After a completely barren 2008, this year is turning out to be a good one for me in terms of publications.  Today sees the publication of Taylor (2009b), entitled Electronic publication of nomenclatural acts is inevitable, and will be accepted by the taxonomic community with or without the endorsement of the code — one of those papers where, if you’ve read the title, you can skip the rest of the paper.   (Although on that score, my effort is knocked into a cocked hat by Hulke 1880.)

The message of the paper will be familiar to anyone who’s been following the Shiny Digital Future thread on this site; as indeed will parts of the text, as the paper is basically a more carefully worked and cohesive form of an argument that I’d previously spread across half a dozen blog posts, a similar number of emails on the ICZN mailing list and any number of comments on other people’s blogs.  The sequence of section headings in the paper tells its own story:

And that conclusion reads as follows:

While we were looking the other way, the digital revolution has happened: everyone but the ICZN now accepts electronic publication. The Code is afforded legitimacy by workers and journals only because it serves them; if we allow it to become anachronistic then they will desert it – or, at best, pick and choose, following only those provisions of the Code that suit them. Facing this reality, the Code has no realistic option but to change – to recognise electronic publishing as valid.

I have no detailed recommendations to make regarding the recently proposed amendments to the Code (ICZN, 2008). Instead I ask only this simple question: will the Code step up to the plate and regulate electronic publications as well as printed publications? Because this is the only question that remains open. Simply rejecting electronic publication is no longer a valid option.

Which I’m sure is familiar rhetoric to long-time SDF advocates, but which I hope will rattle a few cages in the more conservative ranks of specialist taxonomists.  I think it’s a very promising sign that BZN, the official journal of the ICZN, is prepared to publish this kind of advocacy — they didn’t even ask me to tone down the language.  I hope it indicates that in high places, they are sensing which way the wind is blowing.

Here’s a reminder of why electronic publishing is so desirable: figure 3 from Sereno et al.’s (2007) paper on the bizarre skull of the rebbachisaurid Nigersaurus:

Sereno et al. (2007:fig. 3): Nigersaurus taqueti, including photographs of cervical, dorsal and caudal vertebrae in left lateral view.

Let me remind you that this was a paper about skulls — vertebrae were not even on the agenda.  Yet click through the image (go on, you have to) and you will see them each presented in glorious high-resolution detail.  That paper was of course published in the PLoS ONE — a journal that, because it is online only, can provide this quality of figure reproduction, which shames even the very best of printed journals.  To see printed-on-paper figures this detailed and informative, you have to right back to Osborn and Mook (1921).

Which is why I recently decided to put my open-access money where my electronic-only mouth is, and submit the forthcoming Archbishop description to a PLoS journal.  In response to a challenge from Andy Farke, I rather precipitately made a public commitment to do my level best to get that paper submitted this calendar year; and while that may not actually happen, having that goal out there can only help.  Seeing that gorgeous quarry photo of Spinophorosaurus was what tipped me over the edge into wanting to use PLoS.  My plan is to describe the living crap out of that bad boy, photograph every element from every direction and put the whole lot in the paper — make the paper as close as possible as a surrogate for the specimen itself.  Only PLoS (to my knowledge) can do this.

(Of course, once you start wanting to include other kinds of information in your publications — videos, 3d models, etc. — then an electronic-only venue is literally your only option.)

I leave you with two photos of “Cervical P” of the Archbishop; commentary by Matt.  These images are copyright the NHM since it’s their specimen.

Unnamed brachiosaurid NHM R5937, "The Archbishop", Cervical P in right lateral view.

Unnamed brachiosaurid NHM R5937, "The Archbishop", Cervical P in left lateral view.

References

• Hulke, J. W.  1880.  Iguanodon Prestwichii, a new species from the Kimmeridge Clay, distinguished from I. Mantelli of the Wealden Formation in the S.E. of England and Isle of Wight by differences in the shape of the vertebral centra, by fewer than five sacral vertebrae, by the simpler character of its tooth-serrature, &c., founded on numerous fossil remains lately discovered at Cumnor, near Oxford.  Quarterly Journal of the Geological Society 36:433-456. doi:10.1144/GSL.JGS.1880.036.01-04.36
• International Commission on Zoological Nomenclature.  2008.  Proposed amendment of the International Code of Zoological Nomenclature to expand and refine methods of publication.  Zootaxa 1908: 57-67, Bulletin of Zoological Nomenclature 65(4): 265-275 and various other places.
• Osborn, H. F. and C. C. Mook.  1921. Camarasaurus, Amphicoelias and other sauropods of Cope.  Memoirs of the American Museum of Natural History, n.s. 3: 247-387, and plates LX-LXXXV. [HUGE download, but totally worth it.]
• Sereno, Paul C., Jeffrey A. Wilson, Lawrence M. Witmer, John A. Whitlock, Abdoulaye Maga, Oumarou Ide and Timothy A. Rowe.  2007. Structural Extremes in a Cretaceous Dinosaur. PLoS ONE 2 (11): e1230 (9 pages).  doi:10.1371/journal.pone.0001230
• Taylor, Michael P.  2009.  Electronic publication of nomenclatural acts is inevitable, and will be accepted by the taxonomic community with or without the endorsement of the Code.  Bulletin of Zoological Nomenclature 66(3):205-214.

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.

For various arcane reasons, the SV-POW!sketeers are all neck-deep in work, so the blog may actually become somewhat more of the APOD-style picture-n-paragraph thing we originally envisioned, and less of the TetZoo-style monograph-of-the-week thing it’s often leaned toward, at least for a while.

I like it when people decorate their papers with megapixels of vertebral goodness, so here are some caudal vertebrae of the African diplodocine Tornieria, from Remes (2006:fig. 5). Click through to see the figure at its massive native resolution. And check out that pneumaticity! Really, the only question about this image is whether you can settle for just using it as your desktop background, or if you need to print out a wall-sized poster for your bedroom. So the next time you see Kristian Remes, buy him a beer for doing solid work here, on the Humbolt sauropod remount, and on pretty much everything else (including this).

Reference

Remes, K. 2006. Revision of the Tendaguru sauropod Tornieria africana (Fraas) and its relevance for sauropod paleobiogeography. Journal of Vertebrate Paleontology 26 (3): 651–669.

In color, this time, with multiple views, thanks to Xing et al. (2009). They also did a finite element analysis of the tail club and concluded that it was a fairly pathetic weapon. Xing et al. closed by supporting the contention of Ye et al. (2001) that the tail club was a sensory organ. As they stated at the end of the abstract:

The tail club of Mamenchisaurus hochuanensis probably also had limitations as a defense weapon and was more possibly a sensory organ to improve nerve conduction velocity to enhance the capacity for sensory perception of its surroundings.

One thing Xing et al. (2009) cite in support of this is the expanded neural canal inside the club, which they compare to the sacral enlargement in stegosaurs and to the glycogen bodies of birds. They rule out a glycogen body on the grounds that the sacral enlargement in stegosaurs is much bigger than the brain volume, whereas the neural canal enlargement in the M. hochuanensis tail club is much smaller (if you don’t follow that logic, don’t worry, neither do I).

I’m not sure what to make of this thing. On one hand, it would be nice to have more than one club available to rule out the possibility that it’s just a weird paleopathology. On the other hand, it looks oddly regular to be pathological, and the definitive clubs in Shunosaurus and Omeisaurus are at least weak support for this being a genuine feature, although the clubs of the former taxa look very different.

Furthermore, I don’t understand how the authors can rule out the presence of a glycogen body based on the size of the neural expansion alone–especially since the functions of glycogen bodies in extant taxa are very poorly understood (as you may remember from this dustup). Nor can I fathom how a titchy little nerve bundle–if such existed–down at the end of the tail could do much to improve nerve conduction velocity up the rest of the tail. Either my understanding of neuroscience is completely shot, or this hypothesis…lacks support. I am open to being enlightened either way.

Finally, I am disappointed that the authors didn’t pursue the cutting-edge pseudohead hypothesis that has figured prominently here and elsewhere in the blogosphere. There’s a Nobel lurking in there, I just know it.

References

• Xing, L, Ye, Y., Shu, C., Peng, G., and You, H. 2009. Structure, orientation, and finite element analysis of the tail club of Mamenchisaurus hochuanensis. Acta Geologica Sinica 83(6):1031-1040.
• Ye, Y., Ouyang, H., and Fu, Q.-M. 2001. New material of Mamenchisaurus hochuanensis from Zigong, Sichuan. Vertebrata PalAsiatica 39(4):266-271.

In a comment on an earlier article, What’s the deal with your wacky postparapophyses, Shunosaurus?, brian engh asked:

What’s the deal with most Shunosaur “life restorations” showing spikes on the tail club? I can’t find a picture anywhere of a skeleton with any indication of spikes, and yet almost every fleshed-out illustration of Shunosaurs has spikes on it’s tail. Anybody know what that’s about?

It seems we’ve never actually featured the famous Shunosaurus tail-club here before — an amazing oversight, and one that I’m going to remedy right now, thanks to Dong et al. (1989).  This short paper is written in Chinese, so I can’t tell you anything beyond what’s in the figures, captions and English-language abstract.

First up, though, here is his illustration of the famed tail-club:

I can’t help noticing, though, that although the fused clump of enlarged distal caudal vertebrae constitutes a nice club, it’s noticably devoid of spikes.  So it remains a mystery why so many restorations show a spiked club.  Anyone out know why?

