September 22, 2014
We have good descriptions of the proximal parts of the cervical ribs for lots of sauropods. We also have histological cross-sections of a few, mostly thanks to the work of Nicole Klein and colleagues (Klein et al. 2012, Preuschoft and Klein 2013), although histological cross-sections of ribs were also figured as long ago as 1999, by Dalla Vecchia (1999: figs. 29 and 30), and as recently as this month, by Lacovara et al. (2014: supplementary figure 4).
What we have very, very few of is series of cross-sections that show how the cr0ss-section of a cervical rib changes along its length. There may be more out there (and if I have forgotten any, please remind me!), but at the moment I can only think of three such figures: two in Janensch (1950: figs. 83 and 85), both on Giraffatitan, and one in Klein et al. (2012: fig. 1), with cross-sections from Mamenchisaurus, Giraffatitan, and Diplodocus (shown at the top of the post).
Rarer still are images that show cross-sections of overlapped cervical ribs, stacked in situ. You could use the information in Janensch (1950: figs. 83 and 85) to generate the stacked cross-sections, but you wouldn’t know the spacing between the ribs as they were in the ground. I think the image just above, of the cervical rib bundles in the Sauroposeidon holotype, OMNH 53062, may be the first of its kind–again, if you know of any others, please let me know. I took the notes for this figure back in 2004, sitting down with the holotype and some digital calipers to make sure I could scale everything correctly, I just hadn’t ever put it into a presentable form until now. The first C6 section (blue V-shape) is from right at the root where the capitulum and tuberculum meet and the posterior shaft of the rib begins.
It is by now well-understood that the long cervical ribs of sauropods and other dinosaurs are ossified tendons of the long hypaxial neck muscles, specifically the longus colli ventralis and flexor colli lateralis. We argued this back in 200o on comparative anatomical grounds (Wedel et al. 2000b: pp. 378-379), and it has now been demonstrated histologically (Klein et al. 2012, Lacovara et al. 2014). The system of stacked tendons is also found in most birds. Here’s the bundle of stacked tendons in a rhea neck, only slightly fanned out:
And the same neck, with both the epaxial and hypaxial muscles more fully separated:
What I’d really like is an MRI of a rhea or ostrich neck, showing the stacked tendons and their associated belts of muscle, to compare with the stacked cervical ribs of Sauroposeidon and other sauropods. Anyone know of any?
Incidentally, I think the cervical ribs and cervical rib bundles of sauropods are one line of evidence for sauropod necks having been rather slenderly-muscled. The long, multi-segment muscles like the longus colli ventralis are the outermost components of the muscular envelope that surrounds the vertebrae, as you can see in the rhea dissection photos. In sauropod specimens with articulated cervical ribs, the ribs do not deviate from one another or fan out. Rather, they lie in vertically stacked bundles that run from one capitulum-tuberculum intersection to the next. So the depth of that intersection–the “root” of the cervical rib of any given vertebra–plus the thickness of the ribs stacked underneath it, is pretty much the thickness of the muscular envelope around the neck, or at least around the ventral half. And the cervical ribs are typically pretty close to the vertebral centra–only weirdos like Apatosaurus and Erketu displace them very far ventrally (see Taylor and Wedel 2013a: fig. 7 and this post). So, thin jackets of muscle around proportionally large vertebrae–or, if you like, corn-on-the-cob rather than shish-kebabs.
As for why sauropods have long cervical ribs, Mike and I discussed some possibilities in our 2013 PeerJ paper (Taylor and Wedel 2013a), and Preuschoft and Klein addressed the issue last fall in PLOS ONE (Preuschoft and Klein 2013). My favorite hypothesis is that long tendons allow an animal to shift the bulk of the muscle–and therefore the center of gravity–toward the base of the neck, but that long unossified tendons can be distorted through stretching, which wastes muscular energy. Ossifying those long tendons is like putting bony wheelbarrow handles on each vertebra, allowing the muscles to move the vertebra from a distance without so much wasted energy, and probably with finer positional control.
That’s a nifty hypothesis in need of testing, anyway. In fact, cervical ribs and their associated muscles could stand a lot more attention on both the descriptive and analytical fronts. I know that Liguo Li has some research in the works on different conformations of hypaxial muscles, tendons, and cervical ribs in birds (you know, when she’s not describing bizarre new titanosaurs like Yongjinglong — see Li et al. 2014). If you saw Peter Dodson give their talk at SVP last fall, you probably remember some stunning images of dissected bird necks. As a famous legislator once said, we shall watch her career with great interest.
- Dalla Vecchia, F.M. 1999. Atlas of the sauropod bones from the Upper Hauterivian – Lower Barremian of Bale/Valle (SW Istria, Croatia). Natura Nacosta 18:6-41.
