Things to Make and Do, part 3: Butchering a Wallaby

This is part 3 of an emerging and occasional SV-POW! series: part 1 was the pig skull, and part 2 was the lizard feet (though not advertised as such because I couldn’t resist the sauropod pun).

Bennett's wallaby, right lateral view

Today, we’re going to be taking a wallaby apart.  Specifically, a Bennett’s wallaby, the larger of the two subspecies of the red-necked wallaby Macropus rufogriseus.  I was delighted (though of course also saddened) to get a call on Saturday afternoon from the very same mini-zoo that had given me Charlie the monitor — Dick Whittington Farm Park in Longhope, Gloucestershire.  They have a small group of seven wallabies sharing a paddock with goats, and one had died — most likely from being butted by one of the goats, although there were no external signs of injury.

This is going to be the largest animal I’ve prepared the skeleton out of — I measured it at 123 cm from snout to tail and 10.5 kg total weight, which compares with 75 cm and 12 kg for the badger, 100 cm and 5.2 kg for the fox and 111 cm and 3.4 kg for the monitor.  Yes, the badger was heavier, but the awkward shape of the wallaby makes it all-round “bigger” and harder to deal with.  Both the badger and the fox would, just, fit into large plastic toy-boxes which I buried and will exhume after a suitable time has passed, but that wasn’t going to work for the wallaby.  I needed to take that baby apart:

Bennett's wallaby, right ventrolateral view into guts

I was pleasantly surprised at what good condition the guts were in (compared with the horrible state of Charlie innards) — nice and fresh.  If I’d had time, I’d have attempted to learn something from a proper dissection, but as I was pushed for time (trying to get this done in my lunch break) I had to push on.  I discarded the guts and started to carve up the remainder.

Bennett's wallaby in posteroventral view. right leg removed; Homo sapiens for scale

The knife is a Norwegian fisherman’s knife — very sharp, and short enough to be easy to wield.  It’s perfect for dismembering a carcass this size, even though previously I’ve only used it for slicing sushi rolls.  It was a Christmas present from my employer, Index Data, a few years ago.

My plan was to carefully divide the animal into seven portions (head, torso, tail and four legs), remove as much skin and muscle as I could without risking damage to the bone, and to process the parts separately.

Bennett's wallaby, in kit form, mostly dorsal view but with the head and torso in left lateral. WARNING: GRAPHIC CONTENT

After some thought, I decided to prepare the skull and the left fore- and hind-limb by boiling, and to bury the rest in the box.  Here are the relevant divisions:

Bennet's wallaby not looking at all healthy. Top: torso, tail and right fore- and hindlimbs, awaiting burial. Bottom left: head, left fore- and hindlimbs, awaiting cooking. Bottom right: bag full of discarded soft-tissue

Then I put the pot through an hour’s simmering, peeled the skin off the skull and feet, and removed what meat I could; then I simmered a second time and removed more meat.  By this stage, I was able to remove the three most anterior cervicals, which had been attached to the back of the skull — but they are still so covered with attached flesh that they’re not much use yet.  Here’s how the simmered material is looking:

Bennett's wallaby: skull, anterior cervical vertebrae and left hind-limb long bones

And here is the skull as it looks now, after a little more flesh-picking (but not nearly enough):

Bennett's wallaby, partially prepared skull in right lateral view

I think that it (and the other boiled bones pictures above) would benefit from a third simmer-and-pick session before I put them out somewhere for invertebrates to deal with.  While that’s going on, I’ll prep out the foot and the forelimb, which have also been boiled twice but phalanges are a right nasty piece of work.

And then I have to decide what to do with my big yellow box that has the rest of the bits in.  Plan A is still burying, but it is kind of tempting to simmer these parts, too, and get the whole thing completed much more quickly.

On the other hand, now is not a good time for such an effort: I will be away from home all week on a mission of utmost importance, and of great relevance to this blog.  Details to follow!

Finally, I leave you with your weekly sauropod-vertebra goodness!

