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.
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.
What heads tell us about necks, redux
June 5, 2009
I Cannot Brain Today, I Have the Dumb
Man, I hate making mistakes. The only thing worse than making mistakes is making them in public, and the only thing worse than that is finding them in published papers when it’s too late to do anything about them. About the only consolation left–if you’re lucky–is getting to be the one to rat yourself out (we have to do this a lot). So here goes.
In our figure 4 (from Taylor et al. 2009) we showed the skulls of three sauropodomorphs, Massospondylus, Camarasaurus, and Diplodocus, posed with horizontal semicircular canals (HSCCs) level, angled 30 degrees above horizontal, and angled 20 degrees below horizontal, as it is written (by Duijm 1951). We also showed the angle of the occipital condyle when the HSCCs are level; if the craniocervical joint was in osteologically neutral pose (ONP), that line would indicate the angle of the anterior cervicals.
Trouble is, we put the neck lines for Diplodocus and Camarasaurus in the wrong places.
As any idiot can see from Sereno et al. (2008: fig 1), the brain, brainstem, and occipital condyle form a line that runs from roughly the upper part of the orbit (in lateral see-through view) out the back of the head. Now if you look at our fig. 4 you’ll see that the ONP lines for Camarasaurus and Diplodocus are much too inclined, so that if the brain was in line with the anterior neck–which it should be, in ONP–it would be sticking out the back of the head.
If that doesn’t make sense, just look at the above illustration, imagine the brain and spinal cord in a straight line parallel to the black neck line but also dorsal to it, and you’ll see that the brain would be outside the skull. Those incorrect neck lines don’t represent impossible postures, but they don’t represent ONP, either.
Here’s a corrected up version of the figure to show what I mean. The black lines are still the ONP neck lines, and now I’ve put in shadowy necks at +30 and -20 to go with the shadowy heads. The 50 degree spans marked out by the shadowy necks are the ranges within which the neck could articulate in ONP with skulls stuck in the 50-degree “Duijm window”.
Caution: it is very easy to misread the shadowy necks as showing a range of movement within an individual; in fact, the neck lines are ‘anchored’ to the skulls in ONP as the skulls rotate through the 50 degrees allowed by the HSCCs. They are not individual movement but the possible range of taxonomic variation in HSCC orientation according to Duijm (1951).
Worth noting here is the likelihood that Massospondylus had a more elevated neck than any of the neosauropods studied so far–certainly a finding at odds with the traditional depictions of basal sauropodomorphs. (It is just a likelihood, though, since the top, neck-wise, of Massospondylus‘s Duijm window overlaps with the windows of the other taxa a bit.)
In this version I’ve gone one step farther and included Nigersaurus (modified from Sereno et al. (2008: fig 1). Nigersaurus differs from Diplodocus in the angle of the face from the HSCCs and occipital condyle, not in the angle between the HSCCs and the occipital condyle, which is remarkably similar in Camarasaurus, Diplodocus, and Nigersaurus. This suggests that Nigersaurus held its head differently than other sauropods, but not necessarily its neck.
Keep in mind, though, that the difference in facial angle between Diplodocus and Nigersaurus is less than 50 degrees, and that some of the head postures in the respective Duijm windows of the two taxa are identical. So we can’t say for certain that Nigersaurus held its head differently than Diplodocus; it is possible that they held their heads at the same angle and that Nigersaurus just carried its HSCCs at a different angle. If that were the case, the neck of Nigersaurus would have been more inclined than that of Diplodocus. I’m not arguing that that’s likely–it seems perfectly plausible that the two taxa might have held their necks similarly and their heads differently, as suggested above–I’m just pointing out the very wide range of possibilities allowed by the data. To reiterate one of the points of the paper, HSCCs aren’t useless for determining habitual head posture, they just can’t narrow things down very far on their own.
Also note that some of the neck postures allowed by the Duijm window have the anterior cervicals running down, below horizontal, not up. And many of the allowed neck postures for the neosauropods are close to horizontal. So, we were wrong and HSCCs + occipital condyles show that most sauropods held their necks close to level and not strongly elevated after all, right?
Onward and Upward, or Down in Flames?