- Janensch, Werner. 1950. Die Wirbelsaule von Brachiosaurus brancai. Palaeontographica (Suppl. 7) 3: 27-93.
- 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.
- Lacovara, Kenneth J.; Ibiricu, L.M.; Lamanna, M.C.; Poole, J.C.; Schroeter, E.R.; Ullmann, P.V.; Voegele, K.K.; Boles, Z.M.; Egerton, V.M.; Harris, J.D.; Martínez, R.D.; Novas, F.E. (September 4, 2014). A Gigantic, Exceptionally Complete Titanosaurian Sauropod Dinosaur from Southern Patagonia, Argentina. Scientific Reports. doi:10.1038/srep06196.
- Li L-G, Li D-Q, You H-L, Dodson P (2014) A New Titanosaurian Sauropod from the Hekou Group (Lower Cretaceous) of the Lanzhou-Minhe Basin, Gansu Province, China. PLoS ONE 9(1): e85979. doi:10.1371/journal.pone.0085979
- Preuschoft, H., & Klein, N. (2013). Torsion and Bending in the Neck and Tail of Sauropod Dinosaurs and the Function of Cervical Ribs: Insights from Functional Morphology and Biomechanics. PloS one, 8(10), e78574.
- Taylor, Michael P., and Mathew J. Wedel. 2013a. 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., Cifelli, R.L., and Sanders, R.K. 2000b. Osteology, paleobiology, and relationships of the sauropod dinosaurSauroposeidon. Acta Palaeontologica Polonica 45(4): 343-388.
My camera had a possibly-fatal accident in the field at the end of the day on Saturday, so I didn’t take any photos on Sunday or Monday. From here on out, you’re either getting my slides, or photos taken by other people.
On Sunday we were at the John Wesley Powell River History Museum in Green River, Utah, for the Cretaceous talks. There were some fossils on display downstairs, including mounted skeletons of Falcarius and one or two ornithischians,* and this sauropod humerus from the Cedar Mountain Formation (many thanks to Marc Jones for the photo).
* A ceratopsian and Animantarx, maybe? They were in the same room as the sauropod humerus, so it’s no surprise that I passed them by with barely a glance.
There were loads of great talks in the Cretaceous symposium on Sunday, and I learned a lot, about everything from clam shrimp biostratigraphy to ankylosaur phylogeny to Canadian sauropod trackways. But I can’t show you any slides from those talks, so the rest of this post is the abstact from Darren’s and my talk, illustrated by a few select slides.
Sauroposeidon is a giant titanosauriform from the Early Cretaceous of North America. The holotype is OMNH 53062, a series of four articulated cervical vertebrae from the Antlers Formation (Aptian-Albian) of Oklahoma. According to recent analyses, Paluxysaurus from the Twin Mountain Formation of Texas is the sister taxon of OMNH 53062 and may be a junior synonym of Sauroposeidon. Titanosauriform material from the Cloverly Formation of Wyoming may also pertain to Paluxysaurus/Sauroposeidon. The proposed synonymy is based on referred material of both taxa, however, so it is not as secure as it might be.
MIWG.7306 is a cervical vertebra of a large titanosauriform from the Wessex Formation (Barremian) of the Isle of Wight. The specimen shares several derived characters with the holotype of Sauroposeidon: an elongate cervical centrum, expanded lateral pneumatic fossae, and large, plate-like posterior centroparapophyseal laminae. In all of these characters, the morphology of MIWG.7306 is intermediate between Brachiosaurus and Giraffatitan on one hand, and Sauroposeidon on the other. MIWG.7306 also shares several previously unreported features of its internal morphology with Sauroposeidon: reduced lateral chambers (“pleurocoels”), camellate internal structure, ‘inflated’ laminae filled with pneumatic chambers rather than solid bone, and a high Air Space Proportion (ASP). ASPs for Sauroposeidon, MIWG.7306, and other isolated vertebrae from the Wessex Formation are all between 0.74 and 0.89, meaning that air spaces occupied 74-89% of the volume of the vertebrae in life. The vertebrae of these animals were therefore lighter than those of brachiosaurids (ASPs between 0.65 and 0.75) and other sauropods (average ASPs less than 0.65).