Dong et al. (1989) also obligingly includes a figure of the tail-club of Omeisaurus:

And also a photographic plate showing both clubs (though, as is so often the case, the scan has lost a lot of details):

Now, the big question is: why do Shunosaurus and Omeisaurus — and Mamenchisaurus, for that matter — have tail-clubs when they are not closely related, according to modern phylogenies such as those of Wilson (2002) and Upchurch et al. (2004)?  [To be precise, Wilson (2002:fig. 13) had Omeisaurus and Mamenchisaurus clading together, but that clade well separated from Shunosaurus; and Upchurch et al. (2004:fig. 13.18) had all three separate, though with the former two as consecutive branches on the paraphyletic sequence leading to Neosauropoda.]

One possibility is just sheer coincidence: but it’s asking a lot to believe that of the 150 or so known sauropods, the only three for which tail-clubs are known just happened to live more or less at the same time and in the same place.

Another option is some oddity in the environment that strongly encouraged the evolution of tail clubs.  Yes, this is wildly hand-wavy, but you can sort of imagine that maybe all the local theropods thought it was cool to hunt sauropods by biting their tails, and the clubs evolved in response to that.  Or something.  There’s a similar, but even more mystifying, situtation in the late Early Cretaceous Sahara, where the theropod Spinosaurus, the ornithopod Ouranosaurus and arguably even the sauropod Rebbachisaurus all evolved sails.  Why then?  When there?  No-one knows and no-one’s even advanced a hypothesis so far as I know.

Getting back to Jurassic Chinese sauropod tail-clubs, though, there is a third option: could it possibly be that Shunosaurus, Omeisaurus and Mamenchisaurus all form a clade together after all, as proposed back in the day by Upchurch (1998:fig. 19)?  Upchurch’s pioneering (1995, 1998) analyses both recovered a monophyletic “Euhelopodidae” — a clade of Chinese sauropods that included the three genera above plus the early Cretaceous Euhelopus, also from China.  The existence of this clade was one of the two major points of disagreement between Upchurch’s and Wilson’s phylogenies (the other being the position of the nemegtosaurids, Nemegtosaurus and Quaesitosaurus, which Upchurch placed basally within Diplodocoidea but Wilson recovered as titanosaurs).

Upchurch himself has abandoned the idea of the monophyletic Euhelopodidae, as seen in that 2004 analysis and also in Wilson’s and his joint (2009) reassessment of Euhelopus: everyone now agrees that Euhelopus is a basal somphospondyl, i.e. close to Titanosauria, which is a looong way from the basal position that the other Chinese sauropods hold within Sauropoda.)  And so the name Euhelopodidae is no longer used.  But could it be that Upchurch was half-right, and that when Euhelopus is removed that the group that was named after it, a clade remains?

[If so, then that clade is called Mamenchisauridae: as noted by Taylor and Naish (2007), this name was coined by Young and Zhao (1972) and so has priority over the Omeisauridae of Wilson (2002), as Wilson himself now recognises.  Mamenchisauridae was phylogenetically defined (or, as they have it, "diagnosed") by Naish and Martill (2007:498) as "all those sauropods closer to Mamenchisaurus constructus Young, 1954 than to Saltasaurus loricatus Bonaparte".]

As already noted, Omeisaurus and Mamenchisaurus are close together in the recent analyses of both Upchurch and Wilson, so the question becomes: how many additional steps are required to recover Shunosaurus as a member of their clade rather than in its usual more basal position (in the the case of Upchurch’s analysis, to move Omeisaurus up a node)?  And to this, I do not know the answer — to the best of my knowledge, it’s never been tested (or if it has, the result has never been published).  I’d test it myself, but I need to stop working on this post and watch Inca Mummy Girl soonest.  If , say, 20 additional steps are needed, then forget it.  But if we only need, say, three steps, then maybe someone should look at this more closely.  Back in 2004, when he was Young And Stupid, Matt Wedel wrote to me, in a private email which I now quote without permission because I am pretty sure he’s not going to sue me:

Now that I’ve defended the status quo [of using unweighted characters in cladistic analysis], there are some things I’d be happy to bend the rules for.  If an Omeisaurus pops up with a tail club, then Wilson and Sereno be damned, Omeisaurus and Shunosaurus belong in the same clade. [...] So my final word is unweighted characters, please, except for sauropod tail clubs.

Food for thought.

Finally, I leave you with the skeletal reconstruction of Omeisaurus from Dong et al. (1989:fig 3).  Long-time readers will notice a more than passing resemblance to the reconstruction from He et al. (1988:fig. 63), which you can see in Omeisaurus is Just Plain Wrong.

It looks very much as though Dong et al. produced their reconstruction by flipping that of He et al. horizontally and pasting on a tail-club.  Well, we can’t hold that against them — I’d have done the same.

References

• Dong Zhiming, Peng Guangzhao and Huang Daxi. 1988. The Discovery of the bony tail club of sauropods. Vertebrata PalAsiatica 27(3):219-224.
• He Xinlu, Li Kui and Cai Kaiji. 1988. The Middle Jurassic dinosaur fauna from Dashanpu, Zigong, Sichuan, vol. IV: sauropod dinosaurs (2): Omeisaurus tianfuensis. Sichuan Publishing House of Science and Technology, Chengdu, China. 143 pp. + 20 plates.
• Naish, Darren, and David M. Martill. 2007. Dinosaurs of Great Britain and the role of the Geological Society of London in their discovery: basal Dinosauria and Saurischia. Journal of the Geological Society, London, 164: 493-510. (Bicentennial Review issue.)
• 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
• 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, 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.
• Wilson, Jeffrey A. 2002. Sauropod dinosaur phylogeny: critique and cladistic analysis. Zoological Journal of the Linnean Society 136: 217-276.
• Wilson, Jeffrey A. and Paul Upchurch. 2009. Redescription and reassessment of the phylogenetic affinities of Euhelopus zdanskyi (Dinosauria – Sauropoda) from the Early Cretaceous of China. Journal of Systematic Palaeontology 7: 199-239. doi:10.1017/S1477201908002691
• Young, Chung-Chien, 1954. On a new sauropod from Yiping, Szechuan, China. Acta Palaeontologica Sinica II(4):355-369.
• Young, Chung-Chien, and X. Zhao. 1972. [Chinese title. Paper is a description of the type material of Mamenchisaurus hochuanensis]. Institute of Vertebrate Paleontology and Paleoanthropology Monograph Series I, 8:1-30. English translation by W. Downs.

A comment by Charles Epting on the recent article about self-publication led me to check the relevant section of the draft Phylocode, which I’ve read once or twice before but not recently enough for this to have hit me with the force it ought:

From Chapter II. Publication, and specifically Article 4. Publication Requirements:

4.2. Publication, under this code, is defined as distribution of text (but not sound), with or without images. To qualify as published, works must be peer-reviewed, consist of numerous (at least 50 copies), simultaneously obtainable, identical, durable, and unalterable copies, some of which are distributed to major institutional libraries (in at least five countries on three continents) so that the work is generally accessible as a permanent public record to the scientific community, be it through sale or exchange or gift, and subject to the restrictions and qualifications in the present article.

[...]

4.3. The following do not qualify as publication: (a) dissemination of text or images solely through electronic communication networks (such as the Internet) or through storage media (such as CDs, diskettes, film, microfilm and microfiche) that require a special device to read.

I am … flabbergasted, if that’s the word I want.  (I always want to spell that with an “h” after the “g”.)  This language is obviously derived from what’s in the ICZN — for example, “must have been produced in an edition containing simultaneously obtainable copies by a method that assures numerous identical and durable copies” becomes “must consist of numerous (at least 50 copies), simultaneously obtainable, identical, durable, and unalterable copies”.

And the result is that, just like the ICZN, the draft Phylocode does not recognise electronic publication.

Just think about that.  It means that if you define a clade in most of the PLoS journals, it won’t count (unless the journal does one of its inkjet-and-staples special print runs for you).  It also means that any clades you define in Proceedings of the Royal Society of London will not count when the initial online article is published, but only when the later printed edition comes out.  In other words, it means that both the science journals that are growing most quickly in influence and prestige and the oldest science journal in the world will both be useless for phylogenetic nomenclature.

I am sure that’s not what the Phylocode authors want.

That’s particularly true in light of the code’s further requirement that in order to be valid, clade definitions need to be registered.  Really, once a name is officially registered in the Phylocode database and its definition is in a paper published by a reputable publisher and existing in thousands of bit-for-bit-identicial copies in every country in the world, what else is needed for stability?  Fifty stapled inkjet copies?

It seems particularly startling in light of the fact that even the notoriously slow-moving ICZN seems now to be recognising that electronic publishing is inevitable; it would be pretty horrible if by the time the Phylocode is finally implemented, the ICZN has accepted its electronic publishing amendment and the Phylocode is seen to be trailing behind the ICZN in recognising the reality of the world we live in.  (For anyone who is not yet convinced of that reality, I recommend *cough* Taylor 2009, which is a pleasantly easy read.)

Is it too late?  Can the Phylocode be fixed before it’s implemented?  Can it just be done, or will it need lengthy discussion first?  If this doesn’t get fixed, will anyone take the Phylocode seriously?  Is there even a serious argument for keeping the Article 4.2 language as it is now?

I don’t know the answers to any of these questions.  Does anyone else out there?

FIGURE 27. Proximal caudal vertebrae (FMNH PR 2209) of Rapetosaurus krausei in A, anterior view; B, posterior view; C, D, left lateral view. Abbreviations: posl, postspinal lamina; prsl, prespinal lamina; pozg, postzygapophysis. Scale bar equals 3 cm. (Curry Rogers 2009:fig. 27. I'm not sure what part C of this figure is doing here, since it's identical to the rightmost portion of part D. I don't just mean similar, I mean the identical photograph.)

In other news …

I am astounded at the lack of response to University of California vs. Nature, which seems to me just about the most significant thing that’s happened in the world of academic literature since, well, forever.  Can it really be that everyone else’s response is, and I quote, “meh”?

