Apatosaurus cervical ribs and the tyranny of 2D images
December 8, 2022
Some quick backstory: lots of sauropods have long, overlapping cervical ribs, like the ones shown here in Sauroposeidon (diagram from this old post):
These long cervical ribs are ossified tendons of ventral neck muscles, presumably longus colli ventralis. We know they’re ossified tendons because of their bone histology (Klein et al. 2012), and we suspect that they’re longus colli ventralis because those tendons look the same in birds, just less ossified, as in this rhea (same specimens as these even older posts: 1, 2):
Diplodocoids have apomorphically short cervical ribs, which never extend very far past the end of their respective centra and sometimes don’t overlap at all. Still, we assume the long ventral neck muscles were there, just without long ossified tendons. Which brings me to Apatosaurus, which has cervical ribs that are anteroposteriorly short but famously massive, extending very below and/or to the sides of the cervical centra — for a truly breathtaking example see this post. Here are C3 through C7 in CM 3018, the holotype of Apatosaurus lousiae (Gilmore 1936: plate 24):
At least for me, it’s hard to resist the temptation to mentally scoot those vertebrae together into articulation, and imagine that the very swoopy-looking and maybe even down-turned cervical ribs allowed the ventral tendon bundles to wrap around the bottom of each cervical rib protuberance, something like this:
But it’s just not so, because like all 2D images, Gilmore’s plate distorts 3D reality. If you get to see the mounted skeleton in person, it’s clear that the cervical ribs are all more or less in line, and none of them are pointed at the big protuberances, which stick way out ventrolaterally.
Here I’ve drawn in the likely trajectories of the longus colli ventralis tendons. My little red pathways don’t precisely match the cervical ribs as mounted, but there’s a lot of distortion and restoration going on. For example, comparing with Gilmore’s plate we can see that the cervical ribs of C5, which point downward compared to all the others, only do that because someone forced them to — the whole anterior portion of the rib, where the shaft would actually join to the capitulum and tuberculum, is reconstructed. Even if I’m a little off, it’s clear that the cervical ribs shafts point backward, they’re all more or less in two parallel lines, and none of them point down and out toward the ventrolateral processes. The photo contains a mountain of useful morphological information that you’d never get from the lateral views.
My takeaways from all this:
- If a person has only seen 2D images of a specimen, and especially if those 2D images have only been orthogonal views with no obliques, their little island of knowledge is surrounded by at least a sizeable lake of ignorance, if not a small ocean.
- That doesn’t mean that seeing specimens in person is the only antidote — 3D models and 3D prints are extremely useful, and for specimens that are difficult to manipulate because of their size or fragility, they may be more useful than seeing or handling the specimen, at least for some questions.
- For Apatosaurus specifically, those ventrolateral processes cry out for explanation. They’re fairly solid knobs of bone that stick way out past the ossified tendons of the ventral-most neck muscles. That’s a super-weird — and super-expensive — place to invest a bunch of bone if you’re not using it for something fairly important, especially in a lineage that had just spent the last 80-100 million years making their necks as light as possible.
- Pursuant to that last point, we’re now in — ugh-ouch-shame — our 8th year of BrontoSMASH!!, with still just the one conference presentation to show for it (Taylor et al. 2015). Prolly time we got moving on that again.
References
- Klein, N., Christian, A., & Sander, P. M. (2012). Histology shows that elongated neck ribs in sauropod dinosaurs are ossified tendons. Biology letters, 8(6), 1032-1035.
- Taylor, M.P., Wedel, M.J., Naish, D., and Engh, B. 2015. Were the necks of Apatosaurus and Brontosaurus adapted for combat? 63rd Symposium on Vertebrate Palaeontology and Comparative Anatomy, Meeting Proceedings, p. 71, and PeerJ PrePrints 3:e1347v1. https://doi.org/10.7287/peerj.preprints.1347v1
Here’s that ventral-view apatosaur cervical anaglyph you ordered
December 4, 2022
Just to wash our mouths out after all the theropod-related unpleasantness yesterday:
What we’re seeing here, in glorious 3D, is the 7th cervical vertebrae of BYU 1252-18531. This is an apatosaurine at the Brigham Young University Museum of Paleontology which the museum has catalogued as “Apatosaurus excelsus” (i.e. Brontosaurus excelsus), and which Tschopp et al. (2015) tentatively referred to Brontosaurus parvus, but which I suspect is most likely good old Apatosaurus louisae.
