridges (2) are a totally different thing. They’re on or near the sagittal midline of the centrum, usually restricted to the anteroposterior middle of the ventral centrum (so, about halfway between the condyle and the cotyle), and as my preferred term implies, highly variable among individuals and even among vertebrae in a series.
Hope that helps! (Original post starts below.)
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Back in 2005 I visited BYU while I was working on my dissertation. Back then I noted ventral ridges in a few diplodocine cervical vertebrae. (I hesitate to call such flimsy things ‘keels’.)
Up above is BYU 16918, a mid-to-posterior cervical vertebra of Diplodocus from the famous Dry Mesa Quarry. Here it is again in posterior view:
The things I have labeled VLF here are ventrolateral flanges, which are huge in Barosaurus, big in Diplodocus, small in Apatosaurus, and nonexistent in Haplocanthosaurus, Camarasaurus, and the brachiosaurids, at least from what I’ve seen. See this post for details. I know that the left VLF here looks like a second ridge, but the cotyle is broken off in such a way that we’re seeing the fossa just dorsal to the VLF margin. The ridge itself is skewed to the right, which could be natural or a result of taphonomy – as you can see from the photo at the top of the post, this vert has seen better days.
Here’s another Dry Mesa vert, BYU 11617, this time an anterior cervical of Barosaurus and in left lateral view:
Again in right lateral view – on this side you can see the fossa in the VLF more clearly:
And here’s the ventral view showing the ridge:
I noted these things in my notebook back when, filed them under, “Huh. How about that?” and went on with life.
Then last week Mike and I were at the North American Museum of Ancient Life in Lehi, Utah, and we saw this super-nice Barosaurus cervical on display in the prep lab (left ventro-lateral view). Check out the monster ventrolateral flanges, and the ridges between them at about mid-centrum.
Here’s another view, a more square-on ventral this time:
We owe a big thank you to Rick Hunter, who let us into the prep lab at the North American Museum of Ancient Life to see the Barosaurus material up close.
So what’s the deal with these ridges? I assume that they’re caused by pneumatic diverticula remodeling the ventral surface of the centrum. We know that such diverticula were down there because there are actual foramina on the ventral centrum in Supersaurus, many apatosaurines (Lovelace et al., 2008), many brachiosaurids, and probably loads of other things that haven’t been checked. Oddly enough, I’ve never seen the ridges in any of those other taxa. It seems that you get foramina or ridges, but not both. I have no idea what’s up with that – to paraphrase Neal Stephenson, Barosaurus cervicals are confections of air and marketing, and you’d think that if any sauropod would have straight-up foramina down there, it would be Barosaurus. But Barosaurus gets ridges and clunky old Apatosaurus gets foramina (sometimes, not all the time).
It’s a sick world, I tell you.
- 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.