October 20, 2015
I just gave an answer to this question on Quora, and it occurred to me that I ought to also give it a permanent home here. So here it is.
This is a great example of a question that you’d think would have a simple, clear answer, but doesn’t. In fact, as a palaeontologist specialising in dinosaur gigantism, I have an abiding fear of being asked this question in a pub quiz, and not being able to produce the name that’s written on the quizmaster’s answer sheet.
First, what do we mean by “biggest”? Diplodocus was longer than Apatosaurus, but Apatosaurus was heavier. Giraffatitan was taller than either. Let’s simplify and assume we want to know the heaviest dinosaur.
Second, estimating the masses of extinct animals is incredibly hard even when we have a pretty complete skeleton. For example, the gigantic mounted brachiosaur skeleton in Berlin (which used to be called “Brachiosaurus” brancai but is now recognised as the separate genus Giraffatitan) has been subject to at least 14 estimates in the published scientific literature,. They vary from 13,618 kg to 78,258 kg — a factor of 5.75 for the same individual. That’s like looking at a human skeleton and not knowing whether its from Kate Moss or Arnold Schwazenegger. (There are reasons for this and I urge you to read the linked article.)
Third, the big dinosaurs tend to be very poorly represented. Giraffatitan is probably the heaviest dinosaur known from a more or less complete skeleton (though even that is put together from several different individuals) so I could give that as the answer to the hypothetical pub-quiz — though the answer sheet would probably be out of date and call it Brachiosaurus.
Fourth, which individual of a given species do we mean? I said Giraffatitan is known from a more or less complete skeleton. And my best guess is that that individual massed, say, 30,000 kg. But an isolated fibula of the same species is known that’s 12.6% longer than the one in the skeletal mount. That suggest an animal that masses 1.126^3 = 1.43 times as massive as the mounted skeleton — say 43,000 kg. There might be yet bigger Giraffatitan individuals. On the other hand, there isthat Apatosaurus, which is usually thought of as not being so big, might have got even bigger.
Fifth, the very biggest specimens tend to be known from only a handful of bones. A good example here is the titanosaur Argentinosaurus, which is known from several vertebrae and a few limb bones, but not all from the same individual. It’s a good bet that it massed 60-70 tonnes — so maybe about twice as much as Giraffatitan, but much less than the often-cited 100 tonnes. Other, more recently discovered, titanosaurs seem to be in the same size class: Puertasaurus, Futalognkosaurus, Dreadnoughtus and more. They they are hard to compare directly due to the paucity of overlapping material, or at least . (Scientists are working on getting more of this stuff properly described in the literature, which will help.)
But, sixth, the very biggest dinosaurs tend to be apocryphal. There’s Amphicoelias fragillimus,. This may have been 50 m long and massed 80 tonnes; but other published estimates say 58 m and 122 tonnes. We really can’t say from the very poor remains.
So if you get asked this question in a pub quiz, your best bet is to roll a dice, pick an answer, close your eyes and hope. Roll 1 for Giraffatitan, 2 for Brachiosaurus, 3 for Apatosaurus, 4 for Argentinosaurus, 5 for Dreadnoughtus and 6 for Amphicoelias fragillimus. Good luck!
October 19, 2015
I imagine that by now, everyone who reads this blog is familiar with Mark Witton’s painting of a giant azhdarchid pterosaur alongside a big giraffe. Here it is, for those who haven’t seen it:
(This is the fifth and most recent version that Mark has created, taken from 9 things you may not know about giant azhdarchid pterosaurs.)
It’s one of those images that really kicks you in the brain the first time you see it. The idea that an animal the size of a giraffe could fly under its own power seems ludicrous — yet that’s what the evidence tells us.
But wait — what do we mean by “an animal the size of a giraffe”? Yes, the pterosaur in this image is the same height as the giraffe, but how does its weight compare?