References

• Curry Rogers, K. 2009. The postcranial osteology of Rapetosaurus krausei (Sauropoda: Titanosauria) from the Late Cretaceous of Madagascar. Journal of Vertebrate Paleontology 29(4):1046-1086.
• Taylor, Michael P. 2009. Electronic publication of nomenclatural acts is inevitable, and will be accepted by the taxonomic community with or without the endorsement of the Code. Bulletin of Zoological Nomenclature 66(3):205-214.

In a comment on the initial Shunosaurus tail-club post, Jaime Headden pointed out the passage in the Spinophorosaurus paper (Remes et al. 2009) that discusses the club of Shunosaurus (as justification for positioning the Spinophorosaurus osteoderms on the end of its tail):

With the holotypic skeleton, two closely associated dermal  ossifications were found originating from contralateral sides  (Fig. 4A–C). These elements have a subcircular base that is  rugose and concave on its medial side, and bear a caudodorsally  projecting bony spike with a rounded tip laterally. Although these  elements were found in the pelvic region under the dislocated  scapula, we regard it as most probable that they were placed on  the distal tail in the living animal for the following reasons: First,  the close association of the contralateral elements indicates they  were originally placed near the (dorsal) midline of the body.  Second, the stiffening of the distal tail by specialized chevrons is  also found in other groups of dinosaurs that exhibit tail armor  [42,43]. Third, osteoderms of similar shape are known from the  closely related basal eusauropod Shunosaurus [26]. In the latter  form, these elements cover the middle part of a tail club formed by  coalesced distal vertebrae; however, the decreasing size of the distal-most caudal vertebrae of Spinophorosaurus indicate that such a  club was not present in this genus. The right osteoderm is slightly  larger and differs in proportions from the left element, indicating  that, as in Shunosaurus [26], originally two pairs of tail spines were  present (Fig. 5).

– Remes et al. (2009:6-8)

And this gives the reference that I needed for the Shunosaurus tail-spikes (as opposed to the club) — reference 26 is Zhang (1988), which, embarrassingly, we’ve featured here on SV-POW! in our first Shunosaurus post.  Evidently I was so focussed on preparapophyses when I looked at that monograph that I completely failed to register the tail-club spikes — but then, which of us can truly say he has not made that mistake?

Anyway, here’s what Zhang has to show us:

And here’s that tail again, this time from the poorly reproduced photographic plate 12, part 1, and in right lateral view:

It’s apparent that this really is the other side of the distal tail (rather than a reversed image of the same side) because the osteoderms are in front of the club vertebrae in the left-lateral figure, but behind them in the right-lateral plate.

It would be great to say more about these, but the English language summary of Zhang’s monograph is understandably brief, constituting six pages of the 90.  What’s not quite so understandable is that neither the diagnosis of the genus Shunosaurus nor that of the species S. lii mentions the tail-club or spikes, which are arguably the most distinctive features.  The “revised diagnosis” on pp. 78-79 does, however — just:

Posterior caudals platycoelous, with small cylindrical centra; neural spines low, rod-like.  In several last caudals swollen ralidly [sic] and forming “tail-mace”; in addition there are two pairs of little caudal spines, being analogous to that of stegosaurs.

Not much to go on, but something.  That’s all, though — there is no further description, and crucially, no indication of whether the tail elements were found articulated or whether the spikes were found isolated and subsequently moved to the end of the tail.  It may be that Remes at al. know something I don’t, of course — they might have a translation of Zhang (1988) — but if not, then it’s amusing to consider that the spikes on the tail of Shunosaurus may or may not be supported by evidence, and that the inference of tail-spikes on Spinophorosaurus might be based on dodgy premises.

The other thing that struck me forcibly, as I looked at the figure and plate above, is that the caudal vertebrae remain fairly complex all the way to the end: they retain distinct and prominent neural spines, unlike the distal caudal vertebrae of diplodocids and brachiosaurs.  I notice that the distal caudals of Spinophorosaurus also seem to be complex, based on fig. 3H-I and also on the skeletal reconstruction that is fig. 5 — both of which we’ve reproduced before, in our old Spinophorosaurus article.

So what’s going on here?  Are Shunosaurus and Spinophorosaurus unusual in having distal caudals that retain complex neural spines?  If so, is this property correlated with the possession of a tail-club and/or spines?  Is it causally related?  Or could it be that this is normal for basal eusauropods, and my ideas of sauropod tails have been too coloured by extreme neosauropodocentricity?  Clearly I ought to go and look at a lot more basal sauropods’ distal tails before publishing this post.  And prosauropods’, theropods’, ornithischians’, pterosaurs’, crocadilians’ and lizards’ distal tails.

As it happens, the one non-neosauropod group of reptiles whose distal tails I do know something about is monitor lizards, thanks to my adventures with the corpse of “Charlie”.  And those caudals do maintain astonishingly detailed structure right to the end of the tail, with even absolutely tiny caudals having distinct processes.  Here are some photographs that show this.

First, one showing all 56 caudal vertebrae (the 1st is half in frame at top right, next to the sacrum; the rest read from left to right on successive rows, like words on a page).

Now here are five representative caudals from different regions on the tail — the last ones from each row in the picture above, as it happens: caudals 1, 10, 21, 30, 42 and 56.  They are in more or less dorsal view, though caudal 1 has fallen forward onto its anterior face.  In this and subsequent pictures, caudal 10 (the second shown) is  for some reason back to front.

Now here are the same vertebrae, in the same order and orientation, but now in left dorsolateral aspect (except caudal 10 which is of course in right dorsolateral):

Finally, here are the three smallest of these vertebrae (numbers 30, 42 and 56) in close-up, again in left dorsolateral view, so you can more easily see how much structure even the distalmost caudal has:

That last caudal is about 2.5 mm long.

(It’s interesting that caudals 30 and 42 have those cute fused chevrons.)

So anyway: we know that caudal vertebrae retain distinct structure all the way down to the tip of the tail in monitor lizards at least some basal eusauropods: could it be that this is the primitive state, and that degenerate caudals are found only in neosauropods and mammals?  Gotta prep out some more animals’ skeletons and find out!

References

• Remes, Kristian, Francisco Ortega, Ignacio Fierro, Ulrich Joger, Ralf Kosma, Jose Manuel Marin Ferrer, for the Project PALDES, for the Niger Project SNHM, Oumarou Amadou Ide, and Abdoulaye Maga. 2009. A new basal sauropod dinosaur from the Middle Jurassic of Niger and the early evolution of Sauropoda. PLoS ONE 4(9):e6924. doi:10.1371/journal.pone.0006924
• Zhang Yihong. 1988. The Middle Jurassic dinosaur fauna from Dashanpu, Zigong, Sichuam, vol. 1: sauropod dinosaur (I): Shunosaurus. Sichuan Publishing House of Science and Technology, Chengdu, China.

This year, I missed The Paleo Paper Challenge over on Archosaur Musings — it was one of hundreds of blog posts I missed while I was in Cancun with my day-job and then in Bonn for the 2nd International Workshop on Sauropod Biology and Gigantism.  That means I missed out on my annual tradition of promising to get the looong-overdue Archbishop description done by the end of the year.

Brachiosauridae incertae sedis NMH R5937, "The Archbishop", dorsal neural spine C, probably from an anterior dorsal vertebra. Top row: dorsal view, anterior to top; middle row, left to right: anterior, left lateral, posterior, right lateral; bottom row: ventral view, anterior to bottom.

But this year, Matt and I are going to have our own private Palaeo Paper Challenge.  And to make sure we heap on maximum pressure to get the work done, we’re announcing it here.

Here’s the deal.  We have two manuscripts — one of them Taylor and Wedel, the other Wedel and Taylor — which have been sitting in limbo for a stupidly long time.  Both are complete, and have in fact been submitted once and gone through review.  We just need to get them sorted out, turned around, and resubmitted.

(The Taylor and Wedel one is on the anatomy of sauropod cervicals and the evolution of their long necks.  It’s based on the last remaining unpublished chapter of my dissertation, and turned up in a modified form as my SVPCA 2010 talk, Why Giraffes Have Such Short Necks.  The Wedel and Taylor one is on the occurrence and implications of intermittent pneumaticity in the tails of sauropods, and turned up as his SVPCA 2010 talk, Caudal pneumaticity and pneumatic hiatuses in the sauropod dinosaurs Giraffatitan and Apatosaurus.)

We’re going to be realistic: we both have far too much going in (incuding, you know, families) to get these done by the end of 2011.  But we have relatively clear Januaries, so our commitment is that we will submit by the end of January 2012.  If either of us fails, you all have permission to be ruthlessly derisive of that person.

… and in other news …

Some time while we were all in Bonn, the SV-POW! hit-counter rolled over the One Million mark.  Thanks to all of your for reading!

This is the third post in a series on neural spine bifurcation in sauropods, inspired by Woodruff and Fowler (2012). In the first post, I looked at neural spine bifurcation in Morrison sauropod genera based on the classic monographic descriptions. In the second post, I showed that size is an unreliable criterion for assessing age and that serial variation can mimic ontogenetic change in sauropod cervicals. In this post I look at the evidence for ontogenetic changes in neural spine bifurcation presented by Woodruff and Fowler (2012). This posts builds on the last two, so please refer back to them as needed.

