Did apatosaurs have unusually large neural spines, too?
June 22, 2018
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.
Listen to Mark Hallett and Matt Wedel on the I Know Dino podcast
December 29, 2017
Hey sports fans, as the year winds down I bring you another podcast appearance. This time out I’m rolling with Mark Hallett, and we’re talking about sauropods through the lens of our still-plausibly-somewhat-newish book, The Sauropod Dinosaurs: Life in the Age of Giants, on the I Know Dino podcast. Many thanks to Sabrina and Garret for having us on the show. While you’re on that page, check out the nice preview of Mark’s 2018 dinosaur calendar, which is available at Pomegranate and Amazon.
The photo shows the Diplodocus carnegii cast mounted in the natural history museum in Vienna, one of Andrew Carnegie’s gifts to the world. A happy seasonal metaphor, sez me. Hope your new year is equally happy!
The “Growth series of one” poster is published
September 22, 2017
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.
Walking with sauropods at Copper Ridge (video)
December 14, 2016
In the summer of 2015, Brian Engh and I stopped at the Copper Ridge dinosaur trackway on our way back from the field. The Copper Ridge site is 23 miles north of Moab, off US Highway 191. You can find a map, directions, and some basic information about the site in this brochure. The BLM has done a great job of making this and other Moab-area dinosaur trackways accessible to the public, with well-tended trails and nice interpretive signage. Brian has gotten to do the art for interp signs at several sites now, including Copper Ridge, and he put together this video to explain a bit about the site, what we know about the trackmaker, and the lines of evidence he used in making his life restoration. I’m in there, too, yammering a bit about which sauropod might have been responsible. We weren’t sure what, if anything, we would end up doing with the footage at the time, so I’m basically thinking out loud. But that’s mostly what I do here anyway, so I reckon you’ll live.
Stay tuned (to Brian’s paleoart channel) for Part 2, which will be about the Copper Ridge theropod trackway. And the next time you’re in the Moab area, go see some dinosaur tracks. This is our heritage, and it’s cool.
Should we still give taxonomic authorities?
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.
Marsh (1878: plate VIII in part). The only illustration of Diplodocus material in the paper that named the genus.
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?
References
- 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.
My comment in support of the Diplodocus carnegii ICZN petition
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:
Tschopp and Mateus (2016: fig. 1). More anterior of the only two reasonably complete caudal vertebrae of the type specimen of Diplodocus longus (YPM 1920) in dorsal (A), anterior (B), left (C), posterior (D), right (E), and ventral (F) views. The neural spine is lost. The estimated position within the caudal column is caudal vertebra 17â24. Note the transverse ridge between the prezygapophyses shared with AMNH 223 (1).
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 iczn@nhm.ac.uk. (But it’s best also to copy your email to iczn@nus.edu.sg, 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.)
References
- 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.
Reference
- 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.
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…
SV-POW! on the road – Sauropocalypse 2016
April 30, 2016
Mounted Diplodocus at the Utah Field House of Natural History State Park Museum in Vernal.
I love Utah. I love how much of the state is given over to exposed Mesozoic rocks. I love driving through Utah, which has a strong baseline of beautiful scenery that is frequently punctuated by the absolutely mind-blowing (Arches, Bryce Canyon, Zion, Monument Valley…). I love doing fieldwork there, and I love the museums, of which there are many. It is not going too far to say that much of what I learned firsthand about sauropod morphology, I learned in Utah (the Carnegie Museum runs a close second on the dragging-Matt-out-of-ignorance scale).
Cast of the juvenile Camarasaurus CM 11338 at Dinosaur National Monument.
There is no easy way to say this so I’m just going to get it over with: Mike has never been to Utah.
I know, right?
But we’re going to fix that. Mike’s flying into Salt Lake City this Wednesday, May 4, and I’m driving up from SoCal to meet him. After that we’re going to spend the next 10 days driving around Utah and western Colorado hitting museums and dinosaur sites. We’re calling it the Sauropocalypse.
Mounted Barosaurus at the Natural History Museum of Utah in Salt Lake City.
Why am I telling you this, other than to inspire crippling jealousy?
First, Mike and I are giving a pair of public talks next Friday evening, May 6, at the USU-Eastern Prehistoric Museum in Price. The talks start at 7:00 and will probably run until 8:00 or shortly after, and there will be a reception with snacks afterward. Mike’s talk will be, “Why giraffes have such short necks”, and my own will be, “Why elephants are so small”.
Second, occasionally people leave comments to the effect of, “Hey, if you’re ever passing through X, give me a shout.” I haven’t kept track of all of those, so this is me doing the same thing in reverse. Here’s our itinerary as of right now:
May 4, Weds: MPT flies in. MJW drives up from Cali. Stay in SLC/Provo area.
May 5, Thurs: recon BYU collections in Provo. Stay in SLC/Provo area.
May 6, Fri: drive to Price, visit USU-Eastern Prehistoric Museum, give evening talks. Stay in Price.
May 7, Sat: drive to Vernal, visit DNM. Stay in Vernal.
May 8, Sun: visit Utah Field House, revisit DNM if needed, drive to Fruita.
May 9, Mon: visit Rabbit Valley camarasaur in AM, visit Dinosaur Journey museum in PM. Go on to Moab.
May 10, Tues: drive back to Provo, visit BYU collections.
May 11, Weds: BYU collections.
May 12, Thurs: drive to SLC to visit UMNH collections, stay for Utah Friends of Paleontology meeting that evening.
May 13, Fri: BYU collections.
May 14, Sat: visit North American Museum of Ancient Life. MPT flies home. MJW starts drive home.
We’re planning lots of time at BYU because we’ll need it, the quantity and quality of sauropod material they have there is ridiculous. As for the rest, some of those details may change on the fly but that’s the basic plan. Maybe we’ll see you out there.
Sauropod Vertebra Picture of the Week 