Sauroposeidon and MIWG.7306 were originally referred to Brachiosauridae. However, most recent phylogenetic analyses find Sauroposeidon to be a basal somphospondyl, whether Paluxysaurus and the Cloverly material are included or not. Given the large number of characters it shares with Sauroposeidon, MIWG.7306 is probably a basal somphospondyl as well. But genuine brachiosaurids also persisted and possibly even radiated in the Early Cretaceous of North America; these include Abydosaurus, Cedarosaurus, Venenosaurus, and possibly an as-yet-undescribed Cloverly form. The vertebrae of Abydosaurus have conservative proportions and solid laminae and the bony floor of the centrum is relatively thick. In these characters, Abydosaurus is more similar to Brachiosaurus and Giraffatitan than to Sauroposeidon or MIWG.7306. So not all Early Cretaceous titanosauriforms were alike, and whatever selective pressures led Sauroposeidon and MIWG.7306 to evolve longer and lighter necks, they didn’t prevent Giraffatitan-like brachiosaurs such as Abydosaurus and Cedarosaurus from persisting well into the Cretaceous.
The evolutionary dynamics of sauropods in the North American mid-Mesozoic are still mysterious. In the Morrison Formation, sauropods as a whole are both diverse and abundant, but Camarasaurus and an efflorescence of diplodocoids account for most of that abundance and diversity, and titanosauriforms, represented by Brachiosaurus, are comparatively scarce. During the Early Cretaceous, North American titanosauriforms seem to have radiated, possibly to fill some of the ecospace vacated by the regional extinction of basal macronarians (Camarasaurus) and diplodocoids. However, despite a flood of new discoveries in the past two decades, sauropods still do not seem to have been particularly abundant in the Early Cretaceous of North America, in contrast to sauropod-dominated faunas of the Morrison and of other continents during the Early Cretaceous.
That final slide deserves some explanation. On the way back from the field on Saturday–the night before my talk–a group of us stopped at a burger joint in Hanksville. Sharon McMullen got a kid’s meal, and it came in this bag. We took it as a good omen that Sauroposeidon was the first dinosaur listed in the quiz.
For the full program and abstracts from both days of talks, please download the field conference guidebook here.
One articulated Sauroposeidon to go, hold the perspective distortion, with a side of stinkin’ mammal
April 24, 2014
Sauroposeidon is stitched together from orthographic views of the 3D photogrammetric models rendered in MeshLab. Greyed out bits of the vertebrae are actually missing–I used C8 to patch C7, C7 to patch C6, and so on forward. The cervical ribs as reconstructed here were all recovered and they are in collections, but they’re in several jackets and boxes and therefore not easily photographed.
The meter bars are both one meter as advertised. The giraffe neck is FMNH 34426 (from this post), which is actually 1.7 meters long, but I scaled it up to 2.4 meters to match that of the tallest known giraffe. I think it’s cool that a world-record giraffe neck is roughly as long as two vertebrae from the middle of the neck of Sauroposeidon.
There are loads of little morphological details in the Sauroposeidon vertebrae that are clearer now than they were in our old photographs, but those will be stories for other posts.
April 18, 2014
I was in Oklahoma and Texas last week, seeing Sauroposeidon, Paluxysaurus, Astrophocaudia, and Alamosaurus, at the Sam Noble Oklahoma Museum of Natural History, the Fort Worth Museum of Science and History, the Shuler Museum of Paleontology at SMU, and the Perot Museum of Nature and Science, respectively. I have a ton of interesting things from that trip that I could blog about, but unfortunately I have no time. Ten days from now, I’m off to Colorado and Utah for the Mid-Mesozoic conference and field trip, and between now and then I need to finish up my bits on three collaborative papers, get my summer anatomy lectures posted for internal peer review here at WesternU, and–oh yeah–actually write my conference talk. Fun times.
BUT after being subjected to the horror of the Yale Brontosaurus skull, I figured you all deserved a little awesome.
So here’s me getting one of 351 photos of the most posterior and largest of the Sauroposeidon jackets (this is not the awesome, BTW, just a stop along the way). This jacket holds what I once inferred to be the back half of C7 and all of C8. Now that Sauroposeidon may be a somphospondyl rather than a brachiosaur, who knows what verts these are–basal somphospondyls have up to 17 cervicals to brachiosaurids’ probable 13 (for a hypothetical view of an even-longer-necked Sauroposeidon, see this probably-prophetic post by Mike). The vertically-mounted skeleton in the background is Cotylorhynchus. Cotylorhynchus got a lot bigger than that–up to maybe 6 meters long and 2 or 3 tons–and was probably the largest land animal that had ever existed back in the Early Permian. Photo by OU grad student Andrew Thomas, whom you’ll be hearing about more here in the future.
I couldn’t crank the model myself on the road, thanks to the pathetic lack of processing power in my 6-year-old laptop (which will be replaced RSN). Andy Farke volunteered to do the photogrammetricizing with Agisoft Photoscan, if only I’d DropBox him the pictures. Here’s a screenshot from MeshLab showing the result:
And my best taken-from-overhead quasi-lateral photograph:
If you’re curious, the meter stick at the top is actually one meter long, it just has the English measurement side showing. The giant caliper at the bottom is also marked off in inches, and it is open to 36.0 inches (it didn’t go to 1 meter, or I would have used that). You can tell that there is some perspective distortion involved here since 36 inches on the caliper is 1380 pixels, whereas the 39.4-inch meter stick is only 1341 pixels. Man, I hate scale bars. But they make good calibration targets.
