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.
This was inspired by an email Mike sent a couple of days ago:
Remind yourself of the awesomeness of Giraffatitan:
Now think of this. Its neck is 8.5m long. Knock of one measly meter — for example, by removing one vertebra from the middle of the neck — and you have 7.5 m.
Supersaurus’s neck was probably TWICE that long.
I replied that I was indeed freaked out, and that it had given me an idea for a post, which you are now reading. I didn’t have a Giraffatitan that was sufficiently distortion-free, so I used my old trusty Brachiosaurus. The vertebra you see there next to Mike and next to the neck of Brachiosaurus is BYU 9024, the longest vertebra that has ever been found from anything, ever.
Regarding the neck length of Supersaurus, and how BYU 9024 came to be referred to Supersaurus, here’s the relevant chunk of my dissertation (Wedel 2007: pp. 208-209):
Supersaurus is without question the longest-necked animal with preserved cervical material. Jim Jensen recovered a single cervical vertebra of Supersaurus from Dry Mesa Quarry in western Colorado. The vertebra, BYU 9024, was originally referred to “Ultrasauros”. Later, both the cervical and the holotype dorsal of “Ultrasauros” were shown to belong to a diplodocid, and they were separately referred to Supersaurus by Jensen (1987) and Curtice et al. (1996), respectively.
BYU 9024 has a centrum length of 1378 mm, and a functional length of 1203 mm (Figure 4-3). At 1400 mm, the longest vertebra of Sauroposeidon is marginally longer in total length [see this post for a visual comparison]. However, that length includes the prezygapophyses, which overhang the condyle, and which are missing from BYU 9024. The centrum length of the largest Sauroposeidon vertebra is about 1250 mm, and the functional length is 1190 mm. BYU 9024 therefore has the largest centrum length and functional length of any vertebra that has ever been discovered for any animal. Furthermore, the Supersaurus vertebra is much larger than the Sauroposeidon vertebrae in diameter, and it is a much more massive element overall.
Neck length estimates for Supersaurus vary depending on the taxon chosen for comparison and the serial position assumed for BYU 9024. The vertebra shares many similarities with Barosaurus that are not found in other diplodocines, including a proportionally long centrum, dual posterior centrodiapophyseal laminae, a low neural spine, and ventrolateral flanges that connect to the parapophyses (and thus might be considered posterior centroparapophyseal laminae, similar to those of Sauroposeidon). The neural spine of BYU 9024 is very low and only very slightly bifurcated at its apex. In these characters, it is most similar to C9 of Barosaurus. However, theproportions of the centrum of BYU 9024 are more similar to those of C14 of Barosaurus, which is the longest vertebra of the neck in AMNH 6341. BYU 9024 is 1.6 times as long as C14 of AMNH 6341 and 1.9 times as long as C9. If it was built like that of Barosaurus, the neck of Supersaurus was at least 13.7 meters (44.8 feet) long, and may have been as long as 16.2 meters (53.2 feet).
Based on new material from Wyoming, Lovelace et al. (2005 [published as Lovelace et al. 2008]) noted potential synapomorphies shared by Supersaurus and Apatosaurus. BYU 9024 does not closely resemble any of the cervical vertebrae of Apatosaurus. Instead of trying to assign its serial position based on morphology, I conservatively assume that it is the longest vertebra in the series if it is from an Apatosaurus-like neck. At 2.7 times longer than C11 of CM 3018, BYU 9024 implies an Apatosaurus-like neck about 13.3 meters
(43.6 feet) long.
Bonus comparo: BYU 9024 vs USNM 10865, the mounted Diplodocus longus at the Smithsonian, modified from Gilmore 1932 (plate 6). For this I scaled BYU 9024 against the 1.6-meter femur of this specimen.
If you’d like to gaze upon BYU 9024 without distraction, or put it into a composite of your own, here you go:
- Gilmore, C. W. 1932. On a newly mounted skeleton of Diplodocus in the United States National Museum. Proceedings of the United States National Museum 81, 1-21.
- Lovelace, David M., Scott A. Hartman and William R. Wahl. 2008. 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.
- Wedel, M.J. 2007. Postcranial pneumaticity in dinosaurs and the origin of the avian lung. PhD dissertation, University of California, Berkeley, Department of Integrative Biology, 303 pp.
June 9, 2014
We’ve touched on this several times in various posts and comment threads, but it’s worth taking a moment to think in detail about the various published mass estimates for the single specimen MB.R.2181 (formerly known as HMN SII), the paralectotype of Giraffatitan brancai, which is the basis of the awesome mounted skeleton in Berlin.
