Sauropod-art-O-rama!

Scaled restoration of the giant titanosaur Puertasaurus by Nima Sassani, from the Art Evolved Sauropod Gallery

Get on over to Art Evolved and scope out the sauroponderous Sauropod Gallery. It’s brobdingnaginormous. I don’t want to seem biased, but there’s a lot of hot brachiosaurian action on display. I’m happy to say that the other clades are not ignored–diplodocids, dicraeosaurids, titanosaurs, mamenchisaurids, basal eusauropods, and even a basal sauropodomorph are all in the mix.

Normally my brachiosaurcentricity would lead me to steal one of the numerous brachiosaur images–perhaps the awesome parade of brachiosaurs that includes both Sauroposeidon and the Archbishop (!!)–BUT my laziness led me to choose another piece by the same artist, Nima Sassani. That would be the Puertasaurus reconstruction shown at top, which includes vertebrae and thus fulfills our titular mandate. That means I can stop writing now and get back to gawking. Go do likewise.

…oh, and don’t forget to stop by Dracovenator and congratulate Adam Yates on his new critter, Aardonyx. You’ll be hearing more about Aardonyx here at SV-POW! in the hopefully not-too-distant future. I can say no more for now…

Futalognkosaurus was one big-ass sauropod

At the 2007 SVP meeting in Austin, Texas, I noticed that the suffix “-ass” was ubiquitiously used as a modifier: where an Englishman such as myself might say “This beer is very expensive”, a Texan would say “That is one expensive-ass beer” — and the disease seemed to spread by osmosis through the delegates, so that by my last day in Austin is was seemingly impossible to hear an adjective without the “-ass” suffix.

All of which is by way of introducing the fact that Futalognkosaurus really was a big-ass sauropod, as this photo of its sacrum (with articulated ilia) shows:

Articulated pelvis (sacrum and ilia) of Futalognkosaurus, in ventral view. Juan Porfiri (175 cm high) for scale. Photo by kind permission of Jorge Calvo.

A version of this photograph (in black and white and with the background chopped out) appeared in Ferdinand Novas’s recent book (Novas 2009) and attracted some discussion on the Dinosaur Mailing List.

Although in the past, we have complained about the lack of measurements in the two papers describing Futulognkosaurus (Calvo et al. 2007, 2008), this photo demonstrates a lower bound on its size: we know that it was, at least, Darned Big.  (I would attempt to calculate some measurements from this photo using Porfiri as my scale-bar, but we all know how variable human proportions are, so it’s probably better to refrain.)  The great news here is that, as explained by Ruben Juarez Valieri in a comment on an earlier article, a third article is on the way that will contain all the measurements we want.

Anyway, here are some more of Calvo’s awesome Futalognkosaurus photos, all used with grateful permission:

Median or posterior cervical vertebra of Futalognkosaurus in right anterolateral view; Juan Porfiri (175 cm) for scale. Photo by kind permission of Jorge Calvo.

(That is an insanely tall cervical.)

Articulated dorsal vertebrae of Futalognkosaurus in ?ventral view. And there is Juan Porfiri again, still 175 cm tall. Photo by kind permission of Jorge Calvo.

How on Earth did they get that jacket out the ground and back to the museum?!

And finally — if you’ll forgive the flagrant appendicularity:

Right ischium and pubis of Futalognkosaurus in ventrolateral view. Where's Juan? Photo by kind permission of Jorge Calvo.

And now for something completely different:

Open Access Week

I’m pleased to say that this week (October 19-23) is Open Access Week.  Get over to the site for statistics about the rise of open access.  Particularly impressive is a sequence of institutions that are introducing open-access mandates, i.e. requiring that all research produced by its staff is made freely available to the world.  We’re on the way!

References

• Calvo, J.O., Porfiri, J.D., Gonzalez-Riga, B.J., and Kellner, A.W.A. 2007. A new Cretaceous terrestrial ecosystem from Gondwana with the description of a new sauropod dinosaur. Anais da Academia Brasileira de Ciencias 79(3):529-541.
• Calvo, J.O., Porfiri, J.D., Gonzalez-Riga, B.J., and Kellner, A.W.A. 2008. Anatomy of Futalognkosaurus dukei Calvo, Porfiri, Gonzalez-Riga & Kellner, 2007 (Dinosauria, Titanosauridae) from the Neuquen Group (Late Cretaceous), Patagonia, Argentina. Arquivos do Museu Nacional, Rio de Janeiro 65(4):511-526.
• Novas, F. 2009. The Age of Dinosaurs in South America. Indiana University Press (Life of the Past series). 480 pages.

How big were the biggest sauropod trackmakers?

UPDATE December 3, 2009

I screwed up, seriously. Tony Thulborn writes in a comment below to correct several gross errors I made in the original post. He’s right on every count. I have no defense, and I am terribly sorry, both to Tony and to everyone who ever has or ever will read this post.

He is correct that the paper in question (Thulborn et al 1994) does discuss track length, not diameter, so my ranting about that below is not just immoderate, it’s completely undeserved. I don’t know what I was thinking. I did reread the paper before I wrote the post, but I got the two switched in my mind, and I assigned blame where none existed. In particular, it was grossly unfair of me to tar Tony’s careful work with the same brush I used to lament the confused hodgepodge of measurements reported in the media (not by scientists) for the Plagne tracks.

I am also sorry that I criticized the 1994 paper and implied that the work was incomplete. I was way out of line.

I regard this post as the most serious mistake in my professional career. I want very badly to somehow unmake it. I am adding corrections to the post below and striking out but not erasing my mistakes; they will stand as a reminder of my fallibility and a warning against being so high-handed and unfair in the future.

I’m sorry. I beg forgiveness from Tony, from all of our readers, and from the broader vertebrate paleontology community. Please forgive me.

–Mathew Wedel

You might have seen a story last week about some huge sauropod tracks discovered in Upper Jurassic deposits from the Jura plateau in France, near the town of Plagne. According to the news reports, the tracks are the largest ever discovered. Well, let’s see.

The Guardian (from which I stole the image above) says the prints are “up to 2 metres (6ft 6 in) in diameter”, but ScienceDaily says “up to 1.5 m in total diameter”. Not sure how ‘total diameter’ is different from regular diameter, but that’s science reporting for you. The BBC clarifies that, “the depressions are about 1.5m (4.9ft) wide”, which might be the key here (see below), but then mysteriously continues, “corresponding to animals that were more than 25m long and weighed about 30 tonnes.” I find it rather unlikely that a pes track 1.5 m wide indicates an animal only as big as Giraffatitan (hence this post).

So there’s some uncertainty with respect to the diameter of the tracks–half a meter of uncertainty, to be precise. But sauropod pes tracks are usually longer than wide, and a print 1.5 m wide might actually be 2 m long.

Not incidentally, Thulborn (1994) described some big sauropod tracks from the Broome Sandstone in Australia, with pes prints up to 1.5 m. Although the photos of the tracks are not as clear as one might wish, they do appear to show digit impressions and are probably not underprints. [See Tony Thulborn's comment below regarding footprints vs underprints.]

I’ll feel a lot better about the Plagne tracks when the confusion about their dimensions is cleared up and when some evidence is presented that they also are not underprints. In any case, the only dimension with any orientation cited for the Plagne tracks is the 1.5 m width reported by the BBC, so we’ll go with that. So the Plagne tracks might only tie, but not beat, Thulborn’s tracks.