Giraffatitan brancai paralectotype HMN SI, cervical vertebrae 2 and 3 in right lateral view, attempting to do DinoMorph

Finite Element Analysis of sauropod vertebrae

Figure 3 from Schwarz-Wings et al. 2009. A is Diplodocus, B-D are Giraffatitan.

Earlier this month Daniela Schwarz-Wings and colleagues published the first finite element analysis (FEA) of sauropod vertebrae (Schwarz-Wings et al. 2009). Above is one of the figures showing some of their results. Following standard convention, stresses are shown on a gradient with cooler colors indicating lower stresses and hotter colors indicating higher stresses. I’m not going to dwell on the on the nuts-n-bolts of FEA in general or of this study in particular. Instead, I want to talk about how sauropod vertebrae are built.

CT cross sections of BYU 12866, a mid-cervical of Brachiosaurus sp.

In cross-section, sauropod vertebrae often have thick bone at the outer edges of the laminae and in the walls and especially the floor of the centrum, as shown in this Brachiosaurus cervical. The bone everywhere else is pretty thin. If you hit one of these vertebrae with some magical forumula that would dissolve away all the bone thinner than, say, 1 cm, all that would be left would be the various apophyses, the outer margins of the laminae connecting them, and probably the bottom half of the centrum. It would be like the outline of a vertebra constructed from tent poles, or tinkertoys.

This is weird because most pneumatic sauropod vertebrae have at least something approaching an I-beam shape in cross-section. You might think that the median septum would be mechanically important, but it’s usually very thin, sometimes perforated (see Hatcher’s [1901] Diplodocus cervicals, for example), and often asymmetrically deviated to one side or the other. Not what you would expect for a piece of bone that was doing any work.

And indeed, Schwarz-Wings et al. (2009) found that:

Comparative stresses are distributed evenly around the vertebrae and mainly on the bone cortex. Peak stresses occur only at points where the tendons and muscles are inserting because the insertion areas used were small resulting in extreme localized stresses. The interior of both vertebrae is nearly stress free. Almost no stresses occur around the cavities and in their bony walls (figure 3).

This reminds me not of I-beams but of the long bones of the limbs of terrestrial vertebrates. There’s a reason why you’ve got a big honkin’ marrow cavity running through the middle of your femur: the stresses are being borne by the walls of the bone. It makes sense that vertebrae would function similarly, especially sauropod cervicals which sometimes approximate limb bones in their proportions.

So how about that median septum? Why aren’t sauropod vertebrae just hollow tubes? My guess–and it is a guess–is that they got as close to being hollow tubes as their evolutionary and developmental origins allowed. The pneumatic diverticula invaded the centra from either side and pushed in lateral-to-medial, and I think the median septum is just the wimpy little bit of bone left in between the two sets of diverticula when they almost meet up in the middle.

Even if that’s correct, there’s another mystery: why don’t the diverticula just go ahead and erode away the median septum? I can think of two possible reasons. One is that, for reasons I don’t know and I’m not sure if anyone else does either, pneumatic diverticula are good at getting into bones but pretty lousy at getting back out. There are comparatively few cases of diverticula inside bones making foramina to get out into the  surrounding tissue. It does happen–in humans, the mastoid air cells sometimes bust out and make subcutaneous pneumatocoels, basically bubbles of air under the skin (Anorbe et al. 2000)–but it seems to be rare. Maybe median septa fall under the same inscrutable rule.

(Incidentally, this makes the perforate laminae in Giraffatitan all the weirder.)

Another, more mundane possibility is that the median septa (and other oddly thin bits of bone) are not never loaded, just infrequently loaded. Not enough to make them straight, thick, or normal-lookin’, but enough to make sure they don’t get resorbed entirely.

Sauropod vertebrae are just loaded with these growth-and-form-related mysteries. Kudos to Schwarz-Wings et al. for pushing us a little farther down the road toward solving them.

References

• Anorbe, E., Aisa, P. and Saenz de Ormijana, J. 2000. Spontaneous pneumatocele and pneumocephalus associated with mastoid hyperpneumatization. European Journal of Radiology 36:158–160. [abstract only for free]
• 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 and plates I-XIII.
• Schwarz-Wings, D., Meyer, C.A., Frey, E., Manz-Steiner, H.-R., and Schumacher, R. 2009. Mechanical implications of pneumatic neck vertebrae in sauropod dinosaurs. Proceedings of the Royal Society B. doi: 10.1098/rspb.2009.1275

How big were the biggest sauropod trackmakers?

