This is the fourth in a series of posts in which I review the Apatosaurus maquette from Sideshow Collectibles. Other posts in the series are:

A long-running theme here at SV-POW! is that the torsos of most sauropods were not just deep and slab-sided, they were unusually deep and slab-sided, more so than in most other tetrapods (see this and this, and for a more pessimistic take, this). This is something that is easy to get wrong; we are used to seeing round mammalian torsos and a lot of toy sauropods have nearly circular cross-sections. A lot of sculptors of collectible dinos do get the torso cross-section right, though, and the folks who made this Apatosaurus are no exception.

Next item: there’s an upward kink at the base of the tail, as there should be. Gilmore was the first to point this out, in his 1932 paper on the mounting of the Smithsonian Diplodocus (that’s plate 6 from that paper above; the skeleton on the bottom is the more correct one). This came up in the comment thread of the first post in this series, and since I haven’t had any deeper thoughts on the issue in the past week, I’m just going to copy and paste what I wrote then:

The upkink at the base of the tail is unavoidable; the sacrum is shaped like an inverted keystone and there’s no way to get the proximal caudals to do anything but angle upward without disarticulating them…. The reverse keystoning of sauropod sacra is weird. And it’s in every sauropod sacrum I can remember seeing with my own eyes, including Brachiosaurus altithorax. And yet the only authors I can think of off the top of my head who have discussed it seriously are Gilmore (1932), Greg Paul (2010, maybe a magazine article or two I haven’t seen), maybe Jim Jensen (1988), and IIRC Salgado et al. (1997). If there are more, please let me know–this is something I’m very curious about.

The back is gently arched, with the highest point about midway between the shoulder and hip joints. Where the highest point in the back falls depends on a host of factors, including the relative lengths of the forelimb and hindlimb bones, the amount of cartilage on the ends of those bones, the position and angle of the scapula on the ribcage, and the intrinsic curvature, if any, of the articulated series of dorsal vertebrae, which were themselves separated by an unknown amount of cartilage. Opinions are all over the map on most of these issues, particularly scapular orientation. As a scientist, I am agnostic on most of these points; I don’t think that they’re beyond being sorted out, but there’s a lot of work in progress right now and I haven’t seen evidence that would definitely convince me one way or another. So in lieu of saying that Apatosaurus must have had this scapular orientation and that dorsal curvature and so on, I’ll just note that the maquette has been a dominant feature in my office for a few weeks now and nothing about the body profile, shoulder position, or limb length has ever struck me as odd or worthy of comment. It looks like Apatosaurus to me. Moving on…

In the last post I talked about the visible bulges in the neck that allow one to count the cervical vertebrae. The maquette also has low bumps along the back that mark the neural spines of the dorsal vertebrae. This doesn’t strike me as unreasonable. Attachment scars for interspinous ligaments run all the way up to the tips of the neural spines in most sauropods, so the entire height of one neural spine was often webbed to the next by a continuous ligamentous sheet, as Janensch (1929: plate 4) drew for Dicraeosaurus in the illustration above (isp.L). I don’t think those ligaments would have prevented the bony tips of the vertebrae from being visible, necessarily, and the epaxial muscles should have been on either side of the interspinous ligaments and in the triangular spaces between the spine tips and the transverse processes.

What might have smoothed out the dorsal body profile are supraspinous ligaments (ssp.L in the plate above). These are present in crocs (Frey 1988: figs. 14, 16, 17) but apparently absent in most birds; at least, I haven’t seen any myself, and the Nomina Anatomica Avium does not mention any (Baumel et al. 1993: 156-157). So on phylogenetic grounds their presence in sauropods is equivocal. That said, the tips of the neural spines in most sauropods are fairly rugose. Does that mean that they were webbed one to the next by interspinous ligaments only, or that they were embedded in supraspinous ligaments as well? I don’t know the answer, and I don’t know if anyone else does, either. The whole issue of intervertebral ligaments in sauropods has received too little attention to date. In the absence of better data, I’ll just say that although I wouldn’t put any money on the proposition that the spines made externally visible bumps in life, neither does it offend me.

There is one fairly nit-picky point that I am honor-bound to mention. Because the dorsal neural spines make bumps, it is possible to count the dorsals, just like the cervicals last time. And this count doesn’t work out quite as well. Apatosaurus should have 10 dorsal vertebrae, but try as I might I can’t see more than 8 bumps along the back, and that’s generously assuming that c14′s spine is pretty well ahead of its rib. Is this pathologically anal to complain about? Quite possibly. On the other hand, by sculpting in those details the artists were basically begging geeks like me to come along and count vertebrae just because we could.

The tail is pretty cool. It is appropriately massive where it leaves the body, and has a visible bulge for the caudofemoralis muscle, which originated in the tail and inserted on the fourth trochanter of the femur. The caudofemoralis is the major femur retractor in lizards and crocs and in most non-avian dinosaurs, and rather than go on about it I’ll just point you to Heinrich Mallison’s awesome post about dinosaur butts. The tail of the maquette also has an awesome whiplash. I could say a ton more about the hypothesized uses of whiplash tails in diplodocids and other sauropods, but I don’t feel like climbing that hill just now. Suffice it to say that the maquette’s whiplash is pretty sweet, and avoids the “scale is too small so I just stuck in a piece of wire” mode of making whiplashes that I’ve seen in other, smaller diplodocid sculpts.