Mark says “The giraffe is a big bull Masai individual, standing a healthy 5.6 m tall, close to the maximum known Masai giraffe height.” He doesn’t give a mass, but Wikipedia, citing Owen-Smith (1988), says “Fully grown giraffes stand 5–6 m (16–20 ft) tall, with males taller than females. The average weight is 1,192 kg (2,628 lb) for an adult male and 828 kg (1,825 lb) for an adult female with maximum weights of 1,930 kg (4,250 lb) and 1,180 kg (2,600 lb) having been recorded for males and females, respectively.” So it seems reasonable to use a mass intermediate between those of an average and maximum-sized male, (1192+1930)/2 = 1561 kg.
So much for the giraffe. What does the azhdarchid weigh? The literature is studded with figures that vary wildly, from the 544 kg that Henderson (2010) found for Quetzalcoatlus, right down to the widely cited 70 kg that Chatterjee and Templin (2004) found for the same individual — and even the astonishing 50 kg that seems to be favoured by Unwin (2005:192). In the middle is the 259 kg of Witton (2008).
It occurred to me that I could visualise these mass estimates by shrinking the giraffe in Mark’s image down to the various proposed masses, and seeing how credible it looks to imagine these reduced-sized giraffes weighting the same as the azhdarchid. The maths is simple. For each proposed azhdarchid mass, we figure out what it is as a proportion of the giraffe’s 1561 kg; then the cube root of that mass proportion gives us the linear proportion.
- 544 kg = 0.389 giraffe masses = 0.704 giraffe lengths
- 259 kg = 0.166 giraffe masses = 0.549 giraffe lengths
- 70 kg =0.0448 giraffe masses = 0.355 giraffe lengths
Let’s see how that looks.
On the left, we have Mark’s artwork, with the giraffe massing 1561 kg. On the right, we have three smaller (isometrically scaled) giraffes of masses corresponding to giant azhdarchid mass estimates in the literature. If Don Henderson (2010) is right, then the pterosaur weighs the same as the 544 kg giraffe, which to me looks pretty feasible if it’s very pneumatic. If Witton (2008) is right, then it weighs the same as the 259 kg giraffe, which I find hard to swallow. And if Chatterjee and Templin (2004) are right, then the giant pterosaur weighs the same as the teeny tiny 70 kg giraffe, which I find frankly ludicrous. (For that matter, 70 kg is in the same size-class as Georgia, the human scale-bar: the idea that she and the pterosaur weigh the same is just silly.)
What is the value of such eyeball comparisons? I’m not sure, beyond a basic reality check. Running this exercise has certainly made me sceptical about even the 250 kg mass range which now seems to be fairly widely accepted among pterosaur workers. Remember, if that mass is correct then the pterosaur and the 259 kg giraffe in the picture above weight the same. Can you buy that?
Or can we find extant analogues? Are there birds and mammals with the same mass that are in the same size relation as these images show?
- Chatterjee, Sankar, and R. J. Templin. 2004. Posture, locomotion, and paleoecology of pterosaurs. Geological Society of America, Special Paper 376. 68 pages.
- Henderson, Donald M. 2010. Pterosaur body mass estimates from three-dimensional mathematical slicing. Journal of Vertebrate Paleontology 30(3):768-785.
- Witton, Mark P. 2008. A new approach to determining pterosaur body mass and its implications for pterosaur flight. Zitteliana 28:143-159.
June 12, 2015
I was contacted recently by David Goldenberg (firstname.lastname@example.org), a journalist who’s putting together a piece on the biggest dinosaurs. He asked me a few questions, and since I’d taken the time to write answers I thought I may as well post them here.
1) Do you think that we will ever know what the largest dinosaur (by mass) was?
In principle, we can never know that we’ve found the largest dinosaur. All we can know (and we probably can’t really know even this, as we’ll see below) is that we’ve found the largest so far. If we were dealing with animals where there’s a good sample size, there would be statistical techniques that we could use to figure out the likely size-range. But most giant dinosaur species are known only a handful of specimens — sometimes only a single one. How big did Puertasaurus get? We can’t possibly say: the best we can do is estimate how big the one known specimen of Puertasaurus was.