Here is the table of published estimates from my 2010 sauropod-history paper, augmented with the two more recent estimates extrapolated from limb-bone measurements:
|Author and date||Method||Volume (l)||Density (kg/l)||Mass (kg)|
|Janensch (1938)||Not specified||—||—||`40 t’|
|Colbert (1962)||Displacement of sand||86,953||0.9||78,258|
|Russell et al. (1980)||Limb-bone allometry||—||—||13,618|
|Anderson et al. (1985)||Limb-bone allometry||—||—||29,000|
|Paul (1988)||Displacement of water||36,585||0.861||31,500|
|Alexander (1989)||Weighing in air and water||46,600||1.0||46,600|
|Gunga et al. (1995)||Computer model||74,420||1.0||74,420|
|Christiansen (1997)||Weighing in air and water||41,556||0.9||37,400|
|Henderson (2004)||Computer model||32,398||0.796||25,789|
|Henderson (2006)||Computer model||—||—||25,922|
|Gunga et al. (2008)||Computer model||47,600||0.8||38,000|
|Taylor (2009)||Graphic double integration||29,171||0.8||23,337|
|Campione and Evans (2012)||Limb-bone allometry||—||—||35,780|
|Benson et al. (2014)||Limb-bone allometry||—||—||34,000|
(The estimate of Russell et al. (1980) is sometimes reported as 14900 kg. However, they report their estimate only as “14.9 t”; and since they also cite “the generally accepted figure of 85 tons”, which can only be a reference to Colbert (1962)”, we must assume that Russell et al. were using US tons throughout.)
The first thing to notice is that there is no very clear trend through time, either upwards or downwards. Here’s a plot of mass (y-axis) against year of estimate (x-axis):
I’ve not even tried to put a regression line through this: the outliers are so extreme they’d render it pretty much useless.
In fact, the lowest and highest estimates differ by a factor of 5.75, which is plainly absurd.
But we can go some way to fixing this by discarding the outliers. We can dump Colbert (1962) and Alexander (1989) as they used overweight toys as their references. We more or less have to dump Russell et al. (1980) simply because it’s impossible to take seriously. (Yes, this is the argument from personal incredulity, and I don’t feel good about it; but as Pual (1988) put it, “so little flesh simply cannot be stretched over the animal’s great frame”.) And we can ignore Gunga et al. (1995) because it used circular conic sections — a bug fixed by Gunga et al. (2008) by using elliptical sections.
With these four unpalatable outliers discarded, our highest and lowest estimates are those of Gunga et al. (2008) at 38,000 kg and Taylor (2009)at 23,337. The former should be taken seriously as it was done using photogrammetrical measurements of the actual skeletal mount. And so should the latter because Hurlburt (1999) showed that GDI is generally the least inaccurate of our mass-estimation techniques. That still gives us a factor of 1.63. That’s the difference between a lightweight 66 kg man and and overweight 108 kg.
Here’s another way of thinking about that 1.63 factor. Assuming two people are the same height, one of them weighing 1.62 times as much as the other means he has to be 1.28 times as wide and deep as the first (1.28^2 = 1.63). Here is a man next to his 1.28-times-as-wide equivalent:
I would call that a very noticeable difference. You wouldn’t expect someone estimating the mass of one of these men to come up with that of the other.
So what’s going on here? I truly don’t know. We are, let’s not forget, dealing with a complete skeletal mount here, one of the very best sauropod specimens in the world, which has been extensively studied for a century. Yet even within the last six years, we’re getting masses that vary by as much as the two dudes above.
June 8, 2014
This is BYU 12867–you’ve seen it here before–in dorsal view. It’s not a brilliant shot–I took it through the glass of the display case while filming a documentary at the North American Museum of Ancient Life in Lehi, Utah, in 2008. Centrum length is 94 cm, total length with the overhanging prezygapophyses is over a meter.
June 5, 2014
Back in 2008, when I did the GDI of Giraffatitan and Brachiosaurus for my 2009 paper on those genera, I came out with estimates of 28688 and 23337 kg respectively. At the time I said to Matt that I was suspicious of those numbers because they seemed too low. He rightly told me to shut up and put my actual results in the paper.
More recently, Benson et al. (2014) used limb-bone measurements to estimate the masses of the same individuals as 56000 and 34000 kg. When Ian Corfe mentioned this in a comment, my immediate reaction was to be sceptical: “I’m amazed that the two more recent papers have got such high estimates for brachiosaurs, which have the most gracile humeri of all sauropods“.
So evidently I have a pretty strong intuition that Brachiosaurus massed somewhere in the region of 35000 kg and Giraffatitan around 30000 kg. But why? Where does that intuition come from?
I can only assume that my strongly held ideas are based only on what I’d heard before. Back when I did my 2008 estimate, I probably had in mind things like Paul’s (1998) estimate of 35000 kg for Brachiosaurus, and Christiansen’s (1997:67) estimate of 37400 for Giraffatitan. Whereas by the time the Benson et al. paper came out I’d managed to persuade myself that my own much lower estimates were right. In other words, I think my sauropod-mass intuition is based mostly on sheer mental inertia, and so should be ignored.