It’s in the rarely seen ventral view, which really emphasizes the ludicrously over-engineered cervical ribs. Get your 3D glasses on and marvel at how they come lunging out of the screen at you, like giant insects in a 1950s B-movie.
So beautiful.
I’ve been in contact recently with Matt Lamanna, Associate Curator in the Section of Vertebrate Paleontology at the Carnegie Museum of Natural History — which is obviously the best job in the world. Among a batch of photos that he sent me recently, I seized on this gem:
Tyrannosaurus rex, Diplodocus carnegii, Apatosaurus louisae and multiple mostly juvenile individuals of Homo sapiens. Photograph taken between 1941 and 1965. Courtesy of Carnegie Museum of Natural History.
There’s so much to appreciate in this picture: the hunchbacked, tail-dragging Tyrannosaurus; the camarasaur-style skull on the Apatosaurus; the hard-to-pin-down archaic air of Diplodocus.
But the thing I love about it is the 1950s kids. (Or, to be fair, maybe the 1940s kids or early 1960s kids, but you get the point.) They way they’ve all been asked to look up at the tyrannosaur skull, and are obediently doing it. How earnest they all appear. How they’re all dressed as tiny adults. How self-consciously some of them have posed themselves — the thoughtful kid one in from the left, his foot up on the plinth and his chin resting on his hand; the cool kid to his right, arms crossed, interested but careful not to seem too impressed.
Where are these kids now? Assuming it was taken in 1953, the midpoint of the possible range, and assuming they’re about 12 years old in this photo, they were born around 1941, which would make them 81 now. Statistically, somewhere around half of them are still alive. I wonder how many of them remember this day, and the strange blend of awe, fascination, and self-consciousness.
This is a time-capsule, friends. Enjoy it.
Apatosaurus louisae from below
May 3, 2022
We’ve shown you the Apatosaurus louisae holotype mounted skeleton CM 3018 several times: shot from the hip, posing with another massive vertebrate, photographed from above, and more. Today we bring you a world first: Apatosaurus from below. Scroll and enjoy!
Obviously there’s a lot of perspective distortion here. You have to imagine yourself lying underneath the skeleton and looking up — as I was, when I took the short video that was converted into this image.
Many thanks to special-effects wizard Jarrod Davis for stitching the video into the glorious image you see here.
The most obvious effect of the perspective distortion is that the neck and tail both look tiny: we are effectively looking along them, the neck in posteroventral view and the tail in anteroventral. The ribs are also flared in this perspective, making Apato look even broader than it is in real life. Which is pretty broad. One odd effect of this is that this makes the scapulae look as though they are sitting on top of the ribcage rather than appressed to its sides.
Apatosaurus louisae: shooting from the hip
April 30, 2022
Here at SV-POW! Towers, we like to show you iconic mounted skeletons from unusual perspectives. Here’s one:
Apatosaurus louisae holotype CM 3018, mounted skeleton in the public gallery of the Carnegie Museum of Natural History: head, neck, torso and hip in right posterolateral view. Photograph by Matt Wedel, 12th March 2019 (my birthday!)
Oh, man, I love that museum. And I love that specimen. And I love the one that’s standing next to it (Diplodocus CM 82, natch.) I’ve got to find a way to get myself back out there.
That’s all: just enjoy.
The Wiren Apatosaurus femur is on exhibit in the Moab Museum
October 19, 2021
The last time we saw the sauropod femur that Paige Wiren discovered sticking out of a riverbank, it had been moved into the prep lab at the Moab Museum, with the idea that it would eventually go on exhibit as a touch specimen for the public to enjoy and be inspired by. That has come to pass.