Another opening digression, on the OMNH baby sauropod material this time

Nearly all of the Morrison Formation material in the OMNH collections comes from Black Mesa in the Oklahoma panhandle. It was collected in the 1930s by WPA crews working under the direction of J. Willis Stovall. Adequate tools and training for fossil preparation were in short supply. A lot of the prep was done by unskilled laborers using hammers, chisels, pen-knives, and sandpaper (apologies if you have experience with fossil preparation and are now feeling a bit ill). Uncommonly for the Morrison, the bones are very similar in color to the rock matrix, and the prep guys sometimes didn’t realize that they were sanding through bone until they got through the cortex and  into the trabeculae. Consequently, a lot of interesting morphology on the OMNH Morrison material has been sanded right off, especially some of the more delicate processes on the vertebrae. This will become important later on.

Do the ‘ontogenetic’ series in Woodruff and Fowler (2012) actually show increasing bifurcation through development?

In the Materials and Methods, Woodruff and Fowler (2012:2) stated:

Study specimens comprise 38 cervical, eight dorsal, and two caudal vertebrae from 18 immature and one adult diplodocid (Diplodocus sp., Apatosaurus sp., and Barosaurus sp.), and two immature macronarians (both Camarasaurus sp.).

However, their Table 1 and Supplementary Information list only 15 specimens, not 18. Of the 15, one is probably not a diplodocid (SMA 0009 ‘Baby Toni’) — a fact that, oddly, the authors knew, as stated in the Supplementary Information.  Of the remaining 14 specimens, 11 are isolated vertebrae, so only three represent reasonably complete probably-diplodocoid series (MOR 592, AMNH 7535, and CM 555). From CM 555 they discuss only one vertebra, the C6; and AMNH 7535 is not mentioned at all outside of Table 1 and a passing mention the Supplementary Information, so the subadult data actually used in the paper consist of isolated vertebrae and one articulated series, MOR 592. (For the sake of comparison, in the first post on this topic I looked up 10 articulated series, only two of which–Diplodocus carnegii CM 84/94 and Camarasaurus lentus CM 11338–are even mentioned in Woodruff and Fowler [2012].)

In light of the previous post, on serial variation, the dangers of using isolated vertebrae should by now be apparent. Recall that even adult diplodocids are expected to have completely unsplit spines as far back as C5 (Apatosaurus) or C8 (Barosaurus) and as far forward as D7 (Apatosaurus) or D6 (Barosaurus), and only partially split spines in the adjacent positions. Furthermore, size is a notoriously unreliable criterion of age; MOR 790 8-10-96-204 from Figure 2 in Woodruff and Fowler (2012) also appears in their Figure 3 as the second-smallest vertebra in this ‘ontogenetic’ series, despite most likely coming from a well-fused adult approximately the same size as the D. carnegii individual that represents the end of the series. So without any evidence other than sheer size (if that size overlaps with the adult size range) and degree of neural spine bifurcation (which cannot help but overlap with the adult range, since the adult range encompasses all possible states), simply picking small vertebrae with unsplit spines and calling them juvenile is at best circular and at worst completely wrong–as in the case of MOR 790 8-10-96-204 examined in the last post.

Unfortunately it is not possible to tell what criteria Woodruff and Fowler (2012) used to infer age in their specimens, because they don’t say. Neural arch fusion is discussed in general terms in the Supplementary Information, but in the text and in the figures everything is discussed simply in terms of size. For example:

In the next largest specimen (MOR 790 7-26-96-89, vertebral arch 9.9 cm high), the neural spine is relatively longer still and widens at the apex…

The Supplementary Information provides more evidence that Woodruff and Fowler (2012) did not consider the confounding effects of size, serial position, and ontogenetic stage. In the section on the Mother’s Day Quarry in the Supplementary Information, they wrote:

Because of this size distribution it is not surprising that there are also different ontogenetic stages present which result in cervical centrum lengths varying between 12 and 30 cm.

Now, there may be different ontogenetic stages present in the quarry, and the cervicals in the quarry may vary in length by a factor of 2.5, but the latter does not demonstrate the former. In D. carnegii CM 84/94 the longest postaxial cervical (C14, 642 mm) is 2.6 times the length of the shortest (C3, 243 mm; data from Hatcher 1901). The size range reported as evidence of multiple ontogenetic stages by Woodruff and Fowler (2012) turns out to be slightly less than that expected in a single individual.

With that in mind, let’s look at each of the putative ontogenetic sequences in Woodruff and Fowler (2012):

Anterior cervical vertebrae

Woodruff and Fowler (2012:fig. 3)

The proposed ontogenetic series used by Woodruff and Fowler (2012) for anterior cervical vertebrae consists of:

• CMC VP7944, an isolated ?Diplodocus vertebra from the Mother’s Day site, which is described in the text but not pictured;
• MOR 790 7-30-96-132, an isolated vertebra from the same site;
• MOR 790 8-10-96-204, another isolated vertebra from the same site;
• MOR 592, from a partial cervical series of a subadult Diplodocus but with the serial position unspecified;
• ANS 21122, C6 of Suuwassea (included in Fig. 3, but not discussed as evidence in the accompanying text)
• CM 555, C6 of a nearly complete (C2-C14) cervical series of a subadult Apatosaurus;
• CM 84/94, C7 of Diplodocus carnegii

CMC VP7944 is not pictured, but from the description in the text it’s perfectly possible that it represents a C3, C4, or C5, all of which have undivided spines even in adult diplodocids. It therefore contributes no information: the hypothesis that the spine is undivided because of ontogeny is not yet demonstrated, and the hypothesis that the spine is undivided because of serial position is not yet falsified.

MOR 790 7-30-96-132 is shown only from the front, so the centrum proportions and the shape of the neural spine cannot be assessed. The neural arch appears to be fused, but the cervical ribs are not. Again, we cannot rule out the possibility that it comes from an very anterior cervical and therefore its undivided spine could be an artifact of its serial position. It therefore contributes no information on possible ontogenetic changes in neural spine bifurcation.

As shown in the previous post, MOR 790 8-10-96-204 is probably a C4 or C5 of an adult or near-adult Diplodocus about the same size as or only slightly smaller than D. carnegii CM 84/94. It is small and has an undivded spine because it is an anterior cervical, not because it is from a juvenile. It therefore contributes no support to the ontogenetic bifurcation hypothesis.

The pictured vertebra of MOR 592 has a shallow notch in the tip of the spine, which is expected in C6 in Apatosaurus and Diplodocus and in C9 and C10 in Barosaurus. The serial position of the vertebra is not stated in the paper, but about half of the anterior cervicals even in an adult diplodocid are expected to have unsplit or shallowly split spines based on serial position alone. Based on the evidence presented, we cannot rule out the possibility that the shallow cleft in the pictured vertebra is an artifact of serial position rather than ontogeny. It therefore contributes no support to the ontogenetic bifurcation hypothesis.

ANS 21122 has an incompletely divided neural spine, which is in fact expected for the sixth cervical in adult diplodocids as shown by A. parvus CM 563/UWGM (in which C6 is missing but C5 has an unsplit spine and C7 a deeply bifid spine) and D. carnegii CM 84/94 (in which C6 is also shallowly bifid). A. ajax NMST-PV 20375 has a wider split in the spine of C6, but the exact point of splitting appears to vary by a position or two among diplodocids. The hypothesis that the spine of ANS 21122 C6 is already as split as it would ever have gotten cannot be falsified on the basis of the available evidence.

CM 555 C6: see the previous paragraph. Note that in ANS 21122 the neural arch and cervical ribs are fused in C6, and in C6 of CM 555 they are not.

CM 84/94 C7 has a deeply split spine, but this expected at that position. C6 of the same series has a much more shallow cleft, and C5 would be predicted to have no cleft at all (recall from the first post that the neural spines of C3-C5 of this specimen are sculptures). So any trend toward increasing bifurcation is highly dependent on serial position; if serial position cannot be specified then it is not possible to say anything useful about the degree of bifurcation in a given vertebra.

Summary. CMC VP7944 and MOR 790 7-30-96-132 could be very anterior vertebrae, C3-C5, in which bifurcation is not expected even in adults. Since they are isolated elements, that hypothesis is very difficult to falsify. MOR 790 8-10-96-204 is almost certainly a C4 or C5 of an adult or near-adult Diplodocus. ANS 21122 and CM 555 C6 are incompletely divided, as expected for vertebrae in that position even in adults. CM 84/94 has a shallowly divided spine in C6 and more deeply bifid spines from C7 onward, just like CM 555.

Verdict: no ontogenetic change has been demonstrated.

Posterior cervical vertebrae

Woodruff and Fowler (2012:Fig. 4A)

The proposed ontogenetic series includes:

• OMNH 1267 and 1270
• MOR 790 7-26-96-89
• MOR 592
• CM 84/94

OMNH 1267 and 1270 are isolated neural arches of baby sauropods from the Black Mesa quarries. OMNH 1267 does not appear to be bifurcated, but it has a very low neural spine and it was probably sanded during preparation, so who knows what might have been lost. OMNH 1270 actually shows a bifurcation–Woodruff and Fowler (2012:3) describe it as having “a small excavated area”–but again it is not clear that the spines are as intact now as they were in life. More seriously,  since these are isolated elements (you can all join in with the refrain) their serial position cannot be determined with any accuracy, and therefore they are not much use in determining ontogenetic change. Although they are anteroposteriorly short, that does not necessarily make them posterior cervicals. The cervical vertebrae of all sauropods start out proportionally shorter and broader than they end up (Wedel et al. 2000:368-369), and the possibility that these are actually from anterior cervicals–not all of which are expected to have bifurcations–is difficult to rule out.