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.

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.

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? 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?

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). 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.

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.

Small African primate possibly sheds light on soft-tissue morphology of Cretaceous diplodocoid

One of the most bizarre of sauropods – and arguably one of the most bizarre of dinosaurs – is the Patagonian dicraeosaurid diplodocoid Amargasaurus cazaui Salgado & Bonaparte, 1991. Here’s a picture of a replica specimen, provided by Nizar Ibrahim. You’ll note that this skeleton has been posed with an extended cranio-cervical junction, and a slightly extended cervico-dorsal junction: virtually the opposite of what you’d expect in ‘normal’ posture (Taylor et al. 2009).

Thanks to its appearance in virtually every post-1991 book on dinosaurs, Amargasaurus is a relatively familiar animal these days, which is sort of a shame: we’ve somehow forgotten how frikkin’ weird it is. We’ve looked at Amargasaurus on SV-POW! before: in this article we drew attention to the fact that its dorsals are pretty interesting too, and as usual with sauropods we could write whole essays on all manner of its vertebral osteology. Perhaps, in time, we will.

However, let’s be honest and note that it’s those ridiculous cervical spines that keep us awake at night. What the hell did the animal do with these structures, and exactly what did it look like when alive? Early reconstructions (most notably Brian Franczak’s drawing from 1992 (see Hecht 1992)) all went with the idea of two paired sails, running in parallel along the length of the neck. All artists thereafter followed suit. Until 1994, when Greg Paul noted that the circular cross-sections of the spines and tapering tips might suggest that the paired neural spines instead protruded from the neck as horn-covered spikes (Paul 1994). Greg also suggested that having paired sails along the neck might be a really bad idea as it could restrict neck mobility, but this idea never made much sense as the tissue between the spines would presumably be flexible and hence not such a problem (the circular cross-section is not really a problem either, since other animals that are thought to have had skin sails, like Dimetrodon, also have rounded neural spines). Since 1994 few opinions have been expressed on this subject, but those that have have sided with Greg in noting that spines are more likely than sails (Salgado 1999).

At SVP Bristol last month, your favourite blog authors spent many happy hours discussing sauropods and their vertebrae. And, inevitably, the subject of Amargasaurus and its weird cervical spines came up. Fact is, sails AND spines are both equally stupid, and the idea of independent neck spines is particularly stupid because animals just don’t ever have their neural spines poking out beyond the musculature and skin of the neck. Right?

Wrong. Hardly known and rarely mentioned is that the last four (or so) cervical neural spines and first three thoracic spines of the Potto Perodicticus potto (a peculiar African strepsirrhine primate, closely related to the Asian lorises) protrude from the skin of the neck and shoulders, have sharply pointed tips, and are usually stated to serve a defensive function [potto skeleton above provided by Mo Hassan]. The spines don’t protrude as naked bone (though this was claimed by Ivan Sanderson), but are covered with skin and some hair, and form a series of tubercles in the live animal. When threatened, a potto is supposed to hide its head and point those protruding neck spines towards the danger, sometimes even butting the predator or aggressor with the spines. Other functions have been suggested for the spines, but a defensive function remains most popular and in the absence of any special knowledge I’ll go with the majority.

At left, Potto with flexed neck, showing its cervical and thoracic spines. From Walker (1970). At right, Amargasaurus as reconstructed by Salgado (1999). Salgado imagined this to be the animal's normal posture: an idea that we find somewhat unlikely (see Taylor et al. 2009).

You might wonder, quite reasonably, whether the anatomy of a small arboreal modern primate has any bearing whatsoever on the soft-tissue morphology of a Cretaceous sauropod (even a ‘small’ one). The point is, however, that neural spines can protrude from the neck after all (albeit sheathed in skin and other tissue), and hence there is at least some precedent for this sort of thing. All that’s needed now is for someone to study the histology and microscopic surface texture of those protruding potto spines and see if there are any correlates for their protruding nature (so far as I can tell this hasn’t been done). Then go look at Amargasaurus. There’s clearly a paper in this, please report back and include me on the authorship, thanks.