That said, we can sort of get a feel for size classes. There are quite a few sauropods that seem to come in at around 30-40 tonnes — Brachiosaurus, Giraffatitan, Supersaurus, Dreadnoughtus — which suggests there might be some kind of a limit there. But there are bigger titanosaurs (Argentinosaurus, Puertasaurus, Futalognkosaurus) which show that if the barrier exists at all, it’s a “soft” one. And of course the tantalising hints of super-giant sauropods.
There are at least three of these: Amphicoelias fragillimus, a diplodocid known from a drawing of a vertebral arch which has since been lost or destroyed, which could well have massed 100 tonnes. Bruhathkayosaurus, a giant titanosaur known from a two-meter tibia, since destroyed, which could conceivably have massed twice that; and the Broome Sandstone track-maker, known only from footprints, which might have been somewhere in between.
Any one of those, we might write off and say it’s too good to be true — all three stories are pretty vague as to evidence and require a lot of guesswork in the inferences. But the fact that we have all three of these makes me feel pretty certain that there were indeed sauropods out there in the 100-200 tonne range (i.e. the size of big whales). I only hope we find solid, verifiable, curated evidence for them some time soon.
2) What bones do you need to have before you can make an accurate measurement?
You can’t ever make an accurate measurement. Consider even a really well represented, essentially complete specimen such as MB.R.2181 (previously known as HM S II), the giant mounted skeleton in the Museum für Naturkunde Berlin. Peer-reviewed published estimates of the mass of that one individual have varied between 13,618 and 78,258 kg — a factor of 5.75. Even if you discard these obvious outlier estimates, recent and credible estimates vary from 23,337 to 38,000 kg, which is still a factor of 1.63.
And this is not completely crazy. Two humans with essentially identical skeletons can weigh 70 and 114 kg, after all. Soft tissue is essentially impossible to predict.
3) What do you make of the fact that so many different species have been given the title? Is that the fault of the media or scientists or what?
A big part of is that it depends on what you count. That Berlin brachiosaur is the biggest dinosaur known from an essentially complete skeleton, so Giraffatitan is a legitimate holder of the crown. (Confusing matters further, it used to be thought to be a species of Brachiosaurus). But there were definitely bigger sauropods than that — just not known from such complete specimens. Argentinosaurus was certainly bigger, for example. But there’s no way to put a meaningful whole-body mass estimate on it.
But yes, there is also an understandable tendency towards sensationalism, both from scientists and the press. There have been plenty of new discoveries that can legitimately be described as “could be the biggest yet”.
March 3, 2015
I’ve been taking a long-overdue look at some of the recently-described giant sauropods from China, trying to sort out just how big they were. Not a new pursuit for me, just one I hadn’t been back to in a while. Also, I’m not trying to debunk anything about this animal – as far as I know, there was no bunk to begin with – I’m just trying to get a handle on how big it might have been, for my own obscure purposes.
‘Huanghetitan’ ruyangensis was named by Lu et al. (2007) on the basis of a sacrum, the first 10 caudal vertebrae, some dorsal ribs and haemal arches, and a partial ischium. The holotype is 41HIII-0001 in the Henan Geological Museum. Lu et al. (2007) referred the new animal to the genus Huanghetitan, which was already known from the type species H. liujiaxiaensis (You et al., 2006). However, Mannion et al. (2013) found that the two species are not sister taxa and therefore ‘H.’ ruyangensis probably belongs to another genus, which has yet to be erected. Hence my use of scare quotes around the genus name.
Here’s the sacrum of ‘H.’ ruyangensis from Lu et al. (2007: fig. 2). The original small scale bar is supposed to be 10cm. You know how I feel about scale bars (or maybe you don’t, in which case read this and this), but in this case the scale seems pretty legit based on limited measurements that are also given in the paper. I comped in the sacrum of Brachiosaurus altithorax FMNH P25107 from this post (many thanks to Phil Mannion for the photos!), and scaled it according to the max width across the second pair of sacral ribs, which Riggs (1904: p. 236) gives as 105 cm. The sacrum of ‘H.’ ruyangensis is a little bigger, but not vastly bigger. ‘H.’ ruyangensis had six sacrals to Brachiosaurus‘s five, so extra length is mostly illusory, whereas the extra width is mostly legit.