The tail has a row of little spines running down the dorsal midline, which have been de rigeur for life restorations of diplodocids and many other sauropods (ahem) since they were first reported by Czerkas (1993). AFAIK, such spines have only been found preserved in the tail region of diplodocids. That’s not to say that they weren’t present in the neck or the back of diplodocids, or in other sauropod taxa, just that the only good fossil traces of them to date have been from the tails of diplodocids, and maybe just one or two tails. So the presence of little spines in the tail of the maquette and not the back or the neck is perfectly–one might even say slavishly–consistent with the fossil evidence. I’ll discuss the flamboyancy or lack thereof in the maquette in another post, so I’ll say no more about this design choice for now.

The limbs are mostly good. The muscles under the skin look plausible, with one exception. As noted before in this series, Apatosaurus was a freakishly robust critter, and the limbs look appropriately sturdy and well-muscled, except where the thigh meets the hip. There is a visible bulge for the ilium, and the anterior margin of the thigh should converge with the most forward point on the ilium. That’s what the preacetabular blade of the ilium is for: to anchor thigh muscles (discussed here, and also nicely illustrated here). Unless the animal had some kind of wasting disease, there was no bone sticking out beyond the muscle, and so the anterior-most point of the ilium has to be the start of the anterior margin of the thigh.

On the positive side, there’s a little ridge running down from the anterior arm onto the forearm for the biceps tendon, which is a nice touch. The manus shows the short, solid arc of metacarpals typical for diplodocids, and an inward-curving thumb claw. The hind feet have the big laterally-curving claws on the first three digits that one expects.

In a way that is difficult to describe in words, the feet really look they are bearing a lot of weight, and this impression of solidity helps to ground the whole maquette. It doesn’t look like a sauropod-shaped balloon that just happens to be poling itself along with limbs that barely touch the ground–an impression that I have gotten occasionally from some other sculptures with overly skinny limbs and too-small feet. This critter looks big, heavy, and powerful, and those are exactly the adjectives one wants to come to mind when looking at Apatosaurus. (I do wonder if doing a Diplodocus in the same scale would be more difficult. How do you convey ‘multi-ton animal’ and ‘gracile’ at the same time?)

To sum up, in the trunk, tail, and limbs I find much to like and little to criticize. The only noteworthy problems are the insufficient dorsal count and the mismatch between the ilium and anterior thigh profile. On one hand these are puzzling goofs, given the overall attention to detail and the numerous points at which the sculpt is not just good but surprisingly good. On the other hand, I didn’t notice the dorsal thing until I bothered to count, and I didn’t notice the thigh thing until the other day when I was writing the first draft of this post, so both problems went unnoticed for weeks and are probably below the threshold of perception for the vast majority of people. The accuracy of the sculpt is so high that my approach to its problems has not been, “Where do I begin?” but rather, “What is keeping this thing from being perfect?” And the answer is, not very much.

Headless sauropod. NOW it's perfect.

The base is nice. It’s not just a generic slab of earth, it’s a muddy surface marked with the tracks of other dinosaurs, including a couple of theropods. The base sits nice and flat, and the Apatosaurus sits nice and flat on it, with no rocking at either point of contact. Not only do the feet of the Apatosaurus fit neatly into the sculpted footprints, one of the hindfeet has a little metal rod that slots into a socket in one of the hindfoot prints, to keep the maquette firmly on the base. That means that if you want to display the maquette off the base, you’ll have to either cut off the rod or make sure that your alternative surface will accommodate it.


The skull is…less satisfying. It’s a nice enough rendition of an Apatosaurus skull, and if it had come by itself I would have been very happy with it. The trouble is that the maquette is considerably more detailed, so when the skull sits next to the maquette it suffers by comparison. But what else are you going to do with it? Make a separate shrine to Apatosaurus somewhere else?

The difference in sculpt quality between the maquette and base on one hand and the skull on the other is apparent even on casual inspection. My copies are sitting on a bookcase adjacent to my office door. Sometimes people walking down the hall pop their heads in, and so far the most common comments are that the maquette is “awesome” and that the base is “cool”. People have been genuinely impressed that the base is a realistically detailed chunk of the environment and not just a flat slab. The only people who have commented on the skull have said that it seems “lame” compared to the maquette.

The base is included in the basic package with the maquette, in a limited edition of 500, which as of this writing goes for $289.99 (here). The package with the skull accessory is in an edition of 100, and goes for $299.99 (here). So the skull is only $10 more, and although it is not quite as nice as the maquette, I think it’s a steal at the price. Mine is certainly not going anywhere.

So much for the gross anatomy. You probably noticed that I haven’t said anything about how the maquette is posed or textured or colored. Those will all be topics for next time.