References

• Hecht, J. 1992. Dollars for dinosaurs. New Scientist 136 (1845), 50.
• Paul, G. S. 1994. Dinosaur art & restoration notes: dicraeosaurs. The Dinosaur Report Summer 1994, 8.
• Salgado, L. 1999. The macroevolution of the Diplodocimorpha (Dinosauria; Sauropoda): a developmental model. Ameghiniana 36, 203-216.
• Taylor, M. P., Wedel, M. J. & Naish, D. 2009. Head and neck posture in sauropod dinosaurs inferred from extant animals. Acta Palaeontologica Polonica 54, 213-220.
• Walker, A. 1970. Nuchal adaptations in Perodicticus potto. Primates 11, 135-144.

Electronic publishing is inevitable and even the ICZN is beginning to accept it

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:

Background: the availability of the name Darwinius masillae

The Code is in danger of becoming an irrelevance

Paper journals are going away

The time to act is now

Electronic documents are different from electronic media

We must come to terms with the ubiquity of PDF

The current rules are too hard to get right

Conclusion

Background: the availability of the name Darwinius masillae

The Code is in danger of becoming an irrelevance

Paper journals are going away

The time to act is now

Electronic documents are different from electronic media

We must come to terms with the ubiquity of PDF

The current rules are too hard to get right

Conclusion

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.

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.

What a 23% longer torso looks like

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

Thought not.  Right, then, here we go!

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

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

(Yes, brachiosaurs probably had 12 dorsals.)

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

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

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

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

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

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

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

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

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

Meanwhile some good news:

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

And finally …

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

References

• Janensch, Werner.  1950.  Die Wirbelsaule von Brachiosaurus brancai.  Palaeontographica (Suppl. 7) 3: 27-93.
  Paul, Gregory S.  1988.  The brachiosaur giants of the Morrison and Tendaguru with a description of a new subgenus, Giraffatitan, and a comparison of the world’s largest dinosaurs.  Hunteria 2 (3): 1-14.
  Taylor, Michael P.  2009.  A re-evaluation of Brachiosaurus altithorax Riggs 1903 (Dinosauria, Sauropoda) and its generic separation from Giraffatitan brancai (Janensch 1914).  Journal of Vertebrate Paleontology 29(3):787-806.
  Vesalius, A.  1543.  Andreae Vesalii Bruxellensis, Scholae medicorum Patauinae professoris, de Humani corporis fabrica Libri septem [facsimile]. Ex Officina Ioannis Oporini, Basel, 659 pp.
  Wilson, Jeffrey A.  2006.  Anatomical nomenclature of fossil vertebrates: standardized terms or “lingua franca”?  Journal of Vertebrate Paleontology 26(3): 511-518.
• Janensch, Werner.  1950.  Die Wirbelsaule von Brachiosaurus brancai.  Palaeontographica (Suppl. 7) 3: 27-93.
• Paul, Gregory S.  1988.  The brachiosaur giants of the Morrison and Tendaguru with a description of a new subgenus, Giraffatitan, and a comparison of the world’s largest dinosaurs.  Hunteria 2 (3): 1-14.
• Taylor, Michael P.  2009.  A re-evaluation of Brachiosaurus altithorax Riggs 1903 (Dinosauria, Sauropoda) and its generic separation from Giraffatitan brancai (Janensch 1914).  Journal of Vertebrate Paleontology 29(3):787-806.
• Vesalius, A.  1543.  Andreae Vesalii Bruxellensis, Scholae medicorum Patauinae professoris, de Humani corporis fabrica Libri septem [facsimile]. Ex Officina Ioannis Oporini, Basel, 659 pp.
• Wilson, Jeffrey A.  2006.  Anatomical nomenclature of fossil vertebrates: standardized terms or “lingua franca”?  Journal of Vertebrate Paleontology 26(3): 511-518.