The other three vertebrae in the series have deeply bifurcated spines. In the text, Woodruff and Fowler (2012:3) make the case that the bifurcation in MOR 592 is deeper than in the preceding vertebra, MOR 790 7-26-96-89. However, the proportions of the two vertebrae are very different, suggesting that they are from different serial positions, and the centrum of MOR 790 7-26-96-89 is actually larger in diameter than that of the representative vertebra from MOR 592. So unless centrum size decreased through ontogeny, these vertebrae are not comparable. As usual, we don’t know where in the neck the isolated MOR 790 vertebra belongs, and we only see it in anterior view. Nothing presented in the paper rules out possibility that is actually an anterior cervical, and in fact the very low neural spines suggest that that is the case.

Allowing for lateral crushing, the vertebra from MOR 592 (again, we are not told which one it is) looks very similar to the D. carnegii CM 84/94 vertebra (C15–again, I had to look it up in Hatcher), and is probably from a similar position in the neck. In comparing the two, Woodruff and Fowler (2012:4) say that in CM 84/94, “the bifurcated area has broadened considerably”, but this clearly an illusion caused by the lateral compression of the MOR 592 vertebra — its centrum is also only half as wide proportionally as in the CM 84/94 vertebra.

Summary. The OMNH vertebrae are of unknown serial position and probably lost at least some  surface bone during preparation, so their original degree of bifurcation is hard to determine. The other three vertebrae in the series all have deeply bifid spines, but they are out of order by centrum size, MOR 790 7-26-96-89 might be an anterior cervical based on its low neural spines, and the “broadening” of the trough between MOR 792 and CM 84/94 is an artifact of crushing.

Verdict: no ontogenetic change has been demonstrated.

Anterior dorsal vertebrae

Woodruff and Fowler (2012:Fig. 5A)

The ontogenetic series here consists of:

• MOR 790 7-17-96-45
• MOR 592
• CM 84/94

As usual, the serial positions of the MOR 592 and CM 84/94 vertebrae are presumably known but not stated in the paper. The D. carnegii CM 84/94 vertebra is D4. Comparisons to the MOR 592 vertebra are not helped by the fact that it is shown in oblique posterior view. Nevertheless, the two vertebrae are very similar and, based on the plates in Hatcher (1901), the MOR 592 vertebra is most likely a D4 or D5 of Diplodocus. The spines in the larger two vertebrae are equally bifurcated, so the inference of ontogenetic increase in bifurcation rests on the smallest of the three vertebrae, MOR 790 7-17-96-45.

MOR 790 7-17-96-45 is an isolated unfused neural arch, clearly from a juvenile. Its serial position is hard to determine, but it is probably not from as far back as D4 or D5 because it appears to lack a hypantrum and shows no sign of the parapophyses, which migrate up onto the neural arch through the cervico-dorsal transition. The element is only figured in anterior view, so it is hard to tell how long it is proportionally. Still, based on the single photo in the paper (which is helpfully shown at larger scale in Fig. 5B), it seems to be reasonably long, with the prezygapophyses, transverse processes, neural spines, and postzygapophyses well separated from anterior to posterior. In fact, I see no strong evidence that it is a dorsal neural arch at all–the arch of a posterior cervical would look the same in anterior view.

Given that MOR 7-17-96-45 lacks a hypantrum and parapophyses, it is not directly comparable to the two larger vertebrae. Although we cannot determine its position in the presacral series, its spine is shallowly bifurcated, to about half the distince from the metapophyses to the postzygapophyses. In Apatosaurus louisae CM 3018, the notch in D3 is about equally deep, and in C15 it is only slightly deeper, still ending above the level of postzygapophyses. So there is some variation in the depth of the bifurcation in the posterior cervicals and anterior dorsals in the North American diplodocids. Without knowing the precise serial position of MOR 7-17-96-45, it is difficult to derive inferences about the ontogeny of neural spine bifurcation.

Diplodocid anterior dorsal vertebrae. Left and right, dorsal vertebrae 3 and 4 of adult Apatosaurus louisae holotype CM 3018, from Gilmore (1936: plate XXV). Center, juvenile neural arch MOR 7-17-96-45, modified from Woodruff and Fowler (2012: fig. 5B), corrected for shearing and scaled up.

What this element does conclusively demonstrate is that the neural arches of posterior cervicals or anterior dorsals in even small, unfused juvenile diplodocids were in fact bifurcated to to a degree intermediate between  D3 and D4 in the large adult Apatosaurus louisae CM3018 — in fact, so far as neural cleft depth is concerned, it makes rather a nice intermediate between them.  (It differs in other respects, most notable that it is proportionally broad, lacks a hypantrum and parapophyses, etc.)

Summary. The two larger specimens in the ‘ontogenetic series’ are from similar serial positions and show the same degree of bifurcation. MOR 7-17-96-45 is from a more anterior position, based on its lack of hypantrum and parapophyses.  Although it is a juvenile, its degree of bifurcation is similar to that of anterior dorsal vertebrae in adult Apatosaurus (and that of C15 in A. louisae CM 3018, if MOR 7-17-96-45 is, in fact, a cervical).

Verdict: no ontogenetic change has been demonstrated.

Posterior dorsal vertebrae

Woodruff and Fowler (2012:Fig. 6A)

The ontogenetic series consists of:

• OMNH 1261
• MOR 592
• CM 84/94

The D. carnegii CM 84/94 vertebra is D6, and based on its almost identical morphology the MOR 592 vertebra is probably from the same serial position. They show equivalent degrees of bifurcation.

OMNH 1261 is another isolated juvenile neural arch. The portion of the spine that remains is unbifurcated. However, the spine is very short and it is possible that some material is missing from the tip. More importantly, the last 3-4 dorsals in Apatosaurus, Diplodocus, and Barosaurus typically have extremely shallow notches in the neural spines or no notches at all. If OMNH 1261 is a very posterior dorsal, it would not be expected to show a notch even when fully mature.

Verdict: no ontogenetic change has been demonstrated.

Woodruff and Fowler (2012:Fig. 7)

Caudal vertebrae

The ontogenetic series here consists of:

• MOR 592
• CM 84/94

The first thing to note is that the ‘bifurcation’ in MOR 592 is at right angles to that in the proximal caudals of D. carnegiiCM 84/94, so the one can hardly be antecedent to the other.

More importantly, antero-posterior ‘bifurcations’ like that in MOR 592 are occasionally seen in the caudal vertebrae of adult sauropods. Below are two examples, caudals 7 and 8 of A. parvus CM 563/UWGM 15556. In other words, in this character MOR 592 already displays adult morphology.

Verdict: no ontogenetic change has been demonstrated.

A. parvus CM 563/UWGM 15556 caudals 8 and 7 in right lateral view, from Gilmore (1936:pl.. 33)

Camarasaurus

The ontogenetic series here consists of:

• OMNH 1417
• AMNH 5761

OMNH 1417 is an isolated cervical neural spine, and the pictured vertebra of Camarasaurus supremus AMNH 5761 is a posterior cervical. In C. grandis and C. lewisi, all of the cervical vertebrae eventually develop at least a shallow notch in the tip of the neural spine, but as shown in the previous post there seems to be some variation between Camarasaurus species, and, likely, between individuals. In the absence of information about its serial position and the species to which it belonged, the lack of bifurcation in OMNH 1417 is uninformative; it could belong to an anterior cervical of C. supremus that would not be expected to develop a bifurcation.

Verdict: no ontogenetic change has been demonstrated. There is evidence that neural spine bifurcation developed ontogenetically in Camarasaurus, but it comes from the juvenile C. lentus CM 11338, described by Gilmore (1925), and the geriatric C. lewisi, described by McIntosh, Miller et al. (1996)–see the first post in this series for discussion.

Conclusions

The ‘ontogenetic’ series of Woodruff and Fowler (2012) are not really ontogenetic series. In all of the diplodocid presacral vertebrae and in Camarasaurus, the smallest elements in the series are isolated vertebrae or neural arches for which the serial position is almost impossible to determine (and for the reader, completely impossible given the limited information in the paper) and even the taxonomic identifications are suspect (e.g., the OMNH material–how one reliably distinguishes the Apatosaurus and Camarasaurus neural arches is beyond me). The larger vertebrae in the presacral series are all compromised in various ways: one includes an adult masquerading as a juvenile (MOR 790 8-10-96-204 in the anterior cervicals), one is out of order by centrum size (MOR 790 7-26-96-89 and MOR 592 in the posterior cervicals), and two show no change in degree of bifurcation from the middle of the series to the upper end (MOR 592 and CM 84/94 in the anterior and posterior dorsals). The shallow longitudinal bifurcation in the MOR 592 caudal vertebra is similar to those found in caudal vertebrae of adult diplodocids, and is not antecedent to the transverse bifurcations discussed in the rest of the paper.

Crucially, when information on size and serial position is taken into account, none of the ‘ontogenetic series’ in the paper show any convincing evidence that neural spine bifurcation increases over ontogeny. The best evidence that bifurcation does increase over ontogeny comes from Camarasaurus, specifically the juvenile C. lentus CM 11338 described by Gilmore (1925) and geriatric C. lewisi BYU 9047 described by McIntosh et al. (1996), it was already recognized prior to Woodruff and Fowler (2012), and it has not caused any taxonomic confusion.

There is an asymmetry of interference here. To call into question the conclusions of Woodruff and Fowler (2012), all one has to do is show that the evidence could be explained by serial, intraspecific, or interspecific variation, taphonomy, damage during preparation, and so on. But to demonstrate that bifurcation develops over ontogeny, one has to falsify all of the competing hypotheses. I know of only one way to do that: find a presacral vertebral column that is (1) articulated, (2) from an individual that is clearly juvenile based on criteria other than size and degree of bifurcation, which (3) can be confidently referred to one of the known genera, and then show that it has unbifurcated spines in the same serial positions where adult vertebrae have bifurcated spines. Isolated vertebrae are not enough, bones from non-juveniles are not enough, and juvenile bones that might pertain to new taxa are not enough. It may be that this is not yet possible because the necessary fossils just haven’t been found yet. I am not suggesting that we stop doing science, or that the ontogenetic hypothesis of neural spine bifurcation is unreasonable. It’s perfectly possible that it’s true (though MOR 7-17-96-45 ironically suggests otherwise). But it’s not yet been demonstrated, at least for diplodocids, and to the extent that the taxonomic hypotheses of Woodruff and Fowler (2012) rely on an ontogenetic increase in bifurcation in diplodocids, they are suspect. That will be the subject of the next post.