According to Lu et al. (2007), the anterior face of the first caudal vertebra in ‘H.’ ruyangensis measures 26.9 cm tall by 32 cm wide, and the centrum is 18.2 cm long. The same measurements in Brachiosaurus are 28 x 33 cm for the anterior face and 16 cm for the centrum length. It’s basically a tie.
What about the big rib? Lu et al. (2007) show a complete dorsal rib of ‘H.’ ruyangensis that is 293 cm long. That’s nothing to sniff at – the longest rib of Brachiosaurus, and the cause for the specific name altithorax (‘tall-bodied’), measures 274.5 cm, so the ‘H.’ ruyangensis rib is about 7% longer. But it’s not the longest rib known for any sauropod. As far as I know, that honor goes to a Supersaurus dorsal rib measuring 305 cm (Lovelace et al., 2008). The biggest Supersaurus caudal also blows away the caudals of both ‘H.’ ruyangensis and Brachiosaurus, with a anterior face 39 cm tall by 46 cm wide. But then diplodocids were all about that bass, so there’s not much point in comparing tail size with a titanosauriform if you’re trying to get a handle on overall body size. Still, the 35-40 ton Supersaurus shows that you can have 3-meter ribs without being anywhere near Argentinosaurus territory, mass-wise.
So what’s the verdict? ‘H.’ ruyangensis was a little bigger than the holotype of Brachiosaurus altithorax, but only by a few percent. It might have been about the same size as the XV2 specimen of Giraffatitan brancai. Or, who knows, it could have had completely different proportions and massed considerably more (or less). But on the current evidence, it doesn’t seem to have been one of the biggest sauropods of all time. I hope we get some more of it one of these days.
- 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.
- Lu J., Xu, L., Zhang, X., Hu, W., Wu, Y., Jia, S., and Ji, Q. 2007. A new gigantic sauropod dinosaur with the deepest known body cavity from the Cretaceous of Asia. Acta Geologica Sinica, 81: 167-176.
- Mannion, P.D., Upchurch, P., Barnes, R.N., and Mateus, O. 2013. Osteology of the Late Jurassic Portuguese sauropod dinosaur Lusotitan atalaiensis (Macronaria) and the evolutionary history of basal titanosauriforms. Zoological Journal of the Linnean Society, 168(1): 98-206.
- Riggs, E.S. 1904. Structure and relationships of opisthocoelian dinosaurs. Part II, the Brachiosauridae. Field Columbian Museum, Geological Series 2, 6, 229-247.
- You, H., Li, D., Zhou, L., and Ji, Q. 2006. Huanghetitan liujiaxiaensis. a new sauropod dinosaur from the Lower Cretaceous Hekou Group of Lanzhou Basin, Gansu Province, China. Geological Review, 52 (5): 668-674.
December 12, 2014
If you’ve been reading around about Aquilops, you’ve probably seen it compared in size to a raven, a rabbit, or a cat. Where’d those comparisons come from? You’re about to find out.
Back in April I ran some numbers to get a rough idea of the size of Aquilops, both for my own interest and so we’d have some comparisons handy when the paper came out.
I started with the much more completely known Archaeoceratops. The measurements of Scott Hartman’s skeletal recon (shown above and on Scott’s website – thanks, Scott!) match the measurements of the Archaeo holotype given by Dodson and You (2003) almost perfectly. The total length of Archaeoceratops, including tail, is almost exactly one meter. Using graphic double integration, I got a volume of 8.88L total for a 1m Archaeoceratops. That would come down to 8.0L if the lungs occupied 10% of body volume, which is pretty standard for non-birds. So that’s about 17-18 lbs.