Bifid Brachiosaurs, Batman!

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

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

References

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

And finally … two announcements!

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

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

Right, that’s it — time for the revolution

UPDATE (from Matt): I also bring good news … and bad news.

The good news is that the entire dinosaur issue of Anatomical Record is open access after all. So this post is mainly of historical interest now, and you should get on over to the page for this issue and download all the free dinosaurian goodness.

The bad news is that the representatives from Wiley never told anyone any of this when inquiries were made two weeks ago–if they had, this particular teacup could have stayed storm-free–and that they apparently still want institutions to pay $575 for a single Open Access issue of the journal. Whether those moves are predatory or just clueless, they are not earning Wiley any friends.

—————-

I bring good news … and bad news.

Good news! Tom Holtz reported in a message to the Dinosaur Mailing List that there is new issue of The Anatomical Record out that is concerned entirely with dinosaurs!  The online table of contents shows that there’s lots of good stuff.

Bad news! It’s not open access.

Good news! You can buy access to the articles.

Bad news! The price of the articles is NOT STATED.  That’s right, folks: you have to register with Wiley InterScience before they will EVEN TELL YOU THE PRICE!  Way to go, Wiley!  THAT’s the way to make sure important research is widely disseminated!

Good news! B tH wrote to ask the publisher for a price, and got a reply, which he shared in another Dinosaur Mailing List message:

Bad news! This is the reply (which I can’t format better, thanks to totally unnecessary limitations in WordPress):

Date: Mon, 31 Aug 2009 12:48:21 -0700 (PDT)

From: B tH <soylentgreenistrex@yahoo.com>

To: dinosaur@usc.edu

Subject: re: special all-dino issue

I wrote to ask them how much ordering this singl issue was – they wanted to know if I was ordering for an institution or myself. This is the price they quoted me to buy and read it at night with a flashlight under the blankey – and I am totally serious:

$575.00 US

That’s right, five HUNDRED and seventy-five buckeroos.   I assured them they were quite mad, and have to face the fact I won’t get to see it.   Waaah.

Good news! B tH realised that Wiley had quoted him the institutional rate and wrote to clarify.  The exchange is documented in yet another Dinosaur Mailing List message.

Bad news! This is the exchange:

Sent: Monday, August 31, 2009 6:07 PM

To: cs-journals@wiley.com

Subject: RE: wanting to purchase an issue of the magazine [pfCase:1078353,

pfTicket:10108736]

Um, I think you’ve made an error.

Five-Hundred and Seventy-Five dollars for an issue of a magazine?  ??

==============

From: <cs-journals@wiley.com>

Dear __________

The Anatomical Record, Volume 292, Issue 9

Thank you for your email.

As we do not have Individual rates for this title, hence the Institutional single issue rate was quoted instead.

Please provide us with a billing and shipping address if you require a proforma invoice for this order and I will happy to assist you.

Kind Regards,

Jacqueline Choong

Customer Services Advisor

Journal Customer Services for John Wiley & Sons

Good news! The revolution is coming, and things like this can only bring it on.  And Wiley’s InterScience department are a bunch of mindless jerks who will be first up against the wall when the revolution comes.

Yes, Wiley’s behaviour here is totally absurd and absolutely unethical.  No, Wiley didn’t themselves write the articles that they want to charge FIVE HUNDRED AND SEVENTY-FIVE FREAKIN’ DOLLARS for.  Neither did they pay the authors to do so.  Do you know how it comes to be that Wiley are the owners of these articles, and thus in a position to extort for access?  Happily, the reason is right here in the Instructions to Authors:

MISCELLANEOUS

[...]

Upon acceptance of an article for publication, the author will be asked to sign a Copyright Transfer Agreement transferring rights to the publisher, who reserves copyright.

Yes, it’s as simple as that.  Like all of us do most times we submit a manuscript, the authors just signed away the ownership of their work.  Just like that.  Work that was funded, if at all, by public funds, just handed over to a grossly exploitative for-profit commercial enterprise that — quite clearly, from the exchanges above — has no interest whatsoever in the advancement or dissemination of science.