…Then again, Thulborn only said that the biggest tracks were up to 150 cm in diameter. What does that mean–length? Width? Are the tracks perfect circles? Does no one who works on giant sauropod tracks know how to report measurements? These questions will have to wait, because despite the passing of a decade and a half, the world’s (possibly second-) biggest footprints–from anything! ever!–have not yet merited a follow-up paper. [Absolutely wrong and unfair; please see the apology at top and Tony Thulborn's comment below.]

Nevertheless, for the remainder of this post we’ll accept that at least some sauropods were leaving pes prints a meter and a half wide. Naturally, it occurs to me to wonder how big those sauropods were. I don’t know of any studies that attempt to rigorously estimate the size of a sauropod from its tracks or vice versa, so in the finest tradition of the internet in general and blogging in particular, I’m going to wing it.

How Big?

First we need some actual measurements of sauropod feet. When Mike and I were in Berlin last fall (gosh, almost a year ago!), we measured the feet (pedes) of the mounted Giraffatitan and Diplodocus for this very purpose. The Diplodocus feet were both 59 cm wide, and the Giraffatitan feet were 68 and 73 cm wide. The Diplodocus feet are trustworthy, the Giraffatitan bits less so. Unfortunately, the pes is the second part of the skeleton of Giraffatitan that is less well known than I would like (after the cervico-dorsal neural spines). The reconstructed feet look believable, but “believability” is hard to calibrate and probably a poor predictor of reality when working with sauropods.

One thing I won’t go into is that Giraffatitan (HM SII) probably massed more than twice what Diplodocus (CM 84/94) did, but on the other hand G. bore more of its weight on its forelimbs. It would be interesting to calculate whether the shifted center of mass would be enough to even out the pressure exerted by the hindfeet of the two animals; Don Henderson may have done this already.

Anyway, let’s say for the sake of argument that the hindfeet of the mounted Giraffatitan are sized about right. The next problem is figuring out how much soft tissue surrounded the bones. In other words, how much wider was the fleshy foot–deformed under load!–than the articulated pes skeleton? I am of two minds on this. On one hand, sauropods probaby had a big heel pad like that of elephants, and it seems reasonable that the heel pad plus the normal skin, fat, and muscle might have expanded the fleshy foot considerably beyond the edges of the bones. On the other hand, the pedal skeleton is widest across the distal ends of the phalanges, and in well-preserved tracks like the one below the fleshy foot is clearly not much wider than that (thanks, Brian, for the photo!).

Bear in mind that a liberal estimate of soft tissue will give a conservative estimate of the animal’s size, and vice versa. Looking at the AMNH track pictured above, it seems that the width added by soft tissue could possibly be as little as 5% of the width of the pes skeleton. Skewing hard in the opposite direction, an additional 20% or more does not seem unreasonable for other animals (keep in mind this would only be 10% on either side of the foot). Using those numbers, Diplodocus (CM 84/94) would have left tracks as narrow as 62 cm or as wide as 71 cm. For Giraffatitan (HM SII) I’ll use the wider of the two pes measurements, because the foot is expected to deform under load and the 73 cm wide foot looked just as believable as the 68 cm foot (for whatever that’s worth). Applying the same scale factors (1.05 and 1.20) yields a pes track width of 77-88 cm.

These numbers are like pieces of legislation, or sausages: the results are more pleasant to contemplate than the process that produced them. They’re ugly, and possibly wrong. But they give us someplace to start from in considering the possible sizes of the biggest sauropod trackmakers. Something with a hindfoot track 1.5 meters wide would be, using these numbers, conservatively more than twice as big as (2.11x) the mounted Carnegie Diplodocus or 170% the size of the mounted Berlin Giraffatitan. That’s right into Amphicoelias fragillimus/Bruhathkayosaurus territory. The diplo-Diplodocus would have been 150 feet long, and even assuming a very conservative 10 tons for Vanilla Dippy (14,000L x 0.7 kg/L = 9800 kg), would have had a mass of 94 metric tons (104 short tons). The monster Giraffatitan-like critter would have been “only” 130 feet long, but with a 14.5 meter neck and a mass of 113 metric tons (125 short tons; starting from a conservative 23 metric tons for HM SII).

Keep in mind that these are conservative estimates, for both the size of the trackmakers and the masses of the “known” critters. If we use the conservative soft tissue/liberal animal size numbers, the makers of the 1.5 meter tracks were 2.4 times as big as the mounted Diplodocus or almost twice as big as the mounted Giraffatitan, in which case masses in the blue whale range of 150-200 tons become not just probable but inevitable.

Mike measuring Giraffatitan's naughty bits. Check out the hindfeet. Also note the sauropod vertebrae in the background--titular obligation fulfilled!

Too Big?

Going the other way, I can think of only a handful of ways that the “conservative” trackmaker estimates might still be too big:

First, the pes of Giraffatitan might have been bigger than reconstructed in the mounted skeleton. Looking at the photo above, I can image a pes 10% wider that wouldn’t do any violence to the “believability” of the mount. That would make the estimated track of HM SII 10% wider and the estimated size of the HM-SII-on-steroids correspondingly smaller. But that wouldn’t affect the scaled up Diplodocus estimate, and the feet of Giraffatitan would have to be a LOT bigger than reconstructed to avoid the reality of an animal at least half again as big as HM SII.

Second, the amount of soft tissue might have been greater than even the liberal soft tissue/conservative size estimate allows. But I think that piling on 20% more soft tissue than bone is already beyond what most well-preserved tracks would justify, so I’m not worried on that score. (What scares me more is the thought that the conservative estimates are too conservative, and the real trackmakers even bigger.)

Third, I suppose it is possible that sauropod feet scaled allometrically with size and that big sauropods left disproportionately big tracks. I’m also not worried about this. For one thing, when they’ve been measured sauropod appendicular elements tend to scale isometrically, and it would be weird if feet were the undiscovered exception. For another, the allometric oversizing of the feet would have to be pronounced to make much of a dent in the estimated size of the trackmakers. I find the idea of 100-ton sauropods more palatable than the idea of 70-ton sauropods with clown shoes.

Fourth, the meta-point, what if the Broome and Plagne tracks are underprints? [Please see Tony Thulborn's comment below regarding footprints and underprints.] I’ve seen some tracks-with-undertracks where the magnification of the apparent track size in the undertracks was just staggering. The Broom tracks have gotten one brief note and The Plagne tracks have not been formally described at all, so all of this noodling around about trackmaker size could go right out the window. Mind you, I don’t have any evidence that the either set are underprints, and at least for the Broome tracks the evidence seems to go the other way, I’m just trying to cover all possible bases.

Conclusions

So. Sauropods got big. As usual, we can’t tell exactly how big. Any one individual can leave many tracks but only one skeleton, so we might expect the track record to sample the gigapods more effectively than the skeletal record. Interestingly, the largest fragmentary skeletal remains (i.e., Amphicoelias and Bruhathkayosaurus, assuming they’re legit) and the largest tracks (i.e., Plagne and Broome) point to animals of roughly the same size.

It’s also weird that some of the biggest contenders in both categories have been so little published. I mean, if I had access to Bruhathkayosaurus or a track 1.5 m wide, you can bet that I’d be dropping everything else like a bad habit until I had the gigapod evidence properly written up. What gives? [The implication that the Broome tracks were not properly written up is both wrong and unfair; please see the apology at top.]