Not so fast. Remember that all of the neck lines in the above figures show the angle of the anterior neck if the neck was in ONP with the skull. But Vidal et al. (1986) found that the skull is habitually flexed on the neck, even in lizards, and we have since verified this for salamanders, turtles, and more. And sometimes the flexion is dramatic.
Our figure 1 (from Taylor et al. 2009) shows the cranium, cervicals, and first few dorsals from a hare in ONP and in the posture shown by Vidal et al. (1986: fig. 4b). The difference between the anteriorly-directed ONP pose and the backward-leaning Vidal-compliant pose is striking. I measured the angle between the cervical column and the maxillary toothrow to be ~110 degrees in the ONP pose and ~70 degrees in the Vidal-compliant pose (try it yourself with Paint or Photoshop, or download some free image manipulation software). That means the head is flexed on the neck by 40 degrees! That is a big angle. If sauropods did the same, you could take the neck lines shown above and crank them down by 40 degrees (remember that the heads are “fixed” into the 50-degree Duijm windows allowed by the HSCCs), which would make Mike’s elevated Diplodocus look not just achievable, but perhaps even conservative.
Where does all that leave us? In sauropods for which HSCC orientation is known, putting the HSCCs level the anterior neck is still inclined, and even with the HSCCs angled 20 degrees down the ONP neck would only be slightly below horizontal, and if the head was Vidal-compliant (strongly flexed on the neck), the neck would have to be above horizontal. So heads still tell us about necks, and in particular they tell us that the necks angled up. Our neck lines for Camarasaurus and Diplodocus are not correct for ONP, but probably represent attainable postures. My first head ‘n necks post has the angles too exaggeraged for ONP, too, but again all of those poses are not just possible but likely if the head was flexed on the neck.
Miscellanea
We owe mad props to Brian Engh, a.k.a. The Historian, who burst on the paleo-rap scene with a rap video about crocodilian predation and almost certainly the first ever kung-fu rap video to name-check titanosaurs. Brian stumbled across Mike’s extra goodies page for the new paper about week before the paper was due out, and kindly suppressed the information until after D-Day. You can and should download his entire album, Earth Beasts Awaken (open access, yo), and kick it old school.
Congratulations to Francisco “Paco” Gasco, who just got funding for a PhD to do a complete morphological and paleobiological workup on the giant Spanish sauropod Turiasaurus. You’ll be hearing more about Paco in the not-too-distant future, we promise.
Finally, here’s that video of an elephant grabbing an ostrich by the neck that you ordered.
The End of the Beginning?
This brings us to the end of ten solid days of new posts, which is a new record for us and one not likely to be broken for a long time, if ever. We never planned to do all this; in the beginning we each were going to contribute one post and that would have been that. But we kept finding things that we felt needed to be discussed.
As all of us have been saying in every available medium, this is not the end of anything. The sauropod neck posture debate is not over; in a few years we may look back and see that in 2009 we were still stumbling to the real starting line. We don’t think this stuff is unimportant or unknowable, and we’re going to keep working on it, and we hope lots of others do as well.
We’ll see you out there.
References
- Duijm, M. 1951. On the head posture in birds and its relation to some anatomical features. II. Proceedings of the Koninklijke Nederlandse Akademie van Wetenschappen, Series C 54: 260–271.
- 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, M.P., Wedel, M.J., and Naish, D. 2009. Head and neck posture in sauropod dinosaurs inferred from extant animals. Acta Palaeontologica Polonica 54 (2): 213–220.
SuperCroc’s sidekick
June 15, 2008
Paul Sereno’s Project Exploration has a traveling exhibit called The Science of SuperCroc, which I recently visited at my old stomping grounds, the Oklahoma Museum of Natural History. The exhibit focuses on Sarcosuchus, the improbably large and possibly Kryptonian crocodilian from the Cretaceous of Niger, but it also includes nice mounted skeletons of the spinosaur Suchomimus and–relevant to our purposes here–the just plain improbable sauropod Nigersaurus.
At least one of my co-bloggers probably thinks I should stop pandering to the crowds with my mounted skeleton posts and get back to hardcore vertebral anatomy. After all, that’s the raison d’etre of SV-POW!, and I have been falling behind a little lately. Still, I’m going to risk the Wrath of Mike and go ahead and post about the mounted Nigersaurus skeleton, and why you should definitely go see it if you get the chance.