I was in Moab last month with Drs. Jessie Atterholt and Thierra Nalley and we stopped in the Moab Museum to digitize some vertebrae from SUSA 515, an unusual specimen of Camarasaurus that I’ve blogged about before, and will definitely blog about again. While we were there, we got to see and touch the Wiren femur. The museum folks told us that femur has been the first dinosaur bone that a lot of schoolkids and tourists have seen up close, or gotten to touch. As a former dinosaur-obsessed kid who never stopped being excited about touching real dinosaur bones–and as one of the lucky folks that got to rescue this particular fossil from erosion or poaching–that pleases me deeply.
So, obviously, you should go see this thing. And the rest of the museum–as you can see from the photos above, the whole place has been renovated, and there are lots of interesting fossils from central and eastern Utah on display, not to mention loads of historical artifacts, all nicely presented in a clean, open, well-lit space that invites exploration. Go have fun!
Happy Valentine’s Day from Apatosaurinae
February 15, 2021
This is RAM 1619, a proximal caudal vertebra of an apatosaurine, in posterior view. It’s one of just a handful of sauropod specimens at the Raymond M. Alf Museum of Paleontology. It’s a donated specimen, which came with very little documentation. It was originally catalogued only to a very gross taxonomic level, but I had a crack at it on a collections visit in 2018, when I took these photos. I told Andy Farke and the other Alf folks right away, I just never got around to blogging about it until now.
Why do I think it’s an apatosaurine? A few reasons:
- it’s slightly procoelous, which is pretty common for diplodocids, whereas caudals of Haplocanthosaurus, Camarasaurus, and Brachiosaurus are all either amphicoelous or amphiplatyan;
- it has big pneumatic fossae above the transverse processes, unlike Haplo, Cam, and Brachio, but it lacks big pneumatic fossae below the transverse processes, unlike Diplodocus and Barosaurus.
- and finally the clincher: the centrum is taller than wide, and broader dorsally than ventrally.
In the literature this centrum shape is described as ‘heart-shaped’ (e.g., Tschopp et al. 2015), and sometimes there is midline dorsal depression that really sells it. That feature isn’t present in this vert, but overall it’s still much closer to a heart-shape than the caudals of any non-apatosaurine in the Morrison. Hence the literal 11th-hour Valentine’s Day post (and yes, this will go up with a Feb. 15 date because SV-POW! runs on England time, but it’s still the 14th here in SoCal, at least for another minute or two).
RAM 1619 in postero-dorsal view.
Back to the pneumaticity. Occasionally an apatosaurine shows up with big lateral fossae ventral to the transverse processes–the mounted one at the Field Museum is a good example (see this post). And the big Oklahoma apatosaurine breaks the rules by having very pneumatic caudals–more on that in the future. But at least in the very proximal caudals of non-gigantic apatosaurines, it’s more common for there to be pneumatic fossae above the transverse processes, near the base of the neural arch. You can see that in caudal 3 of UWGM 15556/CM 563, a specimen of Brontosaurus parvus:
I don’t think I’d figured out this difference between above-the-transverse-process (supracostal, perhaps) and below-the-transverse-process (infracostal, let’s say) pneumatic fossae when Mike and I published our caudal pneumaticity paper back in 2013. I didn’t start thinking seriously about the dorsal vs ventral distribution of pneumatic features until sometime later (see this post). And I need to go check my notes and photos before I’ll feel comfortable calling supracostal fossae the apatosaurine norm. But I am certain that Diplodocus and Barosaurus have big pneumatic foramina on the lateral faces of their proximal caudals (see this post, for example), Haplocanthosaurus and brachiosaurids have infracostal fossae when they have any fossae at all in proximal caudals (distally the fossae edge up to the base of the neural arch in Giraffatitan), and to date there are no well-documented cases of caudal pneumaticity in Camarasaurus (if that seems like a hedge, sit tight and W4TP).