Finally, IF the biggest fragmentary gigapods and the biggest tracks are faithful indicators of body size, they suggest that gigapods were broadly distributed in space and time (and probably phylogeny). I wonder if these were representatives of giga-taxa, or just extremely large individuals of otherwise vanilla sauropods. Your thoughts are welcome.

Epilogue: What About Breviparopus?

It’s past time someone set the record straight about damn Breviparopus. The oft-quoted track length of 115 cm is (A) much smaller than either the Broome or Plagne tracks, and (B) the combined length of the manus and pes prints together; I know, I looked it up (Dutuit and Ouazzou 1980). Why anyone would report track “length” that way is beyond me, but what is more mysterious is why anyone was taken in by it, since the width of 50 cm (pathetic!) is usually quoted along with the 115 cm “length”, indicating an animal smaller than Vanilla Diplodocus (track length is much more likely than width to get distorted by foot motions during locomotion) [This part is wrong; see the update below.]. But people keep stumbling on crap (thanks, Guiness book!) about how at 157 feet long (determined how, exactly?) Breviparopus was possibly the largest critter to walk the planet. Puh-leeze. If there’s one fact that everyone ought to know about Breviparopus, it’s that it was smaller than the big mounted sauropods at museums worldwide. The only thing super-sized about it is the cloud of ignorance, confusion, and hype that clings to the name like cheap perfume. Here’s the Wikipedia article if you want to do some much-needed revising.

UPDATE (Nov 17 2009): The width of the Breviparopus pes tracks is 90 cm, not 50 cm. The story of the 50 cm number is typically convoluted. Many thanks to Nima Sassani for doing the detective work. Rather than steal his thunder, I’ll point you to his explanation here. Point A above is still valid: Breviparopus was dinky compared to the Broome and Plagne trackmakers.

Parting Shot

You know I ain’t gonna raise the specter of a beast 1.7 times the size of HM SII without throwing in a photoshopped giant cervical. So here you go: me with C8 of Giraffatitan blown up to 170% (the vert, not me). Compare to unmodified original here.

References

• Dutuit, J.M., and A. Ouazzou. 1980. Découverte d’une piste de Dinosaure sauropode sur le site d’empreintes de Demnat (Haut-Atlas marocain). Mémoires de la Société Géologique de France, Nouvelle Série 139:95-102.
• Thulborn, R.A., T.Hamley and P.Foulkes. 1994. Preliminary report on sauropod dinosaur tracks in the Broome Sandstone (Lower Cretaceous) of Western Australia. Gaia 10:85-96.

What a 23% longer torso looks like

Just checking: no-one’s bored of brachiosaurs yet, are they?

Thought not.  Right, then, here we go!

Greg Paul’s (1988) study of the two “Brachiosaurus” species — the paper that proposed the subgenus Giraffatitan for the African species — noted that the trunk is proportionally longer in Brachiosaurus than in Giraffatitan due to the greater length of its dorsal centra. Paul (p. 7) stated that the difference is “25%-30%” on the basis of his figure 2.

Having seen the dorsal vertebrae of the type specimens of both species, my gut reaction was that the difference was nowhere near this great, so I recalculated it for myself (Taylor 2009:table 3).  Dorsal column length is the sum of the “functional length” of the centra of the dorsal vertebrae, where functional length is the length of the centrum not counting the condyle (which of course is nestled in the preceding vertebra’s cotyle when the column is articulated).  For Brachiosaurus, Riggs (1904) did not give this measurement, but did give total heights, and using these for scale I was able to measure the functional lengths from his plate LXXII.  For Giraffatitan, Janensch’s (1950:44) superbly comprehensive table supplied measurements for D4 and D8; for D11 and D12 I was able to determine the length by measuring from Janensch’s (1950:fig. 62) figure, knowing the height from his table; and for D5-D7, D9 and D10, I interpolated linearly between the measurements that I had.  Summing the functional lengths of D6-D12, I got 226 cm for Brachiosaurus and 183 cm for Giraffatitan.  So Brachiosaurus is 226/183 = 1.23 times as long as Giraffatitan — in other words, 23% longer, which is pretty much what Greg Paul said.  So I learned something there.

(Yes, brachiosaurs probably had 12 dorsals.)

So: is a 23% longer torso a big deal?  Back when I was trying to answer that question for myself, I figured it would help to take an image of a familiar animal and stretch it — so here is a horse, stolen from here and stretched:

Horse (top); and evil mutant horse with 23% longer torso (bottom).

To me, that second picture is wrong enough to hurt my eyes a little; your mileage may vary, but I suspect those among you who love horses will feel ill when you look at it.  This image was one of the reasons — one of many — that I concluded that generic separation was unavoidable.

But here’s an odd thing: tonight, for this blog post, I did the same thing to a human body, expecting it to seem even more horrible in light of how familiar we are with our own bodies.  Here it is:

Flayed Homo sapiens in orthograde anatomical position, from Vesalius (1543) "Tertia Musculorum Tabula". Modified from Wilson (2006:fig. 1). Left, as drawn; right, with torso elongated by 23%.

To my surprise, the elongated human doesn’t look appallingly wrong to me.  It doesn’t look right, of course, but it seems within the realms of, for example, what might appear as a representation of a human body in the early issues of Fantastic Four.  I am not sure what to make of that fact.  I don’t believe I have a more finely tuned sense of horse anatomy than human anatomy: it might be that I am more used to badly drawn humans than badly drawn horses; or that there is more variation in human proportions than in horse proportions; or maybe weirdness just looks less weird when it’s upright than when it’s horizontal.  I’ll be interested to hear in the comments whether the Long Horse or the Long Human looks most wrong to readers.

(By the way, I casually talk about the type specimens of both “Brachiosaurus” species: while the situation is simple in the case of Brachiosaurus altithorax, whose holotype is FMNH P25107, things are more complex in the case of Giraffatitan brancai.  Janensch nominated “Skelett S” as the holotype of his new species “Brachiosaurus” brancai, but that turned out to be a chimera, composed of the two skeletons which he subsequently designated SI and SII — but Janensch never designated one of these as the type, and so far as I’ve been able to determine, neither has anyone else done so.  SI is represented by cranial elements and the first seven cervicals, but that’s all; SII is a much larger animal and is represented by most of the skeleton, and has been informally treated as though it were the type specimen most of the while, so I formally proposed HMN SII as the lectotype of the species (Taylor 2009:788) — just a bit of housekeeping.)

Here’s our old friend, the 8th cervical vertebra of HMN II, in a rare posterodorsal aspect, showing just how thin and, well, lamina-like the spinopostzygapophyseal laminae are.  All that space in between them?  Filled with diverticula, mostly.  Amazing.

Giraffatitan brancai lectotype HMN SII, 8th cervical vertebra, in posterodorsal view

Meanwhile some good news:

Remember the good news and bad news about the all-dinosaurs special volume of The Anatomical Record?  Well, since we posted that, the entire issue has been made open access!  Fantastic stuff there: details from D. Schachne of the Wiley-Blackwell Communications Team.  It’s not clear why the articles were all paywalled when originally posted, but all’s well that ends well.