References

  • Baumel, J.J., King, A.S., Breazile, J.E., Evans, H.E., and Vanden Berge, J.C. (eds.) 1993. Handbook of Avian Anatomy: Nomina Anatomica Avium, 2nd ed. Publications of the Nuttall Ornithological Club, No. 23. Cambridge, Massachusetts, 779 pp.
  • Czerkas, S.A. 1993. Discovery of dermal spines reveals a new look for sauropod dinosaurs. Geology 20:1068–1070.
  • Frey, E. 1988. Anatomie des Körperstammes von Alligator mississippiensis Daudin.
  • Gilmore, C. W. 1932. On a newly mounted skeleton of Diplodocus in the United States National Museum. Proceedings of the United States National Museum 81:1-21.
  • Janensch, W. 1929. Die Wirbelsäule der Gattung Dicraeosaurus. Palaeontographica Suppl. 7(1), 3(2), 37-133.
  • Jensen, J.A. 1988. A fourth new sauropod dinosaur from the Upper Jurassic of the Colorado Plateau and sauropod bipedalism. Great Basin Naturalist 48(2):121-145.
  • Paul, G.S. 2010. The Princeton Field Guide to Dinosaurs. Princeton University Press, 320 pp.
  • Salgado, L., R.A. Coria, and J.O. Calvo. 1997. Evolution of titanosaurid sauropods. I: Phylogenetic analysis based on the postcranial evidence. Ameghiniana 34:3-32.

Back when Darren and I did the Xenoposeidon description, we were young and foolish, and only illustrated the holotype vertebra NHM R2095 in four aspects: left and right lateral, anterior and posterior.  No dorsal or ventral views.

Also, because the figure was intended for Palaeontology, which prints only in greyscale, I stupidly prepared the figure in greyscale, rather than preparing it in colour and then flattening it down at the last moment.  (Happily I’d learned that lesson by the time we did our neck-posture paper: although it was destined for Acta Palaeontologia Polonica, which also prints in greyscale, and though the PDF uses greyscale figures, the online full-resolution figures are in colour.)

As if that wasn’t dumb enough, I also composited the four featured views such that the two lateral views were adjacent, and above the anterior and posterior views — so it wasn’t easy to match up features on the sides and front/back between the views.  Since then, I have landed on a better way of presenting multi-view figures, as in my much-admire’d turkey cervical and pig skull images.

So, putting it all together, here is how we should have illustrated illustrated Xenoposeidon back in 2007 (click through for high resolution):

(Top row: dorsal view, with anterior facing left; middle row, from left to right: anterior, left lateral, posterior, right lateral; bottom row, ventral view, with anterior facing left.  As always with images of NHM-owned material, this is copyright the NHM.)

Of course, if we’d published in PLoS ONE, then this high-resolution (4775 x 4095), full colour image could have been the published one rather than an afterthought on a blog somewhere.  But we didn’t: back then, we weren’t so aware of the opportunities available to us now that we live in the Shiny Digital Future.

In other news, the boys and I all registered Xbox Live accounts a few days ago.  I chose the name “Xenoposeidon”, only to find to my amazement that someone else had already registered it.  But “Brontomerus” was free, so I used that instead.

A few months ago, prosauropod supremo Adam Yates blogged about the Aardonyx cake that the BPI honours class baked in his honour.  In the comments, I mentioned that my wife Fiona once made me a BMNH R5937:D9 cake (i.e. a cake in the form of the more posterior of the pair of nicely preserved dorsal vertebrae of The Archbishop, in right lateral view). At the time, I couldn’t find the photo that I knew had been taken, and Adam asked me to post it when it turned up.

Voila!

And here, once more, is the real thing for comparison:

(Note that the topology of the lateral lamination is spot on, with a single infradiapophyseal lamina which forks into anterior and posterior branches only some way ventral to the diapophysis.  That’s what you look for in a cake.)

Update (21 April)

Silly me, of course what I should have shown is the cake and the vertebra side by side.  Here they are — together at last!

As we all know, the International Code of Zoological Nomenclature is a large and intimidating document.  As a result, zoologists naming new animals often do not read it in its entirety (I know I haven’t).  It’s probably because of this that many of the more avoidable nomenclatural mistakes occur.

Whatever might or might not eventually be possible in terms of simplifying the Code, everyone recognises that that would be a huge job, and something that would take years to do.  So let’s ignore that possibility for now.

In the short term, what would be much more useful would be if someone could work up a very short document — no more than a single page of A4 and hopefully much shorter — that states in simple bullet-points what MUST be done to ensure that a new name is valid.  Then there would be no excuse for zoologists venturing into nomenclature for the first time not to read such a document — let’s call it the ICZN Cheat Sheet.

Neural spine morphology along the vertebral series of Bonitasaurua salgadoi (MPCA-460) in anterior view. From Gallina 2011, doi:10.1590/S0001-37652011005000001

Because it’s easier to steer a moving ship, I wrote to the ICZN email list this morning proposing an initial set of bullet points.  I did not for a moment expect that they were complete, consistent or even necessarily correct; but I hoped that they could at least serve as a starting point for a very quick process of putting such a list together.

I am pleased to say that response on the list was fairly positive, and at the suggestion of one of the list members I have now posted the in-progress checklist as a page on this site, having revised it in accordance with several suggestions.

If you’re interested in contributing to this effort — helping us to derive a clear, concise and correct one-page guide to naming new zoological genera and species — please head over to the page and comment there.  (Comments on this post are closed, to avoid splitting discussion across two places.)