RAM 1619 has asymmetric pneumatic fossae, which is pretty cool, and also pretty common, and we think we have a hypothesis to explain that now–see Mike’s and my new paper in Qeios.
And if I’m going to make my midnight deadline, even on Pacific Time, I’d best sign off. More cool stuff inbound real soon.
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.
- Taylor, Michael P., and Mathew J. Wedel. 2021. Why is vertebral pneumaticity in sauropod dinosaurs so variable? Qeios 1G6J3Q. doi:10.32388/1G6J3Q
- (2015) A specimen-level phylogenetic analysis and taxonomic revision of Diplodocidae (Dinosauria, Sauropoda) PeerJ 3:e857 https://dx.doi.org/10.7717/peerj.857
- Wedel, Mathew J., and Michael P. Taylor. 2013. Caudal pneumaticity and pneumatic hiatuses in the sauropod dinosaurs Giraffatitan and Apatosaurus.PLOS ONE 8(10):e78213. 14 pages. doi:10.1371/journal.pone.0078213 [PDF]
What a cervical vertebra of an 800m sauropod would look like
February 1, 2021
Gilmore (1936:243) says of the mounted skeleton of Apatosaurus louisae CM 3018 in the Carnegie Museum that “with the skull in position the specimen has a total length between perpendiculars of about 71 feet and six inches. If the missing eighteen terminal caudal vertebrae were added to the tip of the tail, in order to make it conform to known evidence, the skeleton will reach an estimated length of 76 feet, 6 inches.” That’s 23.3 meters.
But what if it was 800 meters long instead? That would be 34.3 times as big in linear dimension (and so would mass 34.3^3 = 40387 time as much, perhaps a million tonnes — but that’s not my point).
What would a cervical vertebra of an 800m sauropod look like?
Gilmore (1936:196) gives the centrum length of CM 3018’s C10 as 530 mm. In our 34.3 times as long Apatosaurus, it would be 18.17 meters long. So here is what that would look like compared with two London Routemaster buses (each 8.38 meters long).
Cervical vertebra 10 of a hypothetical 800 meter long Apatosaurus louisae, with London Routemaster buses for scale. Vertebra image from Gilmore 1936:plate XXIV; bus image by Graham Todman, from Illustrations for t-shirts.
What is the research significance of this? None at all, of course. Still I think further study is warranted. Some look at sauropods that once were, and ask “why?”; but I go further; I look at sauropods that never were, and ask “why not?”
Hey, look, sauropod vertebrae!
January 15, 2021
It’s been a minute, hasn’t it?
Up top, C10 and C11 of Diplodocus carnegii CM 84, from Hatcher (1901). Below, C9 and C10 of Apatosaurus louisae CM 3018, from Gilmore (1936). The Diplodocus verts are in right lateral view but reversed for ease of comparison, and the Apatosaurus verts are in left lateral view. Both sets scaled to the same cumulative centrum length. Just in case you forgot that apatosaurines are redonkulous.
References
- 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.
- Gilmore, Charles Whitney. 1936. Osteology of Apatosaurus, with special reference to specimens in the Carnegie Museum. Memoirs of the Carnegie Museum 11:175–300.
Tutorial 38: little projects as footsteps toward understanding
January 11, 2021
This is a very belated follow-up to “Tutorial 12: How to find problems to work on“, and it’s about how to turn Step 2, “Learn lots of stuff”, into concrete progress. I’m putting it here, now, because I frequently get asked by students about how to get started in research, and I’ve been sending them the same advice for a while. As with Tutorial 25, from now on I can direct the curious to this post, and spend more time talking with them about what they’re interested in, and less time yakking about nuts and bolts. But I hope the rest of you find this useful, too.