Folks, we have got to stop doing this.  I can (just) stomach handing copyright of my work over to professional societies such as the Society of Vertebrate Paleontology (required for the Journal of Vertebrate Paleontology) or the Palaeontological Association (required for Palaeontology) [although frankly there is absolutely no good reason for these journals to make that requirement].  But I will NOT give my work to these parasitic commercial publishers, and I strongly urge you not to, either.  We should all of us be supporting open-access journals where possible; and failing that, at least those published by non-profit organisations.  I am not going to be propping up Elsevier, Wiley and the rest with any of my stuff.

Deep in our heart, we all — Wiley included — know that non-open academic publishing is dead, even if the corpse is still blundering around trying to eat our brains.  This sort of extortion (I mean the FIVE HUNDRED AND SEVENTY-FIVE FREAKIN’ DOLLARS kind) is death throes.  It’s probably going to get messier before the stakes are finally driven through the hearts of the bloodsuckers.  But take heart: morning is coming, and they will all turn to dust.

And finally …

More Good news! I give you NHM 46869, the holotype of Chondrosteosaurus gigas Owen 1876, a badly eroded cervical centrum from some kind of sauropod, in right lateral view:

NHM 46869, holotype of Chondrosteosaurus gigas, a cervical centrum, in right lateral view.

This is the mate of NHM 46870, a specimen that we have already given way too much coverage, and which has sometimes been considered the cotype along with 46869.  Unlike its mate, it has not been sliced down the middle, and is — for what it’s worth — “complete” (i.e. not actually complete at all).

References

• Owen, Richard.  1876.  Monograph of the fossil Reptilia of the Wealden and Purbeck formations.  Supplement 7.  Crocodilia (Poikilopleuron), Dinosauria (Chondrosteosaurus),  Palaeontographical Society of London [Monographs], 29:15-93.

This is the reply:

Date: Mon, 31 Aug 2009 12:48:21 -0700 (PDT)

From: B tH <soylentgreenistrex@yahoo.com>

To: dinosaur@usc.edu

Subject: re: special all-dino issue

I wrote to ask them how much ordering this singl issue was – they wanted to know if I was ordering for an institution or myself. This is the price they quoted me to buy and read it at night with a flashlight under the blankey – and I am totally serious:

$575.00 US

That’s right, five HUNDRED and seventy-five buckeroos.   I assured them they were quite mad, and have to face the fact I won’t get to see it.   Waaah.

Good news!  B tH realised that Wiley had quoted him the institutional rate and wrote to clarify.  The exchange is documented in yet another Dinosaur Mailing List message.

Bad news!  This is the exchange:

Sent: Monday, August 31, 2009 6:07 PM

To: cs-journals@wiley.com

Subject: RE: wanting to purchase an issue of the magazine [pfCase:1078353,

pfTicket:10108736]

Um, I think you’ve made an error.

Five-Hundred and Seventy-Five dollars for an issue of a magazine?  ??

==============

==============

From: <cs-journals@wiley.com>

Dear __________

The Anatomical Record, Volume 292, Issue 9

Thank you for your email.

As we do not have Individual rates for this title, hence the Institutional single issue rate was quoted instead.

Please provide us with a billing and shipping address if you require a proforma invoice for this order and I will happy to assist you.

Kind Regards,

Jacqueline Choong

Customer Services Advisor

Journal Customer Services for John Wiley & Sons

Good news!  The revolution is coming, and things like this can only bring it on.  And Wiley’s InterScience department are a bunch of mindless jerks who will be first up against the wall when the revolution comes.

Yes, Wiley’s behaviour here is totally absurd and absolutely unethical.  No, Wiley didn’t themselves write the articles that they want to charge FIVE HUNDRED AND SEVENTY-FIVE FREAKIN’ DOLLARS for.  Neither did they pay the authors to do so.  Do you know how it comes to be that Wiley are the owners of these articles, and thus in a position to extort for access?  Happily, the reason is right here in the Instructions to Authors:

MISCELLANEOUS

[...]

Upon acceptance of an article for publication, the author will be asked to sign a Copyright Transfer Agreement transferring rights to the publisher, who reserves copyright.