I recently stumbled across this rather good photograph of the holotype vertebra of our old buddy “Ultrasauros“, thanks to Wikipedia contributor Ninjatacoshell, and thought you’d like to see it:

This is a rather legendary vertebra, but until recently there were no good photographs of it on the web (I know because I tried to find one for my talk at the Dinosaurs: A Historical Perspective conference in 2008).

See It’s Ultrasaurus… I mean, um, Ultrasauros… err, Supersaurus! for the now-traditional run-down of the taxonomic mess surrounding this specimen.

In other news, everyone in palaeontology should read Heinrich Mallison’s recent article No 4WD For Plateosaurus over on the Palaeontologia Electronica blog.  He highlights a lot of important issues that have general applicability.

SV-POW! is, as I’m sure you know, devoted to sauropod vertebrae. But occasionally we look at other stuff… and you might have noticed that, in recent months, we’ve been looking at, well, an awful lot of other stuff. I’m going to continue that theme here and talk about salamanders. Yeah: not sauropods, not sauropodomorphs, not saurischians, and not even dinosaurs or archosaurs. But salamanders. Don’t worry, all will become clear. This all started back in May 2010 when I blogged about amphiumas over at Tet Zoo. Amphiumas are very unusual, long-bodied aquatic salamanders.

A Three-toed amphiuma _Amphiuma tridactylum_. Photo by Brad Moon.

As it happens, amphiuma vertebrae are particularly interesting if you work on saurischians because (drum-roll)… they have laminae. The term lamina is not restricted to structures present only in pneumatic saurischians: I would argue that it should be used for any sheet-like bony process on a vertebra, and I hope everyone agrees with me. Laminae are not common outside of Saurischia, but are present here and there: they’re present in stem-archosaurs (like Erythrosuchus), various crurotarsan archosaurs (including aetosaurs), some neosuchian crocodilians, and silesaurids (Desojo et al. 2002, Parker 2003, Nesbitt 2005, Wedel 2007, Butler et al. 2009). Even weirder, they’re present in Aneides lugubris, the Arboreal salamander of California and Baja California (Wedel 2007). But that’s about it.

Why would a salamander ‘want’ vertebral laminae? The laminae of the Arboreal salamander are presumed to be related to the extensive accessory ossification present in the skeleton of this animal, itself a consequence of adaptation to a peculiar climbing lifestyle. In other words, it’s hypothesised that the function (if I may be so bold as to use that word…) of the salamander’s laminae is nothing like that of the archosaurs that have them.

And now we know that A. lugubris isn’t the only salamander with laminae: amphiumas have them too. They’re clearly figured in the amphiuma literature (Gardner 2003), but (so far as I know) no-one has previously drawn attention to them when discussing archosaur laminae.

A mid-dorsal vertebra of _Amphiuma_, from Gardner (2003).

Gardner (2003) figured schematic amphiuma dorsal vertebrae that were based on a combination of features present in two of the three living amphiuma species (namely, Amphiuma means and A. tridactylum). On the lateral sides of the centra are structures that – if seen in an archosaur – would almost certainly be identified as anterior and posterior centrodiapophyseal laminae (using, as always, the nomenclature proposed by Wilson (1999)) [see the digram above, from Gardner (2003)]. There are also structures on the dorsal surfaces of the postzygapophyses that look something like laminae: they extend from the posterolateral parts of the neural arch and run across the tops of the postzygapophyses, hence recalling spinopostzygapophyseal laminae. Actually, I’ve just realised that similar structures are also sometimes present in anurans (frogs and toads) where they’ve been called paraneural crests or paraneural processes. These structures do have a ‘known’ function: in amphiumas they’re associated with complex dorsalis trunci epaxial muscles. Unlike the spinopostzygapophyseal laminae of saurischians, the structures in the amphibians are low ridges that don’t contact the neural spines, so it could be argued that they aren’t so lamina-like after all.

But what about the structures on the sides of the centra? Why are laminae present in a group of long-bodied aquatic salamanders? Why are laminae present at all? This question has been asked a few times here on SV-POW! (here, for example), and there are two primary hypotheses. One is that the laminae keep the various air sacs separate from each other, perhaps because they persist while much of the bone around them is resorbed during ontogeny, while the other is that they somehow provide mechanical support and are aligned along lines of stress (for more on this subject see the piece on finite element analysis).

An assortment of _Amphiuma_ cervical, dorsal and caudal vertebrae, from Gardner (2003). The 'paraneural crests' (the lamina-like structures on the postzygapophyses) are visible in G and L, and the lateral central laminae are visible in some of the other vertebrae figured here.

The pneumaticity explanation can’t work for amphiumas (given that they’re apneumatic): does the ‘mechanical support’ one apply instead? We don’t know anything about stress distribution in amphiuma vertebrae – in fact, I don’t think we know anything about the mechanics of amphiumas at all – but it’s possible that the laminae might play this role, especially given that amphiumas have to bend, twist and push with their bodies while excavating burrows.

In conclusion, we just don’t really know what’s going on here. In fact, all we can really do at the moment is wave our arms around a bit and say “Hey, amphiumas have vertebral laminae, too”, and that’s pretty much all I’m doing here. It’s also possible that the structures I’m talking about in amphiumas are very different in detail from the vertebral laminae present in archosaurs: I’ve never even seen a single amphiuma skeletal element and am basing all of this on photos and diagrams in the literature. Nevertheless, it’s something definitely worth bringing attention to. As usual, we stand poised at the abyss, straining our eyes to see into the infinite darkness ahead.