Assuming, per Tutorial 12, that you’ve picked something to investigate–or maybe you’re trying to pick among things to investigate–what next? You need a tractable way to get started, to organize the things you’re learning, and to create a little structure for yourself. My recommendation: do a little project, with the emphasis on little. Anyone can do this, in any area of human activity. Maybe your project will be creating a sculpture, shooting and editing a video, learning–or creating–a piece of music, or fixing a lawn mower engine. My central interest is how much we still have to discover about the natural world, so from here on I’m going to be writing as a researcher addressing other researchers, or aspiring researchers.
Arteries of the anterior leg, from Gray’s Anatomy (1918: fig. 553). Freely available courtesy of Bartleby.com.
I’ll start with a couple of examples, both from my own not-too-distant history. A few years ago I got to help some of my colleagues from the College of Podiatric Medicine with a research project on the perforating branch of the peroneal artery (Penera et al. 2014). I knew that vessel from textbooks and atlases and from having dissected a few out, but I had never read any of the primary (journal) literature on it. As the designated anatomist on the project, I needed to write up the anatomical background. So I hit the journals, tracked down what looked like the most useful papers, and wrote a little 2-page summary. We didn’t use all of it in the paper, and we didn’t use it all in one piece. Some sentences went into the Introduction, others into the Discussion, and still others got dropped entirely or cut way down. But it was still a tremendously useful exercise, and in cases like this, it’s really nice to have more written down than you actually need. Here’s that little writeup, in case you want to see what it looks like:
Wedel 2013 anatomy of the perforating branch of the peroneal artery
Pigeon spinal cord cross-section, from Necker (2006: fig. 4).
More recently, when I started working with Jessie Atterholt on weird neural canal stuff in dinosaurs, I realized that I needed to know more about glycogen bodies in birds, and about bird spinal cords generally. I expected that to be quick and easy: read a couple of papers, jot down the important bits, boom, done. Then I learned about lumbosacral canals, lobes of Lachi, the ‘ventral eminences’ of the spinal cord in ostriches, and more, a whole gnarly mess of complex anatomy that was completely new to me. I spent about a week just grokking all the weird crap that birds have going on in their neural canals, and realized that I needed to crystallize my understanding while I had the whole structure in my head. Otherwise I’d come back in a few months and have to learn it all over again. Because it was inherently visual material, this time I made a slide deck rather than a block of text, something I could use to get my coauthors up to speed on all this weirdness, as well as a reminder for my future self. Here’s that original slide deck:
Wedel 2018 Avian lumbosacral spinal cord specializations
If you’re already active in research, you may be thinking, “Yeah, duh, of course you write stuff down as you get a handle on it. That’s just learning.” And I agree. But although this may seem basic, it isn’t necessarily obvious to people who are just starting out. And even to the established, it may not be obvious that doing little projects like this is a good model for making progress generally. Each one is a piton driven into the mountainside that I’m trying to climb: useful for me, and assuming I get them out into the world, useful for anyone I’d like to come with me (which, for an educator and a scientist, means everyone).
A view down the top of the vertebral column in the mounted skeleton of Apatosaurus louisae, CM 3018, showing the trough between the bifurcated neural spines.
If you’re not active in research, the idea of writing little term papers may sound like purgatory. But writing about something that you love, that fascinates you, is a very different proposition from writing about dead royalty or symbolism because you have to for a class.* I do these little projects for myself, to satisfy my curiosity, and it doesn’t feel like work. More like advanced play. When I’m really in the thick of learning a new thing–and not, say, hesitating on the edge before I plunge in–I am so happy that I tend to literally bounce around like a little kid, and the only thing that keeps me sitting still is the lure of learning the next thing. That I earn career beans for doing this still seems somewhat miraculous, like getting paid to eat ice cream.
* YMMV, history buffs and humanities folks. If dead royalty and symbolism rock your world but arteries and vertebrae leave you cold, follow your star, and may a thousand gardens grow.