Incidentally, after playing around with the model in orthographic mode in MeshLab, the distortions in the photos of the vertebrae themselves just scream at me. Finally, finally, I can escape the tyranny of perspective. Compare the ends of the big wooden beam at the top of the jacket to get a feel for how much the two views differ.
Working on Sauroposeidon again after all this time made me seriously nostalgic. I love that beast. I don’t think I’m exaggerating when I say that those vertebrae are the most gorgeous physical objects in the universe. Also, an appropriately huge thank-you to preparator Kyle Davies (of apatosaur-sculpting fame), collections manager Jen Larsen, and Andrew Thomas again for help with wrassling those verts around, and for sharing their thoughts and advice. Thanks also to curators Rich Cifelli and Nick Czaplewski for their hospitality and for the go-ahead to undertake this work, and to Andy Farke for generating the model.
I’ll have a lot more to say about this stuff in the future. I didn’t go to all this work just for giggles. For a long time I’ve had a hankering to do a paper on the detailed anatomy of Sauroposeidon, based on all of the things that I’ve noticed in the last decade that didn’t make it into any of the early papers. And now there’s the proposed synonymy of Paluxysaurus with Sauroposeidon. And “Angloposeidon” needs some attention–Darren and I have been thinking about writing “Angloposeidon II” for years now. And…well, plenty more.
So, loads more to come, but not for the next few weeks. Eventually I’ll be publishing all of this–the photos, the 3D models, the whole works. Stay tuned.
UPDATE a few days later
Man, I am frazzled, because I forgot to include the moral of the story: if I can do this, you can do this. There are good, free photogrammetry programs out there–Peter Falkingham published a whole paper on free photogrammetry in 2012, and posted a guide to an even better program, VisualSFM, on Academia.edu. Even Agisoft Photoscan is not prohibitively expensive–under $200 for an educational license. MeshLab is free and has hordes of good free tutorials. For the photography itself, you basically just build a virtual dome of photos around an object. If you need more instructions than that, Heinrich has written a whole series of tutorials. It doesn’t take a fancy camera–I used a point-and-shoot for the Sauroposeidon work shown here (a Canon S100 operating at 6 megapixels, if anyone is curious). What are you waiting for?
That last one really hurts. Here’s the original image, which should have gone in the paper with the interpretive trace next to it rather than on top of it:
Papers referenced in these slides:
- Taylor, M.P., and Wedel, M.J. 2013b. The effect of intervertebral cartilage on neutral posture and range of motion in the necks of sauropod dinosaurs. PLOS ONE 8(10): e78214. 17 pages. doi:10.1371/journal.pone.0078214
- Wedel, M.J. 2007a. What pneumaticity tells us about ‘prosauropods’, and vice versa. Special Papers in Palaeontology 77:207-222.
- Wedel, Mathew J., Richard L. Cifelli and R. Kent Sanders. 2000b. Osteology, paleobiology, and relationships of the sauropod dinosaur Sauroposeidon. Acta Palaeontologica Polonica 45(4): 343-388.
On that last slide, I also talked about two further elaborations: figures that take up the entire page, with the caption on a separate (usually facing) page, and side title figures, which are wider than tall and get turned on their sides to better use the space on the page.
Also, if I was doing this over I’d amend the statement on the last slide with, “but it doesn’t hurt you at all to be cognizant of these things, partly because they’re easy, and partly because your paper may end up at an outlet you didn’t anticipate when you wrote it.”
And I just noticed that the first slide in this group has the word ‘without’ duplicated. Jeez, what a maroon. I’ll try to remember to fix that before I post the whole slide set at the end of this exercise.
A final point: because I am picking illustrations from my whole career to illustrate these various points, almost all fail in some obvious way. The photos from the second slide should be in color, for example. When I actually gave this talk, I passed out reprints of several of my papers and said, “I am certain that every single figure I have ever made could be improved. So as you look through these papers, be thinking about how each one could be made better.”
- Wedel, M.J. 2003b. The evolution of vertebral pneumaticity in sauropod dinosaurs. Journal of Vertebrate Paleontology 23:344-357.
- 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, Mathew J., Richard L. Cifelli and R. Kent Sanders. 2000b. Osteology, paleobiology, and relationships of the sauropod dinosaurSauroposeidon. Acta Palaeontologica Polonica 45(4): 343-388.
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.