And finally …

There’s been a gratifying amount of discussion in the comments on recent articles.  It can be hard to keep track of, but it helped a lot when I found an RSS feed for comments, which is what I now use.  For anyone else who wants it, it’s at http://svpow.wordpress.com/comments/feed/

References

• Janensch, Werner.  1950.  Die Wirbelsaule von Brachiosaurus brancai.  Palaeontographica (Suppl. 7) 3: 27-93.
  Paul, Gregory S.  1988.  The brachiosaur giants of the Morrison and Tendaguru with a description of a new subgenus, Giraffatitan, and a comparison of the world’s largest dinosaurs.  Hunteria 2 (3): 1-14.
  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.
  Vesalius, A.  1543.  Andreae Vesalii Bruxellensis, Scholae medicorum Patauinae professoris, de Humani corporis fabrica Libri septem [facsimile]. Ex Officina Ioannis Oporini, Basel, 659 pp.
  Wilson, Jeffrey A.  2006.  Anatomical nomenclature of fossil vertebrates: standardized terms or “lingua franca”?  Journal of Vertebrate Paleontology 26(3): 511-518.
• Janensch, Werner.  1950.  Die Wirbelsaule von Brachiosaurus brancai.  Palaeontographica (Suppl. 7) 3: 27-93.
• Paul, Gregory S.  1988.  The brachiosaur giants of the Morrison and Tendaguru with a description of a new subgenus, Giraffatitan, and a comparison of the world’s largest dinosaurs.  Hunteria 2 (3): 1-14.
• 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.
• Vesalius, A.  1543.  Andreae Vesalii Bruxellensis, Scholae medicorum Patauinae professoris, de Humani corporis fabrica Libri septem [facsimile]. Ex Officina Ioannis Oporini, Basel, 659 pp.
• Wilson, Jeffrey A.  2006.  Anatomical nomenclature of fossil vertebrates: standardized terms or “lingua franca”?  Journal of Vertebrate Paleontology 26(3): 511-518.

Bifid Brachiosaurs, Batman!

These are the days of miracle and wonder, especially for all you right-minded people out there who are lovers of fine brachiosaurs.  I heard yesterday evening about a new paper in Proceedings of the Royal Society B: You and Li’s (2009, duh) description of a new brachiosaur, the first one known from the Cretaceous of Asia: Qiaowanlong kangxii. Best of all, it’s based primarily on vertebral material:

You and Li (2009:fig. 2) Cervical vertebrae of Qiaowanlong kangxii holotype FRDC GJ 07-14. (a) Photograph and (b) interpretative line drawing of C4-C7 in left lateral view; (c) a distal portion of a cervical rib; C9 in (d) cranial, (e) left lateral, (f) caudal, (g) right lateral, (h) dorsal and (i) ventral views. di, diapophysis; f1-f5, fossa 1-fossa 5; pa, parapophysis; poz, postzygapophysis; prz, prezygapophysis; sp, neural spine. Scale bars, 10 cm.

Brachiosaur aficionados will be gazing slack-jawed at parts d, f and h of this figure (the anterior, posterior and dorsal views of C9), which clearly show that the neural spines of the new taxon are bifid (i.e. have two peaks side by side and a trough between them) — just like the cervical neural spines of flagellicaudatans (diplodocids and dicraeosaurs) and camarasaurs.  And mamenchisaurs.  And some titanosaurs.  And Erketu.  Finding this feature yet again — apparently independently evolved in brachiosaurs — makes it about the most plastic character in the matrix.  Very exciting.

That is, it’s exciting if this really is a brachiosaurid.  Now as it happens, Matt was one of the reviewers for this paper (and by the way did an amazingly professional job of not telling me about it until it came out, the git).  He’s told me in email that he’s satisfied that Qiaowanlong really is a brachiosaur, and I hesitate to question that identification given that (A) unlike the authors I’ve never seen the material, and (B) unlike Matt, I’ve spent most of my brachiosaur-presacral quality time with dorsals rather than cervicals.  But, with that caveat, I’m not sure that a compelling case has yet been made for a brachiosaurian identity.

The authors cite three characters in support of a brachiosaurid identity:

• The most persuasive is the deeply excavated cervical neural spines.
• Next is a transition in neural spine height: this is quite abrupt in “Brachiosaurus” brancai between cervicals 6 and 7, and also in Sauroposeidon — presumably also between C6 and C7, but that can’t be known for sure, since it’s only the assumption that this is the case that led to the identification of the four preserved Sauroposeidon cervicals as C5-C8 in the first place.  In Qiaowanlong, this transition is “much less pronounced”, with spines increasing in height by only 25% rather then 100% in the other taxa — and occurs between C8 and C9.  All in all, not really very similar to the condition in “B.” brancai.
• The final character supporting the brachiosaurid identity of Qiaowanlong is the absence of an anterior centrodiapophyseal lamina.  As the authors point out, though, this lamina does exist in “B.” brancai and is absent only in Sauroposeidon; so if this is evidence of anything, it’s a synapomorphy of a clade uniting Qiaowanlong and Sauroposeidon to the absence of other brachiosaurs — something that seems very unlikely given the proportions of the vertebrae.

Putting it all together, there seems to be only one convincing brachiosaur character cited; and that stands against several non-brachiosaur characters, most obviously the bifurcation of the neural spine and the low Elongation Index (centrum length divided by cotyle height) but also by a few other characters that are not discussed in the paper.  For example, Matt has previously noted that in brachiosaur cervicals, the diapophyses are more anteriorly positioned than the parapophyses whereas in diplodocids the opposite is the case: as shown in fig 2(b) above, C6 at least of Qiaowanlong resembles diplodocids in this respect.

To try to get more of a handle on this, I put together a comparative figure of the 8th and 9th cervicals of various sauropods:

8th/9th cervicals vertebrae of various sauropods, scaled to the same centrum length. From top to bottom and left to right: "Brachiosaurus" brancai, Sauroposeidon; Qiaowanlong, Diplodocus; Haplocanthosaurus, Camarasaurus. Six sauropod vertebrae for the price of one!

Based on overall proportions, I don’t find it intuitively obvious that the Qiaowanlong (middle row, left) more closely resembles the brachiosaurs (top row) than it does the other three.

What does all this mean?  Probably nothing: most likely there are further reasons for the brachiosaurid identification of the new taxon, and lack of space prevented their explanation and illustration.  We can hope that the authors, having placed an initial brief description in Proc. B, will follow it up with a more comprehensive description and analysis in a journal that does not impose such tight length restrictions.  But for now at least, my feeling is that the case for a bifid brachiosaur has yet to be made.

Moving on … Qiaowanlong is also represented by some nice appendicular material: the entire right side of the pelvis (ilium, ischium and pubis).  The ilium certainly looks brachiosaury, so that is another bit of support for the systematic hypothesis, but the proportions of the pelvic bones are very odd:

Right pelvis of "Brachiosaurus" brancai (left), based on composite of Janensch's (1961) figures, and Qiaowanlong (from You and Li 2009: fig. 3a). Scaled to same ilium length.

You and Li (2009) describe their pelvis as having a “much reduced ischium”, but as is apparent by comparison with the pelvis of “Brachiosaurus” brancai, the ischium is in reasonable proportion to the ilium, and the oddity is more that the pubis is enormous.  So much so that it makes me feel a little ill looking at it, and it makes me wonder how certain it is that all three of these bones are from the same individual — sadly, the paper doesn’t discuss the association of the material.