The rest of the series

Links to all of the posts in this series:

• Part 1: what we knew a month ago
• Part 2: why serial position matters
• Part 3: the evidence from ontogenetic series
• Part 4: is Suuwassea a juvenile of a known diplodocid?
• Part 5: is Haplocanthosaurus a juvenile of a known diplodocid?
• Part 6: more reasons why Haplocanthosaurus is not a juvenile of a known diplodocid

and the post that started it all:

• Changes through growth in sauropods and ornithopods

References

• Gilmore, C.W. 1925. A nearly complete articulated skeleton of Camarasaurus, a saurischian dinosaur from the Dinosaur National Monument. Memoirs of the Carnegie Museum 10:347-384.
• Gilmore, C.W. 1936. Osteology of Apatosaurus with special reference to specimens in the Carnegie Museum. Memoirs of the Carnegie Museum 11:175-300.
• Hatcher, J.B. 1901. Diplodocus (Marsh): its osteology, taxonomy, and probable habits, with a restoration of the skeleton. Memoirs of the Carnegie Museum 1:1-63.
• McIntosh, J.S., Miller, W.E., Stadtman, K.L., and Gillette, D.D. 1996. The osteology of Camarasaurus lewisi (Jensen, 1988). BYU Geology Studies 41:73-115.
• Wedel, M.J., Cifelli, R.L., and Sanders, R.K. 2000. Osteology, paleobiology, and relationships of the sauropod dinosaur Sauroposeidon. Acta Palaeontologica Polonica 45(4):343-388.
• Woodruff, D.C, and Fowler, D.W. 2012. Ontogenetic influence on neural spine bifurcation in Diplodocoidea (Dinosauria: Sauropoda): a critical phylogenetic character. Journal of Morphology, online ahead of print.

Caudal pneumaticity in saltasaurines. Cerda et al. (2012: fig. 1).

Earlier this month I was amazed to see the new paper by Cerda et al. (2012), “Extreme postcranial pneumaticity in sauropod dinosaurs from South America.” The title is dramatic, but the paper delivers the promised extremeness in spades. Almost every figure in the paper is a gobsmacker, starting with Figure 1, which shows pneumatic foramina and cavities in the middle and even distal caudals of Rocasaurus, Neuquensaurus, and Saltasaurus. This is most welcome. Since the 1990s there have been reports of saltasaurs with “spongy bone” in their tail vertebrae, but it hasn’t been clear until now whether that “spongy bone” meant pneumatic air cells or just normal marrow-filled trabecular bone. The answer is air cells, loads of ‘em, way farther down the tail than I expected.

Caudal pneumaticity in diplodocines. Top, transverse cross-section through an anterior caudal of Tornieria, from Janensch (1947: fig. 9). Bottom, caudals of Diplodocus, from Osborn (1899: fig. 13).

Here’s why this is awesome. Lateral fossae occur in the proximal caudals of lots of neosauropods, maybe most, but only a few taxa go in for really invasive caudal pneumaticity with big internal chambers. In fact, the only other sauropod clade with such extensive pneumaticity so far down the tail are the diplodocines, including Diplodocus, Barosaurus, and Tornieria. But they do things differently, with BIG, “pleurocoel”-type foramina on the lateral surfaces of the centra, leading to BIG–but simple–camerae inside, and vertebral cross-sections that look like I-beams. In contrast, the saltasaurines have numerous small foramina on the centrum and neural arch that lead to complexes of small pneumatic camellae, giving their vertebrae honeycomb cross-sections. So caudal pneumaticity in diplodocines and saltsaurines is convergent in its presence and extent but clade-specific in its development. Pneumaticity doesn’t get much cooler than that.

Pneumatic ilia in saltasaurines. Cerda et al. (2012: fig. 3).

But it does get a little cooler. Because the stuff in the rest of the paper is even more mind-blowing. Cerda et al. (2012) go on to describe and illustrate–compellingly, with photos–pneumatic cavities in the ilia, scapulae, and coracoids of saltasaurines. And, crucially, these cavities are connected to the outside by pneumatic foramina. This is important. Chambers have been reported in the ilia of several sauropods, mostly somphospondyls but also in the diplodocoid Amazonsaurus. But it hasn’t been clear until now whether those chambers connected to the outside. No patent foramen, no pneumaticity. It seemed unlikely that these sauropods had big marrow-filled vacuities in their ilia–as far as I know, all of the non-pneumatic ilia out there in Tetrapoda are filled with trabecular bone, and big open marrow spaces only occur in the long bones of the limbs. And, as I noted in my 2009 paper, the phylogenetic distribution of iliac chambers is consistent with pneumaticity, in that the chambers are only found in those sauropods that already have sacral pneumaticity (showing that pneumatic diverticula were already loose in their rear ends). But it’s nice to have confirmation.

So, the pneumatic ilia in Rocasaurus, Neuquensaurus, and Saltasaurus are cool because they suggest that all the other big chambers in sauropod ilia were pneumatic as well. And for those of you keeping score at home, that’s another parallel acquisition in Diplodocoidea and Somphospondyli (given the apparent absence of iliac chambers in Camarasaurus and the brachiosaurids, although maybe we should bust open a few brachiosaur ilia just to be sure*).

* I kid, I kid.**

** Seriously, though, if you “drop” one and find some chambers, call me!

Pectoral pneumaticity in saltasaurines. Cerda et al. (2012: fig. 2).

But that’s not all. The possibility of pneumatic ilia has been floating around for a while now, and most of us who were aware of the iliac chambers in sauropods probably assumed that eventually someone would find the specimens that would show that they were pneumatic. At least, that was my assumption, and as far as I know no-one ever floated an alternative hypothesis to explain the chambers. But I certainly did not expect pneumaticity in the shoulder girdle. And yet there they are: chambers with associated foramina in the scap and coracoid of Saltasaurus and in the coracoid of Neuquensaurus. Wacky. And extremely important, because this is the first evidence that sauropods had clavicular air sacs like those of theropods and pterosaurs. So either all three clades evolved a shedload of air sacs independently, or the basic layout of the avian respiratory system was already present in the ancestral ornithodiran. I know where I’d put my money.

There’s loads more interesting stuff to talk about, like the fact that the ultra-pneumatic saltasaurines are among the smallest sauropods, or the way that fossae and camerae are evolutionary antecedent to camellae in the vertebrae of sauropods, so maybe we should start looking for fossae and camerae in the girdle bones of other sauropods, or further macroevolutionary parallels in the evolution of pneumaticity in pterosaurs, sauropods, and theropods. Each one of those things could be a blog post or maybe a whole dissertation. But my mind is already thoroughly blown. I’m going to go lie down for a while. Congratulations to Cerda et al. on what is probably the most important paper ever written on sauropod pneumaticity.

References

• Cerda, I.A., Salgado, L., and Powell, J.E. 2012. Extreme postcranial pneumaticity in sauropod dinosaurs from South America. Palaeontologische Zeitschrift. DOI 10.1007/s12542-012-0140-6
• Janensch, W. 1947. Pneumatizitat bei Wirbeln von Sauropoden und anderen Saurischien. Palaeontographica, Supplement 7, 3:1–25.
• Osborn, H. F. 1899. A skeleton of Diplodocus. Memoirs of the American Museum of Natural History 1:191–214.

Last night London and I spent the night in the Natural History Museum of Los Angeles County (LACM), as part of the Camp Dino overnight adventure. So we got lots of time to roam the exhibit halls when they were–very atypically–almost empty. Above are the museum’s mounted Triceratops–or one of them, anyway–and mounted cast of the Mamenchisaurus hochuanensis holotype, presented in glorious not-stygian-darkness (if you went through the old dino hall, pre-renovation, you know what I mean).

We got there early and had time to roam around the museum grounds in Exposition Park. The darned-near-life-size bronze dinos out front are a minor LA landmark.

The rose garden was already closed, but we walked by anyway, and caught this rainbow in the big fountain.

After we checked in we had a little time to roam the museum on our own. I’ve been meaning to blog about how much I love the renovated dinosaur halls. The bases are cleverly designed to prohibit people touching the skeletons without putting railings or more than minimal glass in the way, and you can walk all the way around the mounted skeletons and look down on them from the mezzanine–none of that People’s Gloriously Efficient Cattle Chute of Compulsory Dinosaur Appreciation business. Signage is discreet and informative, and so are the handful of interactive gizmos. London and I spent a few minutes using a big touch-screen with a slider that controlled continental drift from the Triassic to the present–a nice example of using technology to add value to an exhibit without taking away from the real stuff that’s on display. There are even a few places to sit and just take it all in. That’s pretty much everything I want in a dinosaur hall.

Also, check out the jumbotron on the left in the above photo. It was running a (blessedly) narration-free video on how fossils are found, collected, prepared, mounted, and studied, on about a five-minute loop. Lots of pretty pictures. Including this next one.

There are a couple of levels of perspective distortion going on here, both in the original photo and in my photo of that photo projected on the jumbotron. Still, I feel confident positing that that is one goldurned big ilium. I’m not going to claim it’s the biggest bone I’ve ever seen–that rarely ends well–but sheesh, it’s gotta be pretty freakin’ big. And apparently a brachiosaurid, or close to it. Never mind, it’s almost certainly an upside-down Triceratops skull. Thanks to Adam Yates for the catch. I will now diminish, and go into the West.