References

Butler, R. J., Barrett, P. M. & Gower, D. J. 2009. Postcranial skeletal pneumaticity and air-sacs in the earliest pterosaurs. Biology Letters 5, 557-60.

Desojo, J. B., Arcucci, A. B. & Marsicano, C. A. 2002. Reassessment of Cuyosuchus huenei, a Middle–Late Triassic archosauriform from the Cuyo Basin, west-central Argentina. Bulletin of the New Mexico Museum of Natural History and Science 21, 143–148.

Gardner, J. D. 2003. The fossil salamander Proamphiuma cretacea Estes (Caudata; Amphiumidae) and relationships within the Amphiumidae. Journal of Vertebrate Paleontology 23, 769-782.

Nesbitt, S. J. 2005. Osteology of the Middle Triassic pseudosuchian archosaur Arizonasaurus babbitti. Historical Biology 17, 19–47.

Parker, W. G. 2003. Description of a new specimen of Desmatosuchus haplocerus from the Late Triassic of northern Arizona. Unpublished MS thesis, Northern Arizona University, Flagstaff, AZ, 312 pp.

Wedel, M. J. 2007. What pneumaticity tells us about ‘prosauropods’, and vice versa. Special Papers in Palaeontology 77, 207-222.

Wilson, J. A. 1999. A nomenclature for vertebral laminae in sauropods and other saurischian dinosaurs. Journal of Vertebrate Paleontology 19, 639-653.

Here at SV-POW! Towers, we have often lamented that so much dinosaur research is locked up behind the paywalls of big for-profit commercial publishers, and that even work that’s been funded by public money is often not available to the public.

One of the quiet delights of the last couple of years has been watching the hide-research-from-researchers edifice slowly crumbling, and indeed we have a whole section of the site dedicated to that very thing: the Shiny Digital Future.  The process is slow, which should surprise nobody given that large, powerful, profit-motivated corporations are trying to prevent it, but it does feel increasingly inevitable.

This week has brought two more steps towards the open-access utopia: one of them specific and immediate, the other more long term but potentially much more far-reaching.

  • In the immediate, the New Mexico Museum of Natural History has made all issues of its Bulletin up to 2008 freely available.  Although the quality of the articles in these issues is hugely variable, there is a lot of good and important stuff in there, and it’s a boon to the community that they are now open to anyone who wants to read them.
  • I just heard today about the Federal Research Public Access Act (HR 5037), brought before Congress six days ago by a bipartisan group of six representatives (four Democrats and two Republicans).  If passed it would ensure that all research funded by eleven U.S. federal agencies was made open-access.  If you’re American, follow the link to see what you do to help ensure that it’s passed!

As Galadriel said, the world is changing.

Holotype dorsal vertebra of Nopcsaspondylus, apparently, from Mannion (2010:fig. 5), although it bears little resemblance to Apesteguia (2007:fig. 2) which is the illustration of that element in the paper that named the genus. Mannion's figure seems to be reproduced from Nopcsa (1902), which I really ought to get hold of so I can check this for myself. Ever notice how Haplocanthosaurusy rebbachisaurid dorsals look? Just sayin', is all.

Finally: I know that whenever we talk about proprietary publishers, I always tell people to go and read Scott Aaronson’s essay on the subject, but seriously: if you’ve not read it before, go and read it now.  It’s brilliant.

Update (22 April 2010)

Thanks to Phil for the clarification below on whether the pictured vertebra is or is not the Nopcsaspondylus holotype (it is).  Phil also sent me a scan of Nopcsa’s original figure of this plate, which is rather better than the reduced version that made it into the new paper, so here it is!

Definitely the holotype dorsal vertebra of Nopcsasopondylus, from Nopcsa (1902). Still looking kind of Haplocanthosaurusy. The main differences apparent here seem to be that this vertebra has a more vertically compressed centrum and more medially directed SPOLs than the Haplo dorsals. Still, I find myself wondering how many steps it would take to move Haplo into a basal rebbachisaurid position. Funny, really, Rebbachisauridae is just about the only sauropod clade that it hasn't been referred to.

Dorsal vertebrae from Argentinosaurus (center) and Supersaurus (either side). The vert on the left is the holotype of Ultrasauros, and the one on the right is the holotype of Dystylosaurus, but both of those taxa have been sunk into Supersaurus. Found on teh intert00bz.

As often happens here, a comment thread got to be more interesting than the original post and ended up deserving a post of its own. In this case, I’m talking about the thread following the recent Mamenchisaurus tail club post, which got into some interesting territory regarding mass estimates for the largest sauropods. This post was inspired by a couple of comments in particular.

Zach Armstrong wrote:

I don’t trust Mazzetta et al.’s (2004) estimate, because it is based off of logarithmic-based regression analyses of certain bone lengths, which a recent paper by Packard et al. (2009) have shown to overestimate the mass by as much as 100 percent! This would mean the estimate of 73 tonnes given my Mazzetta would be reduced to 36 tonnes.