[Not to flog a dead horse, but this kind of omission shows once more the perils of publishing new taxa in general-interest journals such as Proc. B that impose draconian length limits.  This paper just creeps onto page 7, and I simply don't believe that it's possible to do anything like justice to the description of a new taxon in that little space, especially when there is also geography, geology, phylogeny and discussion to be got through.  I don't want to go all This Is How To Do It, but I can't help remembering that Darren and I took 18 pages, nearly three times as long, to describe the single partial vertebra that is Xenoposeidon (Taylor and Naish 2007), and it's not as though that paper wastes a lot of words.  To give You and Li credit, they did squeeze in photos of a representative vertebra from all six cardinal directions, which is great; but only at the cost of the photos being too tiny to be much use.  Please, folks: send your new taxon descriptions to a proper descriptive journal, not to a tabloid!  </hobbyhorse>]

Back on the Dinosaur Mailing List, B tH asked how big Qiaowanlong was.  According to the BBC, the authors say that “the dinosaur would have been a relatively small sauropod about 12m long, 3m high, and weighing perhaps 10 tonnes”.  Can we confirm that?  Well, the excellently comprehensive table of measurements in the paper gives centrum lengths, not counting the condyle, totalling 267 cm for the seven vertebrae C5-C11.  Janensch (1950a:44) gave measurements for the corresponding vertebrae of “Brachiosaurus” brancai HMN SII totalling 577 cm, which is more than twice as long.  If Qiaowanlong was 267/577 = 0.46 times as long as HMN SII, which Janensch (1950b:102) gave as 22.46 m, then it would have been 10.4 m long; it’s not obvious how the authors got the larger figure of 12 m unless they had reason to think the neck was proportionally shorter than in HMN SII.  If Qiaowanlong was isometrically similar to HMN SII, then it was 0.46^3 = 0.99 0.099 times as heavy.  Using my own in-press mass of 23337 kg for HMN SII, this would make Qiaowanlong only 2312 kg in mass — pretty pathetic for a sauropod.

That’s it for now.  I’d be the first to admit that there’s an awful lot of speculation in this post based on relatively little published information.  Hopefully You Hai-Lu will drop by and comment — I’ll be letting him know that I’ve posted this.

References

• Janensch, Werner.  1950.  Die Wirbelsaule von Brachiosaurus brancai. Palaeontographica (Suppl. 7) 3: 27-93.
  Janensch, Werner.  1950.  Die Skelettrekonstruktion von Brachiosaurus brancai.  Palaeontographica (Suppl. 7) 3: 97-103.
  Janensch, Werner.  1961.  Die Gliedmaszen und Gliedmaszengurtel der Sauropoden der Tendaguru-Schichten.  Palaeontographica, suppl. 7 (1), teil 3, lief. 4: 177-235.
  Taylor, Michael P. and Darren Naish.  2007.  An unusual new neosauropod dinosaur from the Lower Cretaceous Hastings Beds Group of East Sussex, England.  Palaeontology 50 (6): 1547-1564.  doi: 10.1111/j.1475-4983.2007.00728.x
  You, Hai-Lu, and Li, Da-Qing.  2009.  The first well-preserved Early Cretaceous brachiosaurid dinosaur in Asia.  Proceedings of the Royal Society B: Biological Sciences.  doi: 10.1098/rspb.2009.1278.
• Janensch, Werner.  1950.  Die Wirbelsaule von Brachiosaurus brancai. Palaeontographica (Suppl. 7) 3: 27-93.
• Janensch, Werner.  1950.  Die Skelettrekonstruktion von Brachiosaurus brancai.  Palaeontographica (Suppl. 7) 3: 97-103.
• Janensch, Werner.  1961.  Die Gliedmaszen und Gliedmaszengurtel der Sauropoden der Tendaguru-Schichten.  Palaeontographica, suppl. 7 (1), teil 3, lief. 4: 177-235.
• Taylor, Michael P. and Darren Naish.  2007.  An unusual new neosauropod dinosaur from the Lower Cretaceous Hastings Beds Group of East Sussex, England.  Palaeontology 50 (6): 1547-1564.  doi: 10.1111/j.1475-4983.2007.00728.x
• You, Hai-Lu, and Li, Da-Qing.  2009.  The first well-preserved Early Cretaceous brachiosaurid dinosaur in Asia.  Proceedings of the Royal Society B: Biological Sciences.  doi: 10.1098/rspb.2009.1278.

And finally … two announcements!

Traumador the Tyrannosaur has asked us to point out that over on ART Evolved (the palaeo-art blog), the next big art gallery is to be sauropod themed.  Details are on the site, so get over there and submit your sauropod art!

And Matt and I will shortly be teaming up with Andy Farke, the open-source paleontologist, on a new project where we plan to actually do some of this Shiny Digital Future that we keep on talking about.  Andy will be announcing the details on Tuesday 8th September.  Mark the date well!  For now, I shall say no more …

How big was Alamosaurus?

Here’s a skeletal reconstruction of Alamosaurus modified from Lehman and Coulson (2002:fig. 11). I cloned the neck and rotated it a few degrees to see where it would put the head.

The skeleton in the figure is scaled to the size of the individuals in the Smithsonian and at UT Austin. The scale bar is 1 meter, which by my calculations gives that individual the following dimensions:

• Total length: 15.8 meters (52 feet)
• Neck length: 5.2 meters (17 feet)
• Shoulder height: 4 meters (13 feet)
• Head height (with neck raised): 8.4 meters (27.5 feet)

Here are a couple of articles on a giant sauropod found in Big Bend in 1999. This critter is generally assumed to be Alamosaurus but it could be something new (I have no evidence either way); the material is currently under study at the Dallas Museum of Nature and Science.
http://www.nps.gov/bibe/naturescience/alamosaurus.htm
http://www.geocities.com/stegob/texasdino.html

According to the articles, 10 cervical vertebrae were found in a string 23 feet long. From the pictures, those ten vertebrae look like the ten largest, which should account for almost all of the neck except for the first few cervicals behind the head. Let’s assume that this big individual therefore had a neck just a little longer than 23 feet, and we find that it is almost exactly 1.5 times bigger than the one listed above. If its proportions follow those of the Lehman and Coulson recon, its measurements would be:

• Total length: 24 meters (79 feet)
• Neck length: 7.8 meters (25.5 feet)
• Shoulder height: 6 meters (19.5 feet)
• Head height: 12.6 meters (41 feet)

In the second article Homer Montgomery speculates that the complete neck would have been more than 30 feet long. That’s certainly not impossible, since 30-foot-plus necks are known for the largest individuals in several clades (e.g., Mamenchisaurus, Supersaurus, Sauroposeidon, probably Puertasaurus, possibly Futalognkosaurus, but probably not Aegyptosaurus) If so, then you could just about double all of the proportions from the first individual described above, which would give a truly prodigious animal. The 52-foot animal probably had a mass around 15 tons, so the 79-footer would have been about 50 tons (1.5^3 = 3.375), and the hypothetical 100-footer would have been 120 tons, which is up in Amphicoelias/Bruhathkayosaurus territory. For what it’s worth, I think the numbers for the 79-foot animal are more plausible, but who knows. Anytime you’ve got a partial neck that is longer than the complete neck of Diplodocus, you’re dealing with a wacky big animal.

Reference

Lehman, T.M. & Coulson, A.B. 2002. A juvenile specimen of the sauropod Alamosaurus sanjuanensis from the Upper Cretaceous of Big Bend National Park, Texas. Journal of Paleontology 76(1): 156-172.

How tallweird was Sauroposeidon?