I’m guessing I should ignore your intuitions about sauropod masses, too.
- Benson Roger B. J., Nicolás E. Campione, Matthew T. Carrano, Philip D. Mannion, Corwin Sullivan, Paul Upchurch, and David C. Evans. (2014) Rates of Dinosaur Body Mass Evolution Indicate 170 Million Years of Sustained Ecological Innovation on the Avian Stem Lineage. PLoS Biology 12(5):e1001853. doi:10.1371/journal.pbio.1001853
- Christiansen, Per. 1997. Locomotion in sauropod dinosaurs. Gaia 14:45-75.
- Paul, Gregory S. 1998. Terramegathermy and Cope’s Rule in the land of titans. Modern Geology 23:179-217.
- Taylor, Michael P. 2009. A re-evaluation of Brachiosaurus altithorax Riggs 1903 (Dinosauria, Sauropoda) and its generic separation from Giraffatitan brancai (Janensch 1914). Journal of Vertebrate Paleontology 29(3):787-806.
In recent photo posts on the mounted Brachiosaurus skeleton and its bones in the ground, I’ve lamented that the Field Museum’s online photo archive is so unhelpful: for example, if it has a search facility, I’ve not been able to find it.
But the good news is that there’s a Field Museum Photo Archives tumblr. Its coverage is of course spotty, but it gives us at least some chance of finding useful brachiosaur images. Like this one of the sixth presacral vertebra (i.e. probably D7 in a column of 12 dorsals):
It’s instructive to compare that with Riggs’s (1904: plate LXXII) illustration of the same vertebra in the same aspect, in which he almost literally airbrushed out the jigsaw-puzzle complexity of the preserved bone surface:
More disturbing still, compare that old photograph with my own (terrible) 2005 photo of the same vertebra:
It looks very much as though the vertebra itself — not just Riggs’s illustration — has been “improved” since the older photo was taken exactly a century earlier in 1905. This is a constant problem when dealing with old fossils.
Here are three more interesting photos from the Tumblr. First, the Brachiosaurus fossils in the field:
This is evidently from later in the excavation process than the previous photo of this area, since much of the material is now jacketed. That’s the femur in front of shot, here seen in anteromedial view, with the top towards the right.
Next up, this photo purports to be “Thirteen men including Security Guard unloading dorsal vertebrae of type specimen Brachiosaurus fossil”:
But in fact it’s not Brachiosaurus — the neural spines are too tall and slender. I am pretty sure this is Riggs’s Apatosaurus — the rightmost dorsal has that distinctive notch on the dorsal aspect of the neural spine. And indeed, checking his monograph on that specimen (Riggs 1903: plate XLVI), I see that its dorsals were distorted in this way, and that the front-centre vert is a fine match for its D10.
Finally, there’s this one of the prep room:
On the far left, we have the still-jacketed Brachiosaurus femur; next to it stands Harold W. Menke, who discovered the fossil; and to his right is Elmer S. Riggs, who wrote the description.
Those are all the Brachiosaurus-related images I’ve been able to find on the tumblr. But do let me know if you find any others.
- Riggs, Elmer S. 1903. Structure and relationships of opisthocoelian dinosaurs. Part I: Apatosaurus Marsh. Field Columbian Museum, Geological Series 2:165-196.
- Riggs, Elmer S. 1904. Structure and relationships of opisthocoelian dinosaurs. Part II, the Brachiosauridae. Field Columbian Museum, Geological Series 2:229-247.
June 3, 2014
Continuing our Brachiosaurus series [part 1, part 2, part 3, part 4, part 5, part 6, part 7], here is another historically important photo scanned from the Glut encyclopaedia: this time, from Supplement 1 (2000), page 157.
This is the Brachiosaurus altithorax holotype FMNH P25107 in the field, at Grand Junction, Colorado, in 1900. Clearly visible are the seven presacral vertebrae running across the middle of the photo (upside-down, so we see their ventral surfaces), several ribs on either side, and the end of a long-bone to the left — most likely the distal end of the femur. The flat bone at bottom left is probably part of the ilium, with the circular cut-out being the acetabulum. (The caption also mentions the sacrum, which I can’t see.)
As with the photo of the mounted skeleton in the museum, this is one of the Field Museum’s own photos — neg. #4027 — but I can’t find a better copy online. It’s got to be out there somewhere — can anyone help?
Glut, Donald F. 2000. Dinosaurs: The Encyclopedia: Supplement 1. McFarland & Company, Inc., Jefferson, North Carolina. 442 pages.