To which Mike replied:

Zach, Mazzetta et al. used a variety of different techniques in arriving at their Argentinosaurus mass estimate, cross-checked them against each other and tested their lines for quality of fit. I am not saying their work is perfect (whose is?) but I would certainly not write it off as readily as you seem to have.

Weeeeell…Mazzetta et al. did use a variety of measurements to make their mass estimates, but they did it in a way that hardly puts them above criticism. First, their estimates are based on limb-bone allometry, which is known to have fairly low accuracy and precision (like, often off by a factor of 2, as Zach noted in his comment). Second, the “raw data” for their allometric equation consists of volumetric mass estimates. So their primary estimation method was calibrated against…more estimates. Maybe I’m just lazy, but I would have skipped the second step and just used volumetric methods throughout. Still, I can see the logic in it for critters like Argentinosaurus where we have limb bones but no real idea of the overall form or proportions of the entire animal.

Anyway, the accuracy of their allometric estimates is intertwingled with their volumetric results, so if their volumetric estimates are off…. The volumetric estimates used a specific gravity of 0.95, which to me is unrealistically high. Taking into account the skeletal pneumaticity alone would lower that to 0.85 or 0.8, and if the critter had air sacs comparable to those of birds, 0.75 or even 0.7 is not beyond the bounds of possibility (as discussed here and also covered by Zach in his comment).

Now, Mazzetta et al. (2004) were not ignorant of the potential effects of pneumaticity. Here’s  what they wrote about density (p. 5):

The values from Christiansen (1997) were recalculated using a slightly higher overall density (950 kg/m^3), as the 900 kg/m^3 used in that paper may be slightly too low. Most neosauropods have extensively pneumatised vertebrae, particularly the cervicals, which would tend to lower overall density. However, these animals are also very large, implying a proportionally greater amount of skeletal tissue (Christiansen, 2002), particularly appendicular skeletal tissue, and consequently, they should have had a higher overall density.

This is pretty interesting: they are arguing that the positive allometry of skeletal mass as a fraction of body mass–which is well documented in extant critters–would offset the mass reduction from pneumaticity in animals as big as sauropods. I haven’t given that enough thought, and I definitely need to. My guess–and it is a guess–is that the effects of skeletal allometry were not enough to undo the lightening imposed by both PSP (~10%) and pulmonary air sacs (another ~10%, separate from the lungs), but I haven’t done any math on this yet. Fodder for another post, I reckon.

Getting back to Mazzetta et al., some of the volumes themselves strike me as too high, like ~41,500 liters for HM SII. That’s a LOT more voluminous than Greg Paul, Don Henderson, or Mike found for the same critter. The 16 metric ton Diplodocus and 20.6 metric ton Apatosaurus used by Mazzetta et al. are also outside the bounds of other recent and careful estimates. Not necessarily wrong, but definitely at the upper end of the current spectrum.

Mazzetta et al. got a mass estimate of 73,000 kg for Argentinosaurus, but (1) they used a density that I think is probably too high even if skeletal allometry is considered, (2) at least some of the volumetric mass estimates that form the “data” for the limb-bone regressions are probably too high, and (3) even if those problems were dealt with, there is still the general untrustworthiness of limb-bone regression as a mass estimation technique. 1 and 2, if fixed to my satisfaction, would tend to push the estimated mass of Argentinosaurus down, perhaps significantly (the effect of 3 is, if not unknowable, at least unknown to me). Given that, Zach’s ~52 metric ton estimate for Argentinosaurus is very defensible. (Probably worth remembering that I am a sparse-wing fanatic, though.)

None of this means that Mazzetta et al. (2004) were sloppy or that their estimate is wrong. Indeed, one of the reasons that we can have such a deep discussion of these points is that every link in their chain is so well documented. And there is room for honest disagreement in areas where the fossils don’t constrain things as much as we’d like. You cannot simply take a skeleton, even a complete one, and get a single whole-body volume. The body masses of wild animals often fluctuate by a third over the course of a single year, which pretty well buries any hope of getting precise estimates based on skeletons alone. And no one knows how dense–or sparse–sauropods were. I haven’t actually done any math to gauge the competing effects of skeletal allometry on one hand and PSP and air sacs on the other–and, AFAIK, no one else has either (Mazzetta et al. were guessing about pneumaticity as much as I’m guessing about skeletal allometry). Finally, Argentinosaurus is known from a handful of vertebrae and a handful of limb bones and that’s all, at least for now. If we can’t get a single body volume even when we have a complete skeleton, we have to get real about how precise we can be in cases where we have far less material.

The upshot is not that Argentinosaurus massed 73 metric tons or 52 or any other specific number. As usual, the two-part take home message is that (1) mass estimates of sauropods are inherently imprecise, so all we can do is make our assumptions as clear as possible, and (2) even the biggest sauropods might have been smaller than you think. ;-)

Reference

Mazzetta, G.V., Christiansen, P., and Farina, R.A. 2004. Giants and bizarres: body size of some southern South American Cretaceous dinosaurs. Historical Biology 2004:1-13.