In an email, Vladimir Socha drew my attention to the fact that Tom Holtz’s dinosaur encyclopaedia estimates the maximum height of Sauroposeidon as 20 meters plus, and asked whether that was really possible.  Here’s what Tom actually wrote: “Sauroposeidon was one of the largest of all dinosaurs.  At perhaps 98 to 107 feet (30 to 32.5 meters) long and weighing 70 to 80 tons [...] Sauroposeidon would have been the tallest of all dinosaurs. [...] If it could crane its neck up, it might have been able to hold its head 66 to 69 feet (20 to 21 meters) high or more” (Holtz and Rey 2007:207).  Vladimir was understandably skeptical.  But can it be true?

Wedel and Cifelli (2005: fig. 15) shows Matt’s best skeletal reconstruction of Sauroposeidon, with Boring Old Brachiosaurus and a human for scale:

Sauroposeidon with Boring Old Brachiosaurus and human for scale and 20 m height indicated. Lightly modified from Wedel and Cifelli (2005: fig. 15)

Amazingly, those dummies didn’t include an actual scalebar; but apparently the human figure is 1.8 m tall, so by measuring pixels and cross-scaling, I determined that in this image, Sauroposeidon is a mere 13.43 m tall.  I took the liberty of adding in a marker for the 20 m height proposed by Holtz, and as things stand you’d have to say that it doesn’t look probable.

But let’s see what we can do.  We’ll begin with the classic brachiosaur skeleton of Paul (1988), which shows the well represented species Brachioaurus brancai:

Brachiosaurus brancai skeletal reconstruction in left lateral view. From Paul (1988:fig. 1)

(Some other time, we should take a moment to discuss the differences between this and the Wedel brachiosaur reconstruction; but it will not be today.)

This reconstruction is in a nice erect-necked posture which, in light of our own recent paper, is probably not too extreme.  Since all the neural arches and processes are missing from the only known posterior cervicals of this species, we don’t know how much flexibility they allowed, and so in light of how the rest of the animal is built (high shoulders and all) it seems reasonable to allow a lot of extension at the base of the neck.  So let’s assume that the pose offered by Paul is correct.  By measuring my scan of that figure, and I see that the 2.13 m humerus is 306 pixels long.  The entire reconstruction, from tip of cranial crest down to forefoot, is 1999 pixels tall, which is 1999/306 = 6.53 times as long as the humerus, which scales to 6.53*2.13 = 13.91 m — a little taller than Sauroposeidon (not Brachiosaurus) in Matt’s reconstruction, which seems about right if we imgine Matt’s Brachiosaurus raising its neck into a Paul-compliant posture.

Now Paul’s reconstruction is based on the Berlin mounted skeleton HMN S II.  Cervical 8 is very well preserved in that animal, and has a centrum length of 98 cm (Janensch 1950a:44).  By contrast, the centrum of C8 of Sauroposeidon OMNH 53062 (the only known specimen) is 125 cm long (Wedel et al. 2000a: 110). So if Sauroposeidon is merely an elongated Brachiosaurus brancai, then it’s 125/98 = 1.28 times as long and tall, which would be 17.74 m.

But wait: it seems that Sauroposeidon is to Brachiosaurus brancai as Barosaurus is to Diplodocus — similar overall but more elongate.  And it turns out that Barosaurus has at least 16, maybe 17 cervicals (McIntosh 2005:45) compared with Diplodocus’s 15.  So maybe Sauroposeidon also added cervicals from the brachiosaur base-state — in fact, that would hardly be surprising given that Brachiosaurus brancai has so few cervicals for a long-neck: 13, compared with 15 in most diplodocids, 16 or 17 in Barosaurus, and 19 in Mamenchisaurus.  If you reconstruct Sauroposeidon with two more C8-like cervicals in the middle of the neck, that adds 2*125 = 250 cm, which would give us a total height of 17.74+2.5 = 20.24 m.

So I don’t think Tom Holtz’s estimate is completely unrealistic, even for the one Sauroposeidon specimen we have now — and remember that the chances of that individual being the biggest that species got are vanishingly small.

On the other hand, maybe Sauropodseidon’s neck was the only part of it that was elongated in comparison to Brachiosaurus brancai — maybe its forelimbs were no longer than those of its cousin, so that only the neck elongation contributed to greater height.  And maybe it had no additional cervicals, so its neck was “only” 1.28 times as long as that of Brachiosaurus brancai — 1.28*8.5 = 10.88 m, which is 2.38 m longer; so the total height would be 13.91+2.38 = 16.29 m (assuming the additional neck length was vertical).  And maybe the neck couldn’t get very close to vertical, so that the true height was lower still.

All of this just goes to show the perils of reconstructing an animal based only on a sequence of four cervicals.  (Reconstructing on the basis of a single partial mid-to-posterior dorsal, on the other hand, is a much more exact science.)

Finally: Matt’s reconstruction of Sauroposeidon is really rather conservative, and looks very much like a scaled-up vanilla brachiosaur.  Just to see how it looks, I’ve made a reconstruction of the putative (and very possible) elongated, attenuated version of Sauroposeidon, showing the legs and cervicals 28% longer than in B. brancai, and with two additional cervicals.  I made this by subjecting Greg Paul’s 1988 brachiosaur to all sorts of horrible and half-arsed distortions, so apologies to Greg.  But remember, folks: this is just as likely correct as Matt’s version!

A different view of Sauroposeidon, based on elongation of the cervicals and legs of Brachiosaurus brancai and the insertion of two additional cervicals. Heavily and carelessly modified from Paul (1988: fig. 1)

References

• Holtz, Thomas R., Jr., and Luis Rey.  2007.  Dinosaurs: The Most Complete, Up-to-Date Encyclopedia for Dinosaur Lovers of All Ages.  Random House, New York.  428 pages.
• Janensch, Werenr.  1950.  Die Wirbelsaule von Brachiosaurus brancai.  Palaeontographica (Suppl. 7) 3: 27-93.
• McIntosh, John S.  2005.  The Genus Barosaurus Marsh (Sauropoda, Diplodocidae).  pp. 38-77 in Virginia Tidwell and Ken Carpenter (eds.), Thunder Lizards: the Sauropodomorph Dinosaurs.  Indiana University Press, Bloomington, Indiana.  495 pages.
• Paul, Gregory S.  1988.  The brachiosaur giants of the Morrison and Tendaguru with a description of a new subgenus, Giraffatitan, and a comparison of the world’s largest dinosaurs.  Hunteria 2 (3): 1-14.
• Wedel, Mathew J., and Richard L. Cifelli.  2005.  Sauroposeidon: Oklahoma’s Native Giant.  Oklahoma Geology Notes 65 (2): 40-57.
• Wedel, Mathew J., Richard L. Cifelli and R. Kent Sanders.  2000a.  Sauroposeidon proteles, a new sauropod from the Early Cretaceous of Oklahoma.  Journal of Vertebrate Paleontology 20(1): 109-114.
• 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.

.

Little, big: the reveal

Here’s the answer to last week’s riddle. The big vertebra was obviously cervical 8 of Sauroposeidon, which you’ve seen here more than once. The small vertebra is also a mid-cervical, also from the Early Cretaceous, but from Croatia rather than Oklahoma. The very long centrum, unbifurcated neural spine, and extensive pneumatic sculpturing mark it as a brachiosaurid. It was first described by Dalla Vecchia (1998), and lavishly illustrated with numerous photos by Dalla Vecchia (1999). It was also included by Dalla Vecchia (2005:figs. 18.5 and 18.6) in the Thunder-Lizards volume from Indiana University Press, which is where I figured someone might recognize it from.