Triceratops, Styracosaurus, and Einiosaurus–collect the whole set!

Of course, the centerpiece of the second dinosaur hall–and how great is it that there are two!?–is the T. rex trio: baby, juvenile (out of frame to the right), and subadult. Yes, subadult: the “big” one is not as big as the really big rexes, and from the second floor you can see unfused neural arches in some of the caudal vertebrae (many thanks to Ashley Fragomeni for pointing those out to me on a previous visit).

Awwwww! C’mere, little fella!

Still, this ain’t Vulgar Overstudied Theropod Picture of the Week. Here are some sweet pneumatic diplodocid caudals in the big wall o’ fossils (visible behind Mamenchisaurus in the overhead photo above). The greenish color is legit–in the Dino Lab on the second floor, they’re prepping a bunch of sauropod elements that look like they were carved out of jade.

Sudden violent topic shift, the reason for which will be become clear shortly: London and I have been sculpting weapons of mass predation in our spare time. In some of the photos you may be able to see his necklace, which has a shark tooth he sculpted himself. Here are a couple of allosaur claws I made–more on those another time.

The point is, enthusiasm for DIY fossils is running very high at Casa Wedel, so London’s favorite activity of the evening was molding and casting. Everyone got to make a press mold using a small theropod tooth, a trilobite, or a Velociraptor claw. Most of the kids I overheard opted for the tooth, but London went straight for the claw.

Ready for plaster! Everyone got to pick up their cast at breakfast this morning, with instructions to let them cure until this evening. All went well, so I’ll spare you a photo of this same shape in reverse.

We were split into three tribes of maybe 30-40 people each, and each tribe bedded down in a different hall. The T. rex and Raptor tribes got the North American wildlife halls, but our Triceratops tribe got the African wildlife hall, which as a place to sleep is about 900 times cooler. Someone had already claimed the lions when we got there, so London picked hyenas as our totem animals.

Lights out was at 10:30 PM, and the lights came back on at 7:00 this morning. Breakfast was out from 7:15 to 8:00, and then we had the museum to ourselves until the public came in at 9:30. So I got a lot of uncluttered photos of stuff I don’t usually get to photograph, like this ammonite. Everyone should have one of these.

London’s favorite dino in the museum is Carnotaurus. It’s sufficiently weird that I can respect that choice.

Not that there’s anything wrong with the old standards, especially when they’re presented as cleanly and innovatively as they are here.

Finally, the LACM has a no tripod policy, and if they see you trying to carry one in they will make you take it back to your car. At least during normal business hours. But no one searched my backpack when we went in last night, and I put that sucker to some good use. Including getting my first non-bigfoot picture of the cast Argentinosaurus dorsal. It was a little deja-vu-ey after just spending so much time with the giant Oklahoma Apatosaurus–elements of the two animals really are very comparable in size.

If you’re in the LA area and interested in spending a night at the museum–or at the tar pits!–check out the “Overnight Adventures” page on the museum’s website. Cost is $50 per person for members or $55 for non-members, and worth every penny IMHO. It’s one of those things I wish we’d done years ago.

This is a caudal vertebra from the middle of the tail of an ostrich, LACM Bj342:

The middle row shows it in anterior, left lateral and posterior views; above and below the anterior view are the dorsal and ventral views. It’s about 5 cm across the transverse processes. (This figure is from a manuscript that Matt and I will submit to a journal probably within 24 hours.)

In compositing the different views, I had a heck of a time recognising what was what. The dorsal view looks so much more like what we’d expect a ventral view to look like — indeed, the two are more similar for this vertebra than for any other I’ve seen.

How about those big pnuematic foramina right at the top of the bone? At first, Matt and I thought we’d never seen anything like that before. But then we realised that we sort of had — in a cervical vertebra of Apatosaurus which appears as part one of Taylor and Wedel (2013: figure 9).

This is Apatosaurus sp. OMNH 01341 in right posterodorsolateral view. “las” marks a ligament attachment site — a big, baseball-sized rugose lump — and right next to it is a pneumatic foramen, marked “pfo”.

Just like this, the ostrich caudal is a saurischian vertebra with a bifid neural spine, and with pneumatic foramina within the intermetapophyseal cleft.

From The Dinosaur Heresies.

Part 1.

This is an exciting day: the new PLOS Collection on sauropod gigantism is published to coincide with the start of this year’s SVP meeting! Like all PLOS papers, the contents are free to the world: free to read and to re-use.  (What is a Collection? It’s like an edited volume, but free online instead of printed on paper.)

There are fourteen papers in the new Collection, encompassing neck posture (yay!), nutrition (finally putting to bed the Nourishing Vomit Of Eucamerotus hypothesis), locomotion, physiology and evolutionary ecology. Lots every sauropod-lover to enjoy.

Taylor and Wedel (2013c: Figure 12). CT slices from fifth cervical vertebrae of Sauroposeidon. X-ray scout image and three posterior-view CT slices through the C5/C6 intervertebral joint in Sauroposeidon OMNH 53062. In the bottom half of figure, structures from C6 are traced in red and those from C5 are traced in blue. Note that the condyle of C6 is centered in the cotyle of C5 and that the right zygapophyses are in articulation.

Matt and I are particularly excited that we have two papers in this collection: Taylor and Wedel (2013c) on intervertebral cartilage in necks, and Wedel and Taylor (2013b) on pneumaticity in the tails of (particularly) Giraffatitan and Apatosaurus. So we have both ends of the animal covered. It also represents a long-overdue notch on our bed-post: for all our pro-PLOS rhetoric, this is the first time either of has had a paper published in a PLOS journal.

Wedel and Taylor (2013b: Figure 4). Giraffatitan brancai tail MB.R.5000 (‘Fund no’) in right lateral view. Dark blue vertebrae have pneumatic fossae on both sides, light blue vertebrae have pneumatic fossae only on the right side, and white vertebrae have no pneumatic fossae on either side. The first caudal vertebra (hatched) was not recovered and is reconstructed in plaster.

It’s a bit of a statistical anomaly that after a decade of collaboration in which there was never a Taylor & Wedel or Wedel & Taylor paper, suddenly we have five of them out in a single year (including the Barosaurus preprint, which we expect to eventually wind up as Taylor and Wedel 2014). Sorry about the alphabet soup.

Since Matt is away at SVP this week, I’ll be blogging mostly about the Taylor and Wedel paper this week. When Matt returns to civilian life, the stage should be clear for him to blog about pneumatic caudals.

Happy days!

References

• Taylor, Michael P., and Matthew 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 [PDF]
• Wedel, Mathew J., and Michael P. Taylor. 2013. Caudal pneumaticity and pneumatic hiatuses in the sauropod dinosaurs Giraffatitan and Apatosaurus. PLOS ONE 8(10):e78213. 14 pages. doi:10.1371/journal.pone.0078213 [PDF]

A few bits and pieces about the PLOS Collection on sauropod gigantism that launched yesterday.

First, there’s a nice write-up of one of our papers (Wedel and Taylor 2013b on pneumaticity in sauropod tails) in the Huffington Post today. It’s the work of PLOS blogger Brad Balukjian, a former student of Matt’s from Berkeley days. The introduction added by the PLOS blogs manager is one of those where you keep wanting to interrupt, “Well, actually it’s not quite like that …” but the post itself, once it kicks in, is good. Go read it.

Brad also has a guest-post on Discover magazine’s Crux blog: How Brachiosaurus (and Brethren) Became So Gigantic. He gives an overview of the sauropod gigantism collection as a whole. Well worth a read to get your bearings on the issue of sauropod gigantism in general, and the new collection in particular.

PLOS’s own community blog EveryONE also has its own brief introduction to the collection.

And PLOS and PeerJ editor Andy Farke, recently in these pages because of his sensational juvenile Parasaurolophus paper, contributes his own overview of the collection, How Big? How Tall? And…How Did It Happen?

Finally, if you’re at SVP, go and pick up your free copy of the collection. Matt was somehow under the impression that the PLOS USB drives with the sauropod gigantism collection would be distributed with the conference packet when people registered. In fact, people have to go by the PLOS table in the exhibitor area (booth 4 in the San Diego ballroom) to pick them up. There are plenty of them, but apparently a lot of people don’t know that they can get them.

References

• Wedel, Mathew J., and Michael P. Taylor. 2013. Caudal pneumaticity and pneumatic hiatuses in the sauropod dinosaurs Giraffatitan and Apatosaurus. PLOS ONE 8(10):e78213. 14 pages. doi:10.1371/journal.pone.0078213 [PDF]

“Look at all the things you’ve done for me
Opened up my eyes,
Taught me how to see,
Notice every tree.”

So sings Dot in Move On, the climactic number of Stephen Sondheim’s Pulitzer Prize-winning music Sunday in the Park with George, which on the surface is about the post-impressionist painter Georges Seurat, but turns out to be a study of obsession and creativity.

Un dimanche après-midi à l’Île de la Grande Jatte – 1884 [A Sunday Afternoon on the Island of La Grande Jatte – 1884]

In fact, the psychology of perception is complicated and sophisticated, and the brain does an extraordinary amount of filtering of the visual signals we get, to save us the bother of having to consciously process way too much data. This is a whole scientific field of its own, and I’m going to avoid saying very much about it for fear of making a fool of myself — as scientists so often do when wandering outside their own field. But I think it’s fair to say that we all have a tendency to see what we expect to see.