Shunosaurus lii is a basal eusauropod from the Middle Jurassic of China.  Outside of palaeontological circles, it’s not at all well known — which is kind of surprising, as it’s one of the best represented of all sauropods.  It’s known from numerous complete skeletons, including skulls, and has been described in detail in Zhang’s (1988) monograph: 89 pages and 15 plates.  Here’s a skeleton of one individual, as found in the ground:

Shunosaurus lii, referred adult individual ZDM T5402, skeleton as found. From Zhang (1988:fig. 2)

Apart from being so well represented, Shunosaurus is known mostly for its tail club, which at the time of its discovery was unique among sauropods.  Despite recent discoveries of Mamenchisaurus hochuanensis individuals with preserved tail-clubs, and of Spinophorosaurus, the Shunosaurus tail-club is the best developed and best preserved.

But I don’t want to show you that.  I want to show you something I’ve been wanting to see for many years, and today finally saw for the first time: a feature of the dorsal vertebrae totally unique to Shunosaurus, known as postparapophyses.

Shunosaurus lii, referred juvenile individual ZDM T5401, dorsal vertebrae 3, 7 and 11 in left lateral view. Note postparapophyses on dorsals 7 and 11. From Zhang (1988:figs. 31-32)

Sadly, these are the only figures in the paper that show the postparapophyses (and as far as I know the only published figures anywhere).  So we have them in lateral view only, and lack what would be an informative posterior view.  Plate 10, part 1, supposedly shows one of the posterior dorsals in posterior view, but in my PDF at least the reproduction is so poor as to be wholly uninformative.

What makes things even worse is that the extended English-language abstract on pages 86-91 of Zhang (1988) does not mention this feature at all — in fact it occurs only in the list of anatomical abbreviations on page 6.  So, to the best of my knowledge, here is the entirety of what has been published in the English language about this feature based on observation of the material:

ppp, postparapophysis

Wilson and Sereno (1998:14-15) expanded a little on this, but it’s not clear that what they wrote was based on anything more than the figure above.  Here it is anyway, for completeness:

Comments–Zhang’s (1988:78-79) diagnosis listed numerous features, only a few of which appear to be autapomorphies of Shunosaurus lii.  One of the more striking autapomorphies is an unusual articulation between the ribs and the posterior dorsal vertebrae.  The parapophyseal articulation is split between adjacent vertebrae, with a portion of the articulation in its usual position by the prezygapophyses and an anterior extension located near the postzygapophysis on the preceding vertebra (Zhang, 1988:figs. 31, 32; “postparapophysis”).

That’s your lot.

So if we’re to make anything at all of the PPPs, it will have to be on the basis of the figure reproduced above.  And I don’t really know how much we can say.  The PPPs look sort of like postzygapophyses, havng a distinct ventrally oriented facet.  This makes me wonder whether they are in fact lateral extensions of the postzygapohyseal facets, perhaps connected by a lamina that would be visible in posterior view.

The bottom line is, I don’t know, and I would greatly appreciate comments (or better still, photos!) from anyone who has seen the material first-hand.

I leave you with Zheng’s (1988:fig. 57) skeletal reconstruction of this distinctively dumpy-looking sauropod.  Note by the way that the plantigrade manus reconstruction is almost certainly wrong: the metacarpals should be held in a more or less vertical arcade as in other sauropods.

Shunosaurus lii, referred adult individual ZDM T5402, skeletal reconstruction in left lateral view. From Zheng (1988:fig. 7). Man, that thing is ugly.

References

  • Wilson, Jeffrey A. and Paul C. Sereno.  1998.  Early evolution and Higher-level phylogeny of sauropod dinosaurs.  Society of Vertebrate Paleontology, Memoir 5: 1-68.
  • Zhang Yihong.  1988.  The Middle Jurassic dinosaur fauna from Dashanpu, Zigong, Sichuam, vol. 1: sauropod dinosaur (I): Shunosaurus. Sichuan Publishing House of Science and Technology, Chengdu, China.

Update (9 March 2010)

Rob Taylor found this nice photograph of what is apparently a skeletal mount of Shunosaurus: the original is here.  Any information about this mount will be gratefully received: please comment below if you know anything.

Lovers of fine sauropods will be well aware that, along with the inadequately described Indian titanosaur Bruhathkayosarus, the other of the truly super-giant sauropods is Amphicoelias fragillimus.  Known only from a single neural arch of a dorsal vertebra, which was figured and briefly described by Cope (1878) and almost immediately either lost or destroyed, it’s the classic “one that got away”, the animal that sauropod aficionados cry into their beer about late at night.

Amphicoelias fragillimus, holotype dorsal vertebral neural arch in posterior view. From Osborn and Mook (1921:fig. 21), which in turn was gently tweaked from Cope (1878:unnumbered and only figure).

I’m not going to write about A. fragillimus in detail here, because Darren’s so recently covered it in detail over at Tetrapod Zoology — read Part 1 and Part 2 right now if you’ve not already done so.  The bottom line is that it was a diplodocoid roughly twice as big as Diplodocus in linear dimension (so about eight times as heavy).  That makes it very very roughly 50 m long and 100 tonnes in mass.

But Mike!, you say, Isn’t it terribly naive to go calculating masses and all from a single figure of part of a single bone?

Why, yes!  Yes, it is!  And that is what this post is about.