Archaeoceratops has a rostrum-jugal length of 145mm, compared to 84mm in Aquilops. Making the conservative assumption that Aquilops = Archaeoceratops*0.58, I got a body length of 60cm (about two feet), and volumes of 1.73 and 1.56 liters with and without lungs, or about 3.5 lbs in life. The internet informed me that the common raven, Corvus corax, has an adult length of 56-78 cm and a body mass of 0.7-2 kg. So, based on this admittedly tall and teetering tower of assumptions, handwaving, and wild guesses, Aquilops (the holotype individual, anyway) was about the size of a raven, in both length and mass. But ravens, although certainly well-known, are maybe a bit remote from the experience of a lot of people, so we wanted a comparison animal that more people would be familiar with. The estimated length and mass of the holotype individual of Aquilops also nicely overlap the species averages (60 cm, 1.4-2.7 kg) for the black-tailed jackrabbit, Lepus californicus, and they’re pretty close to lots of other rabbits as well, hence the comparison to bunnies.
Of course, ontogeny complicates things. Aquilops has some juvenile characters, like the big round orbit, but it doesn’t look like a hatchling. Our best guess is that it is neither a baby nor fully grown, but probably an older juvenile or young subadult. A full-grown Aquilops might have been somewhat larger, but almost certainly no larger than Archaeoceratops, and probably a meter or less in total length. So, about the size of a big housecat. That’s still pretty darned small for a non-avian dinosaur.
Although Aquilops represents everything I normally stand against – ornithischians, microvertebrates, heads – I confess that I have a sneaking affection for our wee beastie. Somebody’s just gotta make a little plush Aquilops, right? When and if that happens, you know where to find me.
- Dodson, P., and You, H.L. 2003. Redescription of neoceratopsian dinosaur Archaeoceratops and early evolution of Neoceratopsia. Acta Palaeontologica Polonica 48(2): 261-272.
- Farke, A.A., Maxwell, W.D., Cifelli, R.L., and Wedel, M.J. 2014. A ceratopsian dinosaur from the Lower Cretaceous of Western North America, and the biogeography of Neoceratopsia. PLoS ONE 9(12): e112055. doi:10.1371/journal.pone.0112055
December 10, 2014
Today sees the description of Aquilops americanus (“American eagle face”), a new basal neoceratopsian from the Cloverly Formation of Montana, by Andy Farke, Rich Cifelli, Des Maxwell, and myself, with life restorations by Brian Engh. The paper, which has just been published in PLOS ONE, is open access, so you can download it, read it, share it, repost it, remix it, and in general do any of the vast scope of activities allowed under a CC-BY license, as long as we’re credited. Here’s the link – have fun.
Obviously ceratopsians are much more Andy’s bailiwick than mine, and you should go read his intro post here. In fact, you may well be wondering what the heck a guy who normally works on huge sauropod vertebrae is doing on a paper about a tiny ceratopsian skull. The short, short version is that I’m here because I know people.
The slightly longer version is that OMNH 34557, the holotype partial skull of Aquilops, was discovered by Scott Madsen back in 1999, on one of the joint Cloverly expeditions that Rich and Des had going on at the time (update: read Scott’s account of the discovery here). That the OMNH had gotten a good ceratopsian skull out of Cloverly has been one of the worst-kept secrets in paleo. But for various complicated reasons, it was still unpublished when I got to Claremont in 2008. Meanwhile, Andy Farke was starting to really rock out on ceratopsians at around that time.
For the record, the light bulb did not immediately go off over my head. In fact, it took a little over a year for me to realize, “Hey, I know two people with a ceratopsian that needs describing, and I also know someone who would really like to head that up. I should put these folks together.” So I proposed it to Rich, Des, and Andy in the spring of 2010, and here we are. My role on the paper was basically social glue and go-fer. And I drew the skull reconstruction – more on that in the next post.
Anyway, it’s not my meager contribution that you should care about. I am fairly certain that, just as Brontomerus coasted to global fame on the strength of Paco Gasco’s dynamite life restoration, whatever attention Aquilops gets will be due in large part to Brian Engh’s detailed and thoughtful work in bringing it to life – Brian has a nice post about that here. I am very happy to report that the three pieces Brian did for us – the fleshed-out head that appears at the top of this post and as Figure 6C in the paper, the Cloverly environment scene with the marauding Gobiconodon, and the sketch of the woman holding an Aquilops – are also available to world under the CC-BY license. So have fun with those, too.