###

How big was Alamosaurus?

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

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

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

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

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

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

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

Reference

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

Tutorial 7: the sauropod family tree

We really should have covered this ages ago …  Here we are, blithering on about brachiosaurids and diplodocoids and all, and we’ve never really spelled out what these terms mean.  Sorry!

The family tree of a group of animals (or plants, or fungi, or what have you) is called its phylogeny.  The science of figuring out a phylogeny is called systematics.  And once you’ve got a phylogeny, the business of naming the parts of it (and of course choosing which parts to name) is taxonomy.

For a long time, sauropod systematics was completely up in the air, so that the McIntosh’s (1990) review article on sauropods in The Dinosauria (first edition) said, rather despairingly, that “although recent discoveries are beginning to clarify the problems of sauropod phylogeny, were are still very far from being able to construct a cladogram” (p. 399).  Happily, this changed rapidly thereafter, with the first published numerical phylogenetic analysis appearing in Russell and Zheng’s (1993) description of the new Mamenchisaurus species M. sinocanadorum.  More importantly, in the same year Paul Upchurch submitted his (1993, duh) dissertation on sauropods, and this contained a much larger analysis which was published as Upchurch (1995).  This paper raised the bar significantly, with an analysis of 27 taxa using 174 characters.  Three years later, Upchurch (1998) published a major revision of his own work; in the same year, the other major school of sauropod phylogeny launched with a JVP memoir (Wilson and Sereno 1998), which featured only 10 taxa and 109 characters, but discussed and illustrated them in more detail.  Wilson (2002) followed this up with a much larger analysis of 27 taxa and 234 characters, and Upchurch et al. (2004), in the second editi0n of The Dinosauria, saw his 27×234 and raised him to 41×309.  The good news is that, by this time, the two schools’ phylogenies, having started out rather different, were converging on a consensus topology with only two significant disagreements, which we’ll come to in a minute.

Since then, Jerry Harris (2006) created a union matrix from the character scores in the Wilson (2002) and Upchurch et al. (2004) matrices, and also threw in a few additional characters from other less ambitious phylogenetic analyses.  This analysis came up with a tree that was very similar to Wilson’s, and subsequent work by Wilson and Upchurch (2009) indicates that Upchurch is now also substantially in agreement with this arrangement.

So here it is!

Consensus phylogeny of sauropod, from Harris (2006)

I plucked this from Jerry’s paper, and coloured it in to show two of the more important groups.  Evolution begins at bottom left, so let’s quickly tour the group.

• First of all, note the outgroups. Sauropods’ nearest relatives are the other saurischian dinosaurs, theropods and prosauropods.  (They’re shown the wrong way round here, because in an unrooted tree it makes no difference.  Ignore that.)
• The most basal sauropods include things like Vulcanodon and, it turns out mostly from the work of Adam Yates (e.g. Yates 2007), a whole bunch of things that, if you looked at them you’d probably guess were prosauropods.
• Sauropods as we know them really begin at the boundary of the group Eusauropoda (“true sauropods”), which is roughly speaking everything more derived than Vulcanodon.  (I won’t discuss the naming of nodes and branches in detail in this post, as it would quickly get too long.  Maybe in Tutorial 8.)  This group I have coloured pink in the diagram above.
• Basal eusauropods include quite a few genera, and the order in which the branched off the “main line” leading to the neosauropods is not clear — as the unresolved polytomy above shows.  Cetiosaurus (which for some reason is not shown in this figure) is generally considered quite derived; some of the Chinese sauropods (Mamenchisaurus, Omeisaurus, etc.) may form a group of their own, but that’s not clear.
• Most of the best-known sauropods fall within the great group Neosauropoda (“new sauropods”), which is coloured purple above.  A few genera float around the root of this group, including Haplocanthosaurus and Jobaria, both of which are sometimes considered neosauropods, and sometimes non-neosauropod sauropods (or what I informally call “eosauropods”, or “dawn sauropods”).
• Otherwise the great split within Neosauropoda is between the diplodocoids (on the left) and the macronarians (on the right) — the groups including Diplodocus on one hand, and Saltasaurus on the other.
• The most basal diplodocoids are the rebbachisaurids over on the left.
• Most other diplodocoids fall into the group Flagellicaudata (“whip-tails”), which is itself composed of dicraeosaurids and diplocodids.  (It’s not clear where in that dichotomy, or maybe just outside it, Suuwassea falls.)
• Over in the other half of the Neosauropods, the first macronarians to diverge are the camarasaurids (which currently means, uh, Camarasaurus).
• Most of the other macronarians fall into Titanosauriformes, the group uniting brachiosaurids (yay!) with titanosaurs and their buddies.  Everything closer to titanosaurs falls within Somphospondyli, and that includes Euhelopus — as it turns out.  (Upchurch had found Euhelopus to fall outside Neosauropoda).
• Once you get past Euhelopus, you’re into Titanosauria (though there are various definitions which place the entry point differently).
• And once inside Titanosauria … well, all bets are off at this stage.  There is a rough consensus that things like Malawisaurus and Andesaurus are pretty basal and Saltasaurus is, sort of by definition, derived.  But apart from that, different studies have come up with wildly different phylogenies, with that of Curry Rogers (2005) being particularly left-field.