Here are my reasons for doing so:
1. It is a demonstrated scientific fact, as rock-solid as the value of c or the proposition that the Amazon basin is damp, that Nigersaurus is Damn Weird. In a clade of little-known weirdos (Rebbachisauridae), it promises to be an exceedingly well-known ultra-weirdo, thanks to (1) the large number of skeletons that have been discovered, including both juveniles and adults, and (2) the sheer vastness of its weirdness, which you can sample immediately and without charge courtesy of Sereno et al. (2007) and the kind offices of the Public Library of Science (translation: free paper here).
2. Although Nigersaurus was named in 1999 and has been the subject of three peer-reviewed publications, not much of the skeleton has been figured to date. So the opportunity to see the whole critter up close is pretty remarkable. If sauropods were heavy metal, the traveling Nigersaurus mount would be an evening backstage getting high with Led Zeppelin, circa 1973. Certainly if you work on sauropods, the morphology of Nigersaurus will make you think that someone has been under the influence of powerful illicit substances, and that someone is Mother Nature (or Gaia, or the overused/sexist/quasi-pantheistic biosphere personification of your choice).
3. It’s a really nice mount. It’s fiberglass, but the quality of the casts is first rate. I have seen a lot of traveling skeletons that looked like they were made out of Play-Doh by speed-sculpting chimps, but the mounts in the SuperCroc exhibit are all well cast, gracefully mounted, and nicely displayed, by which I mean that you can get up close to them and walk most or all of the way around them, which is my major pet peeve about mounted skeletons: I want to be able to see them from any angle, or at least many angles. SuperCroc delivers.
4. The exhibit includes a lot of display cases that explain the detailed anatomy of the beasts. For Nigersaurus alone, there were cases on vertebral pneumaticity (yay!), the vertebrae themselves (real bones), the detailed anatomy of the jaws (real bones, from the holotype!), the head and neck skeleton plus life sculpture (shown at the top), adult and baby femora (real bones), probable feeding ecology, and maybe one or two others I can’t remember, plus a giant wall hanging of the full-color life restoration painting that came out with the 2007 paper.
So if you get a chance to see SuperCroc, it’s worth it just for the sauropod.
I’ll have tons more to say about the Nigersaurus vertebrae in future posts, but the short version is that they are small and unbelievably delicate. Mike and I always characterize Camarasaurus vertebrae as coarse, fat, and kind of ugly; the vertebrae of Nigersaurus are the aesthetic opposite. They look like they might have been constructed out of toothpicks and white glue. And they are crazy pneumatic. In one of his essays, outdoor humorist Patrick F. McManus characterized a poorly-maintained country bridge as consisting mostly of holes that were elevated and loosely defined by a few rotting beams. Similarly, the skull and cervical vertebrae of Nigersaurus seem to be mostly holes, with just enough bone around them to suggest the form of a sauropod. One more half-baked comparison: the mounted Nigersaurus looks like the skeleton of a skeleton, at least in the craniocervical region.
I want to make a final point that is not really about vertebrae. As you can see in the photo above, and to beat a dead thunder-lizard, sauropods had erect limbs, compact feet, and deep, slab-sided bodies. You don’t have to be a zoologist to see that this is a body-form made for roaming the land, not for bobbing around slurping up pond scum. That’s not to say that sauropods didn’t go into the water. They probably did so all the time. Elephants do, almost every chance they get. Heck, elephants may even be descended from aquatic ancestors. But no one would characterize elephants as aquatic or even semi-aquatic. Sauropods weren’t elephants, and they weren’t giraffes, and facile comparisons of sauropods to big mammals have probably done more harm than good to sauropod paleobiology. But sauropods weren’t hippos or manatees, either, despite decades of ecological characterization as such. The Aquatic Sauropod era officially ended the same year I was born, so you may rightly wonder why I am tilting at this particular windmill. It’s because ideas are seductive, and sometimes we allow them to make us blind to the obvious. I don’t know of any way to fight that tendency other than to keep asking questions.
And I don’t know of a sauropod that is more likely to provoke questions than Nigersaurus. Go see it if you can.