Here are two of those figures from Dalla Vecchia (1999)–note the thumb and fingers in the left-hand photo. The vertebra is about a foot long (~30 cm), which means it is TINY for a brachiosaurid mid-cervical. Note also that there is no sign of a neurocentral suture, so the critter was probably at least half grown and might have been full grown.

It is worth bearing mind that this super-tiny, pathetically titchy, adorable widdle bwachiosauw ve’tebwa is only a bit smaller than your average giraffe cervical.

Speaking of giraffes, from left to right we have:

• Sauroposeidon, scaled like HM SII x 1.15;
• a 20-foot-tall world record giraffe;
• WNV-1, scaled like 0.22 x Sauroposeidon;
• a 6′2″ human, such as yours truly.

Note that I  could look over the shoulder of WNV-1, but it could not look over the giraffe’s shoulder, nor could the giraffe look over Sauroposeidon’s shoulder. The giraffe could not walk under Sauroposeidon’s stomach, but WNV-1 could walk under the giraffe’s.  If the mass of Sauroposeidon was 40 tons, that of WNV-1 may have been around 450 kg, or a little under half a ton.

I wonder which evolved first in brachiosaurids, stupendous size or stupendous necks?

References

• Dalla Vecchia, F.M. 1998. Remains of Sauropoda (Reptilia, Saurischia) in the Lower Cretaceous (Upper Hauterivian/Lower Barremian) limestones of SW Istria (Croatia). Geologia Croatica 51(2):105-134.
• 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.
• Dalla Vecchia, F.M. 2005. Between Gondwana and Laurasia: Cretaceous sauropods in an intraoceanic carbonate platform; pp. 395-429 in Tidwell, V., and Carpenter, K. (eds.), Thunder-Lizards: The Sauropodomorph Dinosaurs. Indiana University Press, Bloomington.

Sauropods were corn-on-the-cob, not shish kebabs

This is corn on the cob:

Corn on the cob, in cross section. Stolen from http://www.istockphoto.com/file_thumbview_approve/214165/2/istockphoto_214165-co rn-cob-cross-section.jpg

This is a shish kebab:

Shish kebab. Stolen from http://www.mediterraneancafe-flatiron.com/images/shish.jpg

Most tetrapods are like shish kebabs: a whole lot of meat stuck on a proportionally tiny skeleton.  If you don’t believe me, you can look at the human and cow neck torso cross-sections in Matt’s last post, or check out this ostrich-neck cross-section from his 2003 Paleobiology paper:

Ostrich neck in cross section, CT scan. From Wedel (2003a: fig. 2)

Remember that this is a freakin’ ostrich — of all extant animals, one of the ones with a most extreme long, skinny neck.  And yet, if sauropods were muscled like ostriches, then their necks would have looked like this in cross section:

Putative shish kebab-style sauropod neck in cross section. Ostrich soft-tissue from Wedel (2003a: fig. 2), Diplodocus vertebra cross-section from Paul (1997: fig. 4) scaled to match size of ostrich vertebra

And soft-tissue reconstructions would have to look like this:

Diplodocus with its neck as fat as an ostrich's. Modified from Paul (1998: fig. 1F)

Which, happily, no-one is suggesting.  Instead, published reconstructions of sauropod neck soft-tissue are startlingly emaciated.  As exhibit A, I call this pair of Greg Paul cross-sections:

Diplodocus and Brachiosaurus neck cross-sections, showing very light musculature. From Paul (1997: fig. 4)

(Yes, the Diplodocus on the left is the one I used in the photoshopped ostrich cross-section above.  It’s instructive to compare Paul’s original with the What If It Was Like A Big Ostrich version.)

Paul’s reconstructions seem to be widely considered too lightly muscled.  But even the very careful and rigorous more recent reconstructions of Daniela Schwarz and her colleague show a neck much, much thinner than that of the ostrich:

Diplodocus neck cross-sections. From Schwarz et al. (2007: fig. 7a)

Although Schwarz has put a lot more soft tissue onto the neck vertebrae than Paul did, it is still a tiny proportion of what we see in extant animals — even the ostrich, remember, which has a super-thin neck compared with pretty much anything else alive today.  If sauropod necks were muscled as heavily as those of, say, cows, then the soft tissue would pretty much reach down to the ground.  But they weren’t: they were more like corn on the cob, with a broad core of skeleton and relatively little in the way of delicious edibles festooned about it.

So why is this?  Why does everyone agree that sauropod necks were much less heavily muscled than those of any extant animal?

It’s a simple matter of scaling.  A really big ostrich might have a neck 1 m long.  (Actually, ostriches don’t get that big, but let’s pretend they do because it makes the maths easier).  If the x meter-long neck of a sauropod was just a scaled-up ostrich neck, then it would be x times longer, x times taller and x times wider, for a total of x^3 times as voluminous and therefore x^3 times as heavy.  But the cross-sectional area of the tension members that support it is only x times taller and x times wider, for a total of x^2 times the strength.  In total, then, the neck’s mass/strength is x^3/x^2 = x times as great as in the ostrich.  (The sauropod neck’s mass also acts further out from the fulcrum by an additional factor of x, but that is cancelled by the fact that the tension in the neck also acts x times higher above the fulcrum.)

It seems intuitively obvious (which is is code for “I have no way to prove”) that you can’t reasonably expect the neck muscles of a giant ostrich to work ten times as hard as they do in their lesser cousins, which is what you’d need to do for the 10 m neck of, say, Sauroposeidon.  So simple isometric scaling won’t get the job done, and you need to restructure the neck.

But how?  Surely just reducing all the muscle around the vertebrae can’t help?  No indeed — but that is not really what sauropods were doing.  If you look at the typical sauropod-neck life restoration, you’ll see that the proportional thickness of the neck is actually not too dissimilar to that of an ostrich — rather thicker, in fact.  If you scaled an ostrich neck up to sauropod size and compared it with a real sauropod neck, you would find not that the soft tissue was too fat, but that the vertebrae were too thin.

And so we come to it at last: rather than thinking of sauropods as having reduced the amount of soft-tissue hanging on the cervical vertebrae, we do better to think of them as having kept a roughly similar soft-tissue profile to that of an an ostrich, but enlarging the vertebrae within the soft-tissue envelope.  Of course if you just blindly made the vertebrae taller and wider, they would become heavier in proportion, which would defeat the whole purpose of the exercise — but as everyone who reads this blog surely knows by now, sauropod cervicals were extensively lightened by pneumaticity.  By bringing air into the center of the neck, they were effectively able to displace bone, muscle and ligament away from the centre, so that they acted with greater mechanical advantage: higher epaxial tension members, lower hypaxial compression members, and more laterally positioned paraxials.

It’s a rather brilliant system — using the same volume of bone to achieve greater strength by displacing it outwards and filling the center with air (and, in doing so, also displacing soft tissue outwards).  And it will be hauntingly familiar to anyone who loves birds, because it is of course exactly what birds (and pterosaurus) have done in their long bones: the hollow humeri of flying vertebrates famously allow them to attain greater strength — specifically, resistance to bending — for the same volume and mass of bone.  It’s a neat trick when done with long bones, but it takes a truly awesome taxon to do it with the neck.

So maybe sauropods were not corn on the cob after all.  Maybe they were Hostess Twinkies.