Doing little projects is such a convenient and powerful way to make concrete progress that it has become my dominant mode. As with the piece that I wrote about the perforating branch of the peroneal artery, the products rarely get used wholesale in whatever conference presentation or research paper I end up putting together, but they’re never completely useless. First, there is the benefit to my understanding that I get from assembling them. Second, they’re useful for introducing other people to the sometimes-obscure stuff I work on, and nothing makes you really grapple with a problem like having to explain it to others. And third, these little writeups and slideshows become the Lego bricks from which I assemble future talks and papers. The bird neural canal slide deck became a decent chunk of our presentation on the Snowmass Haplocanthosaurus at the 1st Palaeontological Virtual Congress (Wedel et al. 2018)–and it’s about to become something even better. (Four months later: it did!)
The operative word at the start of the last paragraph is ‘concrete’. I don’t think this was always the case, but now that I’m in my mid-40s ‘what I know’ is basically equivalent to ‘what I remember’, which is basically equivalent to ‘what I’ve written down’. (And sometimes not even then–Mike and I both run across old posts here on SV-POW! that we’ve forgotten all about, which is a bit scary, given how often we put novel observations and ideas into blog posts.) Anyway, this is why I like the expression ‘crystallize my understanding’: the towers of comprehension that I build in my head are sand castles, and if I don’t find a way to freeze them in place, they will be washed away by time and my increasingly unreliable cerebral machinery.
Really nice Stegosaurus plate on display at Dinosaur National Monument.
Also, if I divide my life into the things I could do and the things I have done, only the things in the latter category are useful. So if you are wondering if it’s worthwhile to write a page to your future self about valves in the cerebral arteries of rats, or all of the dinosaurs from islands smaller than Great Britain, or whatever strange thing has captured your attention, I say yes, go for it. Don’t worry about finding something novel to say; at the early stages you’re just trying to educate yourself (also, talks and papers need intro and background material, so you can still get credit for your efforts). I’ll bet that if you set yourself the goal of creating a few of these–say, one per year, or one per semester–you’ll find ways to leverage them once you’ve created them. If all else fails, start a blog. That might sound flip, but I don’t mean for it to. I got my gig writing for Sky & Telescope because I’d been posting little observing projects for the readers of my stargazing blog.
A final benefit of doing these little projects: they’re fast and cheap, like NASA’s Discovery missions. So they’re a good way to dip your toes into a new area before you commit to something more involved. The more things you try, the more chances you have to discover whatever it is that’s going to make you feel buoyantly happy.
You may have noticed that all of my examples in this post involved library research. That’s because I’m particularly interested in using little projects to get started in new lines of inquiry, and whenever you are starting out in a new area, you have to learn where the cutting edge is before you can move it forward (Tutorial 12 again). Also, as a practical consideration, most of us are stuck with library research right now because of the pandemic. Obviously this library research is no substitute for time in the lab or the field, but even cutters and diggers need to do their homework, and these little projects are the best way that I’ve found of doing that.
P.S. If you are a student, read this and do likewise. And, heck, everyone else who writes should do that, too. It is by far the advice I give most often as a journal editor and student advisor.
P.P.S. As long as you’re reading Paul Graham, read this piece, too–this whole post was inspired by the bit near the end about doing projects.
References
- Gray, Henry. 1918. Anatomy of the Human Body. Lea & Febiger, Philadelphia. Bartleby.com, 2000. www.bartleby.com/107/.
- Penera, K., Manji, K., Wedel, M., Shofler, D., and Labovitz, J. 2014. Ankle syndesmotic fixation using two screws: Risk of injury to the Perforating Branch of the Peroneal Artery. The Journal of Foot and Ankle Surgery 53(5):534-8. DOI: 10.1053/j.jfas.2014.04.006
- Wedel, M.J., Atterholt, J., Macalino, J., Nalley, T., Wisser, G., and Yasmer, J. 2018. Reconstructing an unusual specimen of Haplocanthosaurus using a blend of physical and digital techniques. 1st Palaeontological Virtual Congress / PeerJ Preprints 6:e27431v1
Sauropod Vertebra Picture of the Week 