Phylogeny of Sauropoda, strict consensus of most parsimonious trees according to Wilson (2002:fig. 13a)

In the case of sauropods, this tendency has meant that we’ve all been startlingly bad at seeing pneumaticity in the caudal vertebrae of sauropods. Because the literature has trained us to assume it’s not there. For example, in the two competing sauropod phylogenies that dominated the 2000s, both Wilson (2002) and Upchurch et al. (2004) scored caudal pneumaticity as very rare: Wilson’s character 119, “Anterior caudal centra, pneumatopores (pleurocoels)”, was scored 1 only for Diplodocus and Barosaurus; and  Upchurch et al. (2004:286) wrote that “A few taxa (Barosaurus, Diplodocus, and Neuquensaurus) have pleurocoel-like openings in the lateral surfaces of the cranial [caudal] centra that lead into complex internal chambers”. That’s all.

And that’s part of the reason that every year since World War II, a million people have walked right past the awesome mounted brachiosaur in the Museum Für Naturkunde Berlin without noticing that it has pneumatic caudals. After all, we all knew that brachiosaur caudals were apneumatic.

But in my 2005 Progressive Palaeontology talk about upper limits on the mass of land animals estimated through the articular area of limb-bone cartilage, I included this slide that shows how much bigger the acetabulum of Giraffatitan is than the femoral head that it houses:

And looking at that picture made me wonder: those dark areas on the sides of the first few caudals (other than the first, which is a very obvious plaster model) certainly look pneumatic.

Then a few years later, I was invited to give a talk at the Museum Für Naturkunde Berlin itself, on the subject “Brachiosaurus brancai is not Brachiosaurus“. (This of course was drawn from the work that became my subsequent paper on that subject, Taylor 2009) And as I was going through my photos to prepare the slides of that talk, I thought to myself: darn it, yes, it does have pneumatic caudals!

So I threw this slide into the talk, just in passing:

Those photos were pretty persuasive; and a closer examination of the specimen on that same trip was to prove conclusive.

Meanwhile …

Earlier in 2009, I’d been in Providence, Rhode Island, with my Index Data colleagues. I’d managed to carve a day out of the schedule to hope along the coast to the Yale Peabody Museum in New Haven, Connecticut. My main goal was to examine the cervicals of the mounted Apatosaurus (= “Brontosaurus“) excelsus holotype (although it was also on that same trip that I first saw the Barosaurus holotype material that we’ve subsequently published a preprint on).

The Brontosaurus cervicals turned out to be useless, being completely encased in plaster “improvements” so that you can’t tell what’s real and what’s not. hopefully one day they’ll get the funding they want to take that baby down off its scaffold and re-prep the material.

But since I had the privilege of spending quality time with such an iconic specimen, it would have been churlish not to look at the rest of it. And lo and behold, what did I see when I looked at the tail but more pneumaticity that we thought we knew wasn’t there!

An isolated pneumatic fossa is present on the right side of caudal vertebra 13 in Apatosaurus excelsus holotype YPM 1980. The front of the vertebra and the fossa are reconstructed, but enough of the original fossil is visible to show that the feature is genuine. (Wedel and Taylor 2013b: Figure 10).

What does this mean? Do other Giraffatitan and Apatosaurus specimens have pneumatic tails? How pervasive is the pneumaticity? What are the palaeobiological implications?

Stay tuned! All will be revealed in Matt’s next post (or, if you can’t wait, in our recent PLOS ONE paper, Wedel and Taylor 2013b)!

References

• 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.
• 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.
• Wedel, Mathew J., and Michael P. Taylor. 2013. Caudal pneumaticity and pneumatic hiatuses in the sauropod dinosaurs Giraffatitan and Apatosaurus. PLOS ONE 8(10):e78213. 14 pages. doi:10.1371/journal.pone.0078213
• Wilson, J.A. 2002. Sauropod dinosaur phylogeny: critique and cladistic analysis. Zoological Journal of the Linnean Society 136:217-276.

This whole section, including the title, is mostly swiped from Mike’s Tutorial 17.

Other posts in this series are here.

Papers referenced in these slides:

• Farke, Andrew A., and Sertich, Joseph J.W. 2013. An abelisauroid theropod dinosaur from the Turonian of Madagascar. PLoS ONE 8(4): e62047. doi:10.1371/journal.pone.0062047 [PDF]
• Taylor, Michael P., Mathew J. Wedel and Richard L. Cifelli. 2011. A new sauropod dinosaur from the Lower Cretaceous Cedar Mountain Formation, Utah, USA. Acta Palaeontologica Polonica 56(1):75-98. doi: 10.4202/app.2010.0073
• Wedel, Mathew J., and Michael P. Taylor. 2013. Caudal pneumaticity and pneumatic hiatuses in the sauropod dinosaurs Giraffatitan and Apatosaurus. PLOS ONE 8(10):e78213. 14 pages. doi:10.1371/journal.pone.0078213 [PDF]

Here’s a working version of that link.

Working link.

Working links:

• Falkingham (2012) on photogrammetry for free
• Mallison photogrammetry tutorial 1
• Mallison photogrammetry tutorial 2
• Mallison photogrammetry tutorial 3
• Mallison photogrammetry tutorial 4

The rest of this series.

Reference

• Powell, Jaime E.  2003.  Revision of South American Titanosaurid dinosaurs: palaeobiological, palaeobiogeographical and phylogenetic aspects.  Records of the Queen Victoria Museum 111: 1-94.

Today for the first time I saw Saegusa and Ikeda’s (2014) new monograph describing the Japanese titanosauriform Tambatitanis amicitiae. I’ve not yet had a chance to read the paper — well, it’s 65 pages long — but it certainly looks like they’ve done a nice, comprehensive job on a convincing new taxon represented by good material: teeth, braincase, dentary, atlas, and as-yet unprepared fragmentary cervical, fragmentary dorsals, sacral spines, some nice caudals, some ribs and chevrons, and pubis and ilium.

What catches the eye immediately is the bizarre forward-curved neural spines of the anterior caudals:

Saegusa and Ikeda (2104: fig. 8): Tambatitanis amicitiae gen. et sp. nov., holotype (MNHAH D-1029280). A, Cd2–Cd11 in right lateral view. B, Cdx1–Cdx2 in right lateral view.

Here’s the third caudal in detail. (The first is fragmentary, and the second has some minor reconstruction near the tip of the spine which sceptical readers might think is covering up a misconstruction):

Saegusa and Ikeda (2014: fig. 11): Tambatitanis amicitiae gen. et sp. nov., holotype (MNHAH D-1029280). A–F, stereopairs of Cd3. A, right lateral view. B, left lateral view of the neural spine. C, anterior view. D, posterior view. E, dorsal view. F, ventral view. G, CT slices through the neural spine of Cd3, part corresponding to the matrix that filling the internal chamber is removed from the image. Greek letters in B and D indicate the position of CT slices shown in G. Scale bar = 10cm.

And here is the right-lateral view in close-up:

Saegusa and Ikeda (2014: fig. 11): Tambatitanis amicitiae gen. et sp. nov., holotype (MNHAH D-1029280) in right lateral view.

A phylogenetic analysis based on that of D’Emic (2012) recovers the new taxon in a polytomy with the Euhelopus clade that’s going to need a new name pretty soon, since it keeps growing and can’t be called Euhelopodidae for historical reasons: [that should probably be called Euhelopodidae: see discussion in comments]:

Saegusa and Ikeda (2014: fig. 23): Phylogenetic relationships of the titanosauriform sauropod Tambatitanis amicitiae gen. et sp. nov. from the Lower Cretaceous Sasayama Group of Tamba, Hyogo, Japan produced using the matrix of D’Emic (2012) with the addition of Tambatitanis. The final matrix, including 29 taxa and 119 characters, was analyzed in PAUP* 4.0b10. Left side, strict consensus of 81 most parsimonious trees (length = 207; CI = 0.609; RI = 0.8010; RC = 0.489), figures below nodes are decay indices. Right side, 50% majority rule consensus, figures above and below nodes represents the percentage of MPTs in which the node was recovered (only those relationships recovered in over 50% of the MPTs are shown).

Nice to see that new sauropods just keep on rolling out of the ground faster than we can blog about them!

References

• D’Emic, Michael D. 2012. The early evolution of titanosauriform sauropod dinosaurs. Zoological Journal of the Linnean Society 166:624-671.
• Saegusa, Haruo, and Tadahiro Ikeda. 2014. A new titanosauriform sauropod (Dinosauria: Saurischia) from the Lower Cretaceous of Hyogo, Japan. Zootaxa 3848(1):1-66. doi:10.11646/zootaxa.3848.1.1

Back in 2013, when we were in the last stages of preparing our paper Caudal pneumaticity and pneumatic hiatuses in the sauropod dinosaurs Giraffatitan and Apatosaurus (Wedel and Taylor 2013b), I noticed that, purely by chance, all ten of the illustrations shared much the same limited colour palette: pale brows and blues (and of course black and white). I’ve always found this strangely appealing. Here’s a composite:

I’m really happy with this coincidence. In fact I think I might get it printed up as a poster for my office.

(Thought: if I did, would anyone else be interested in buying it?)

Update (a couple of hours later)

At Matt’s suggestion, I switched the order of figures 7 and 8 (the last two on the third row) to get the following version of the image. It break the canonical order of the figures, but it’s visually more pleasing.

Now we should write an updated version of the paper that reverses the order in which we refer to figures 7 and 8 :-)

References

• Wedel, Mathew J., and Michael P. Taylor. 2013. Caudal pneumaticity and pneumatic hiatuses in the sauropod dinosaurs Giraffatitan and Apatosaurus. PLOS ONE 8(10):e78213. 14 pages. doi:10.1371/journal.pone.0078213