Hostess Twinkie. Not truly pneumatic, as the internal cavity is filled with adipose tissue rather than air, but do you have any idea how difficult it is to find good images of hollow junk food? Stolen from http://dixiedining.files.wordpress.com/2008/07/twinkie_070918_ms1.jpg

And now for something completely different

Now that I’ve finished my Ph.D at the University of Portsmouth, what am I going to do with the rest of my scientific life?  I’ve always said that I have no intention of going into palaeo full time: my knowledge is far too narrow for that, so that even if paid jobs were not in insanely short supply, I wouldn’t stand much chance of getting one.  And in any case, I’d hate to get into the all-too-common situation of being up against a friend for a position we both wanted. Throw in the fact that I really enjoy my computer-programming day-job and it seems pretty clear that what I need is an unpaid affiliation that lets me get on with lovely research.

Well: I am absolutely delighted to announce that, as of last month, I am an Honorary Research Associate in the Department of Earth Sciences at UCL.  It’s not just that UCL is such a well-respected institution — see that Wikipedia article for some details — more importantly, it’s where Paul Upchurch hangs out, as Senior Lecturer in Palaeobiology.  Sauropod fans will be familiar with Paul’s characteristically detailed and careful work, from his pioneering work on sauropod phylogeny (Upchurch 1995, 1998), through his and John Martin’s indispensible Cetiosaurus makeovers (Upchurch and Martin 2002, 2003) to the state-of-the art review that he lead-authored for Dinosauria II (Upchurch et al. 2004) and the Tokyo Apatosaurus monograph (Upchurch et al. 2005).  What many of you won’t know is what an excellent collaborator he is — quick, conscientious, insightful and diplomatic.  We’ve already collaborated on a few short papers (Upchurch et al. 2009 and a couple of Phylocode companion-volume chapters that are in press), and I hope there will be more in the future.

References

• Paul, Gregory S.  1997.  Dinosaur models: the good, the bad, and using them to estimate the mass of dinosaurs.  pp. 129-154 in: D. L. Wolberg, E. Stump, and G. D. Rosenberg (eds.), DinoFest International: Proceedings of a Symposium Sponsored by Arizona State University.  Academy of Natural Sciences, Philadelphia.
• Paul, Gregory S.  1998.  Terramegathermy and Cope’s Rule in the land of titans.  Modern Geology 23: 179-217.
• Schwarz, Daniela, Eberhard Frey and Christian A. Meyer.  2007. Pneumaticity and soft-tissue reconstructions in the neck of diplodocid and dicraeosaurid sauropods.  Acta Palaeontologica Polonica 52(1): 167-188.
• Upchurch, Paul.  1995.  The evolutionary history of sauropod dinosaurs.  Philosophical Transactions of the Royal Society of London Series B, 349: 365-390.
• Upchurch, Paul.  1998.  The phylogenetic relationships of sauropod dinosaurs.  Zoological Journal of the Linnean Society, 124: 43-103.
• Upchurch, Paul and John Martin.  2002.  The Rutland Cetiosaurus: the anatomy and relationships of a Middle Jurassic British sauropod dinosaur.  Palaeontology, 45(6): 1049-1074.
• Upchurch, Paul and John Martin.  2003.  The anatomy and taxonomy of Cetiosaurus (Saurischia, Suaropoda) from the Middle Jurassic of England.  Journal of Vertebrate Paleontology 23(1): 208-231.
• Upchurch, Paul, Paul M. Barrett and Peter Dodson.  2004.  Sauropoda. pp. 259-322 in D. B. Weishampel, P. Dodson and H. Osmólska (eds.), The Dinosauria, 2nd edition.  University of California Press, Berkeley and Los Angeles.  861 pp.
• Upchurch, Paul, Yukimitsu Tomida, and Paul M. Barrett.  2005.  A new specimen of Apatosaurus ajax (Sauropoda: Diplodocidae) from the Morrison Formation (Upper Jurassic) of Wyoming, USA.  National Science Museum Monographs No. 26.  Tokyo.  ISSN 1342-9574.
• Upchurch, Paul, John Martin, and Michael P. Taylor.  2009.  Case 3472: Cetiosaurus Owen, 1841 (Dinosauria, Sauropoda): proposed conservation of usage by designation of Cetiosaurus oxoniensis Phillips, 1871 as the type species.  Bulletin of Zoological Nomenclature 66(1): 51-55.
• Wedel, Mathew J.  2003.  Vertebral pneumaticity, air sacs, and the physiology of sauropod dinosaurs.  Paleobiology 29(2): 243-255.

Neck posture, yet again: T. rex’s neck is pathetic

Here at SV-POW! Towers, we often like to play Spot The T. rex — a simple drinking game that can be played whenever you have supply of palaeontology-related news reports.  Each player in turn takes a report off the stack, and if T. rex is mentioned anywhere in the report, the player drinks.  We lay in a lot of beer when we play this game, because as it turns out, T. rex is nearly always mentioned (and nearly always spelled “T-Rex”, no italics, no full stop, gratuitous hyphen, capitalised trivial name).  For example, suppose someone publishes an innocent paper arguing that a particular Eocene clam was an obligate scavenger: then the story in the press will be “… just as has been argued for the terrifying T-Rex, which had teeth like steak knives”.  Or if someone names a new Miocene rodent, it will be introduced as “… which lived 50 million years after the terrifying T-Rex, which had teeth like steak knives”.   (Drink twice if the steak knives are mentioned.  Three times if they are described as “banana-sized”.)

So we didn’t feel our neck-posture paper was real until it had somehow been tied in with T-Rex.  Happily, the Great North Museum came to the rescue: by coincidence, they unveiled their T. rex cast the weekend before the paper came out, and the Sunday Sun wanted our opinion on the way the neck had been mounted.  Here’s their mount (not quite ready to exhibit):

Tyrannosaurus rex mounted skeleton at the Great North Museum. From journallive.co.uk

Of course, everything we said about the necks of sauropods in the paper also applies to every other extinct land vertebrate — we only concentrated on sauropods because (A) they are the group whose neck posture has been claimed to depart from the tetrapod norm, and (B) they are cool.  In particular, non-avian theropods such as T. rex are in the same extant phylogenetic bracket as sauropods are (i.e. birds plus crocs), so we’d expect strong extension at the base of the neck and strong flexion at the head joint in habitual pose.

So I replied that “the Newcastle mount has the neck and torso in more of a straight line [than a Vidal-compliant posture], which would probably not have been the habitual pose.  It looks to me as though this animal is crouching down to take a drink”, and I’m pleased that the resulting news story included a rather gracious response from the GNM curator.

I don’t know whether the notoriously litigious Disney corporation would be so mellow, though, regarding their truly horrible mount of a cast of “Sue”:

Tyrannosaurus rex "Sue" cast, at Animal Kingdom, Walt Disney World, Florida. From wwarby's Flickr photostream.

I’m really not sure what the people who mounted this were getting at: unlike the Great North Museum mount, the legs are erect, so it’s not going into or coming out of a crouch; and it’s not going into a drinking posture, because the head is pointing straight forward.  But for some reason, it’s below shoulder height.

Here’s how it should be done:

Tyrannosaurus rex at the American Museum of Natural History. Photo by Mike Taylor

It’s good to see that the biggest natural history musuem in the world is ahead of the curve, and has its T. rex mount in a pose consistent with how other land vertebrates habitually hold their necks.

I leave you with the news the T. rex’s neck is pathetic.  Here is the skull and neck of that same AMNH mount, composited with a single cervical vertebra (C8) of Sauroposeidon.  Please note that the Sauroposeidon cervical is way longer than the whole T. rex neck.

T. rex's neck is pathetic

No references today!

[You don't need to be told the reference for Taylor et al. (2009) again, do you?]