Finally, I need to thank a couple of people. Steve Henriksen, our Vice President for Research here at Western University of Health Sciences, provided funds to commission the art from Brian. And Gary Wisser in our scientific visualization center used his sweet optical scanner to generate the hi-res 3D model of the skull. That model is also freely available online, as supplementary information with the paper. So if you have access to a 3D printer, you can print your own Aquilops – for research, for teaching, or just for fun.
Next time: Aquilöps gets röck döts.
Farke, A.A., Maxwell, W.D., Cifelli, R.L., and Wedel, M.J. 2014. A ceratopsian dinosaur from the Lower Cretaceous of Western North America, and the biogeography of Neoceratopsia. PLoS ONE 9(12): e112055. doi:10.1371/journal.pone.0112055
September 15, 2014
The body mass should be considerably lower because the reconstructed column don’t match with published vertebrae centra lengths. 3D reconstruction also leaves too much space between vertebrae. The reconstruction body trunk is probably 15-20% longer than it really was. Check the supplementary material: http://www.nature.com/srep/2014/140904/srep06196/extref/srep06196-s1.pdf
So I did. The table of measurements in the supplementary material is admirably complete. For all of the available dorsal vertebrae except D9, which I suppose must have been too poorly preserved to measure the difference, Lacovara et al. list both the total centrum length and the centrum length minus the anterior condyle. Centrum length minus the condyle is what in my disseration I referred to as “functional length”, since it’s the length that the vertebra actually contributes to the articulated series, assuming that the condyle of one vertebra sticks out about as far as the cotyle is recessed on the next vertebra. Here are total lengths/functional lengths/differences for the seven preserved dorsals, in mm:
- D4 – 400/305/95
- D5 – 470/320/150
- D6 – 200/180/20
- D7 – 300/260/40
- D8 – 350/270/80
- D9 – 410/ – / –
- D10 – 330/225/105
The average difference between functional length and total length is 82 mm. If we apply that to D9 to estimate it’s functional length, we get 330mm. The summed functional lengths of the seven preserved vertebrae are then 1890 mm. What about the missing D1-D3? Since the charge is that Lacovara et al. (2014) restored Dreadnoughtus with a too-long torso, we should be as generous as possible in estimating the lengths of the missing dorsals. In Malawisaurus the centrum lengths of D1-D3 are all less than or equal to that of D4, which is the longest vertebra in the series (Gomani 2005: table 3), so it seems simplest here to assign D1-D3 functional lengths of 320 mm. That brings the total functional length of the dorsal vertebral column to 2850 mm, or 2.85 m.
At this point on my first pass, I was thinking that Lacovara et al. (2014) were in trouble. In the skeletal reconstruction that I used for the GDI work in the last post, I measured the length of the dorsal vertebral column as 149 pixels. Divided by 36 px/m gives a summed dorsal length of 4.1 m. That’s more than 40% longer than the summed functional lengths of the vertebrae calculated above (4.1/2.85 = 1.44). Had Lacovara et al. really blown it that badly?
Before we can rule on that, we have to estimate how much cartilage separated the dorsal vertebrae. This is a subject of more than passing interest here at SV-POW! Towers–the only applicable data I know of are the measurements of intervertebral spacing in two juvenile apatosaurs that Mike and I reported in our cartilage paper last year (Taylor and Wedel 2013: table 3, and see this post). We found that the invertebral cartilage thickness equaled 15-24% of the length of the centra.* For the estimated 2.85-meter dorsal column of Dreadnoughtus, that means 43-68 cm of cartilage (4.3-6.8 cm of cartilage per joint), for an in vivo dorsal column length of 3.28-3.53 meters. That’s still about 15-20% shorter than the 4.1 meters I measured from the skeletal recon–and, I must note, exactly what Asier stated in his comment. All my noodling has accomplished is to verify that his presumably off-the-cuff estimate was spot on. But is that a big deal?