Without a doubt, Titanosauria is where the action is right now.  As alluded to in the comments of Matt’s Isisaurus post, it’s a big, big group, encompassing many genera and huge morphological range.  It’s also a long-lived group, spanning the whole of the Cretaceous; and it’s where most new genera are being named, as Argentina seems to be packed full of ‘em.

Well, that’s all for now.  Sorry it’s been wordier than usual — probably not much fun to read, but hopefully useful to refer back to in future.

Here’s the famous 8th cervical vertebra of the “Brachiosaurus” brancai lectotype HMN SII, this time in a left-lateral close-up of its left prezygpapophyseal ramus, showing the many pneumatic excavations.  Enjoy!

"Brachiosaurus" brancai lectotype HMN SII, 8th cervical vertebra, left prezygapophyseal ramus in left lateral view.

References

Curry Rogers (2005)

Harris (2006)

McIntosh (1990)

Russell and Zheng (1993)

Upchurch (1993)

Upchurch (1995)

Upchurch (1998)

Upchurch et al. (2004)

Wilson (2002)

Wilson and Sereno (1998)

Wilson and Upchurch (2009)

Yates (2007)

• Curry Rogers, Kristina. 2005. The Evolutionary History of the Titanosauria. pp. 50-103 in: K. Curry Rogers and J. A. Wilson (eds.), The Sauropods: Evolution and Paleobiology. University of California Press, Berkeley.
• Harris, Jerald D. 2006. The significance of Suuwassea emiliae (Dinosauria: Sauropoda) for flagellicaudatan intrarelationships and evolution. Journal of Systematic Palaeontology 4: 185-198.
• McIntosh, John S. 1990. Sauropoda. pp. 345-401 in: D. B. Weishampel, P. Dodson and H. Osmólska (eds.), The Dinosauria, 1st edition. University of California Press, Berkeley and Los Angeles.
• Russell, Dale A., and Zheng, Zhong. 1993. A large mamenchisaurid from the Junggar Basin, Xinjiang, China. Canadian Journal of Earth Science 30(10/11): 2082-2095.
• Upchurch, Paul. 1993. The Anatomy, Phylogeny and Systematics of the Sauropod Dinosaurs. University of Cambridge, unpublished Ph.D. dissertation. 489 pp.
• 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, J. A. and Paul C. Sereno. 1998. Early evolution and Higher-level phylogeny of sauropod dinosaurs. Society of Vertebrate Paleontology, Memoir 5: 1-68.
• 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
• Yates, Adam M. 2007. The first complete skull of the Triassic dinosaur Melanorosaurus Haughton (Sauropodomorpha: Anchisauria). pp. 9-55 in: Paul M. Barrett and David J. Batten (eds.), Special Papers in Palaeontology 77: Evolution and Palaeobiology of Early Sauropodomorph Dinosaurs. The Palaeontological Association, U.K.