As I write, the go-to paper on A. fragillimus is Ken Carpenter’s (2006) re-evaluation, which carefully and tentatively estimated a length of 58 m, and a mass of around 122,400 kg.

As it happens, Matt and a colleague submitted a conference abstract a few days ago, and he ran it past me for comments before finalising.  In passing, he’d written “there is no evidence for sauropods larger than 150 metric tons and it is possible that the largest sauropods did not exceed 100 tons”.  I replied:

I think that is VERY unlikely. [...] the evidence for Amphicoelias fragillimus looks very convincing, Carpenter’s (2006) mass estimate of 122.4 tonnes is conservative, being extrapolated from Greg Paul’s ultra-light 11.5 tonne Diplodocus.

Carpenter’s estimate is based on a reconstruction of the illustrated vertebra, which when complete he calculated would have been 2.7 m tall.  That is 2.2 times the height of the corresponding vertebra in Diplodocus, and the whole animal was considered as it might be if it were like Diplo scaled up by that factor.  Here is his reconstruction of the vertebra, based on Cope’s figure of the smaller but better represented species Amphicoelias altus:

One possible reconstruction of the Alphicoelias fragillimus vertebra, from Carpenter (2006:fig. 1).  Part A is Cope’s original figure annotated with lamina designations; part C is Cope’s illustration of an Amphocoelias altus dorsal; part B is Carpenter’s reconstruction of the former after the latter.

Matt’s answer to me was:

First, Paul’s ultra-light 11.5 tonne Dippy is not far off from my 12 tonne version that you frequently cite, and mine should be lighter because it doesn’t include large air sacs (density of 0.8 instead of a more likely 0.7). If my Dippy had an SG of 0.7, it would have massed only 10.25 tonnes. Second, Carpenter skewed [...] in the direction of large size for Amphicoelias. I don’t necessarily think he’s wrong, but his favoured estimate is at the extreme of what the data will support. Let’s say that Amphicoelias was evenly twice as large as Dippy in linear terms; that could still give it a mass as low as 90 tonnes. And that’s not including the near-certainty that Amphicoelias had a much higher ASP than Diplodocus. If Amphicoelias was to Diplodocus as Sauroposeidon was to Brachiosaurus—pneumatic bones about half as dense—then 1/10 of its volume weighed ½ as much as it would if it were vanilla scaled up Dippy, and we might be able to knock off another 5 tonnes.

There’s lots of good stuff here, and there was more back and forth following, which I won’t trouble you with.  But what I came away with was the idea that maybe the scale factor was wrong.  And the thing to do, I thought, was to make my own sealed-room reconstruction and see how it compared.

So I extracted the A.f. figure from Osborn and Mook, and deleted their dotted reconstruction lines.  Then I went and did something else for a while, so that any memory of where those lines might have been had a chance to fade.  I was careful not look at Carpenter’s reconstruction, so I could be confident mine would be indepedent.  Then I photoshopped the cleaned A. fragillimus figure into a copy the A. altus figure, scaled it to fit the best as I saw it, and measured the results.  Here it is:

My scaling of a complete Amphicoelias fragillimus vertebra: on the left, Cope’s figure of the only known vertebra; on the right, Cope’s figure of an A. altus dorsal vertebra, scaled to match the preserved parts of the former.  Height of the latter scaled according to the measured height of the former.

As you can see, when I measured my scaled-to-the-size-of-A.f. Amphicoelias vertebra, it was “only” 2293 mm tall, compared with 2700 mm in Ken’s reconstruction.  In other words, mine is only 85% as tall, which translates to 0.85^3 = 61% as massive.  So if this reconstruction is right, the big boy is “only” 1.87 times as long as Diplodocus in linear dimension — maybe 49 meters long — and would likely come in well below the 100-tonne threshhold.  Using Matt’s (2005) 12-tonne estimate for Diplodocus, we’d get a mere 78.5 tonnes for Amphicoelias fragillimus.  So maybe Matt called that right.

Amphicoelias altus dorsal vertebra, almost certainly the holotype, in left lateral view, lying on its back.  Photograph by Matt Wedel, from the collections of the AMNH.  I can’t believe — can’t BELIEVE — that I didn’t take ten minutes to look at this vertebra when I was in that basement last February.  What a doofus.

The Punchline

Folks — please remember, the punchline is not “Amphicoelias fragillimus only weighed 78.5 tonnes rather than 122.4 tonnes”.  The punchline is “when you extrapolate the mass of an extinct animal of uncertain affinities from a 132-year-old figure of a partial bone which has not been seen in more than a century, you need to recognise that the error-bars are massive and anything resembling certainty is way misplaced.”

Caveat estimator!

References

  • Carpenter, Kenneth.  2006.  Biggest of the big: A critical re-evalustion of the mega-sauropod Amphicoelias fragillimus Cope, 1878.  pp. 131-137 in J. Foster and S. G. Lucas (eds.), Paleontology and Geology of the Upper Jurassic Morrison Formation.  New Mexico Museum of Natural History and Science Bulletin 36.
  • Cope, Edward Drinker.  1878.  Geology and Palaeontology: a new species of Amphicoelias.  The American Naturalist 12 (8): 563-566.
  • Osborn, Henry Fairfield, and Charles 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.
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