Visually, a 20% shorter torso makes a small but noticeable difference. Check out the original reconstruction (top) with the 20%-shorter-torso version (bottom):
FWIW, the bottom version looks a lot more plausible to my eye–I hadn’t realized quite how weiner-dog-y the original recon is until I saw it next to the shortened version.
In terms of body mass, the difference is major. You’ll recall that I estimated the torso volume of Dreadnoughtus at 32 cubic meters. Lopping off 20% means losing 6.4 cubic meters–about the same volume as a big bull elephant, or all four of Dreadnoughtus‘s limbs put together. Even assuming a low whole-body density of 0.7 g/cm^3, that’s 4.5 metric tons off the estimated mass. So a ~30-ton Dreadnoughtus is looking more plausible by the minute.
For more on how torso length can affect the visual appearance and estimated mass of an animal, see this post and Taylor (2009).
* I asked Mike to do a review pass on this post before I published, and regarding the intervertebral spacing derived from the juvenile apatosaurs, he wrote:
That 15-24% is for juveniles. For the cervicals of adult Sauroposeidon we got about 5%. Why the differences? Three reasons might be relevant: 1, taxonomic difference between Sauroposeidon and Apatosaurus; 2, serial difference between neck and torso; 3, ontogenetic difference between juvenile and adult. By applying the juvenile Apatosaurus dorsal measurement directly to the adult Dreadnoughtus dorsals, you’re implicitly assuming that the adult/juvenile axis is irrelevant (which seems unlikely to me), that the taxonomic axis is (I guess) unknowable, and that the cervical/dorsal distinction is the only one that matter.
That’s a solid point, and it deserves a post of its own, which I’m already working on. For now, it seems intuitively obvious to me that we got a low percentage on Sauroposeidon simply because the vertebrae are so long. If the length-to-diameter ratio was 2.5 instead of 5, we’d have gotten 10%, unless cartilage thickness scales with centrum length, which seems unlikely. For a dorsal with EI of 1.5, cartilage thickness would then be 20%, which is about what I figured above.
Now, admittedly that is arm-waving, not science (and really just a wordy restatement of his point #2). The obvious thing to do is take all of our data and see if intervertebral spacing is more closely correlated with centrum length or centrum diameter. Now that it’s occurred to me, it seems very silly not to have done that in the actual paper. And I will do that very thing in an upcoming post. For now I’ll just note three things:
- As you can see from figure 15 in our cartilage paper, in the opisthocoelous anterior dorsals of CM 3390, the condyle of the posterior vertebra is firmly engaged in the cotyle of the anterior one, and if anything the two vertebrae look jammed together, not drifted apart. But the intervertebral spacing as a fraction of centrum length is still huge (20+4%) because the centra are so short.
- Transferring these numbers to Dreadnoughtus only results in 4.3-6.8 cm of cartilage between adjacent vertebrae, which does not seem unreasonable for a 30- or 40-ton animal with dorsal centra averaging 35 cm in diameter. If you asked me off the cuff what I thought a reasonable intervertebral spacing was for such a large animal, I would have said 3 or 4 inches (7.5 to 10 cm), so the numbers I got through cross-scaling are actually lower than what I would have guessed.
- Finally, if I’ve overestimated the intervertebral spacing, then the actual torso length of Dreadnoughtus was even shorter than that illustrated above, and the volumetric mass estimate would be smaller still. So in going with relatively thick cartilage, I’m being as generous as possible to the Lacovara et al. (2014) skeletal reconstruction (and indirectly to their super-high allometry-derived mass estimate), which I think is only fair.
- Gomani, Elizabeth M. 2005. Sauropod dinosaurs from the Early Cretaceous of Malawi, Africa. Palaeontologia Electronica 8(1):27A (37 pp.)
- 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.
- 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.
- Taylor, Michael P., and Mathew J. Wedel. 2013c. 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