We jumped the gun a bit in asking How fat was Camarasaurus? a couple of years ago, or indeed How fat was Brontosaurus? last year. As always, we should have started with extant taxa, to get a sense of how to relate bones to live animals — as we did with neck posture.

So here we go. I give you a herd of Indian elephants, Elephas maximus (from here):

056-Elephant-c-Gehan-de-Silva-Wijeyeratne-Minneriya-2004-07-27-165-Gathering

You will notice, from this conveniently-close-to-anterior view, that their torsos bulge out sideways, much further than the limbs.

Now let’s take a look at the skeleton of the same animal in the Oxford University Museum of Natural History (downloaded from here but for some reason the photo has now gone away):

4795746797_575d1f0ce6_b

The rib-cage is tiny. It doesn’t even extend as far laterally as the position of the limb bones.

(And lest you think this is an oddity, do go and look at any mounted elephant skeleton of your choice, Indian or African. They’re all like this.)

What’s going on here?

Is Oxford’s elephant skeleton mounted incorrectly? More to the point, are all museums mounting their elephants incorrectly? Do elephants’ ribs project much more laterally in life?

Do elephants have a lot of body mass superficial to the rib-cage? If so, what is that mass? It’s hard to imagine they need a huge amount of muscle mass there, and it can’t be guts. Photos like this one, from the RVC’s televised elephant dissection on Inside Nature’s Giants, suggest the ribs are very close to the body surface:

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I’m really not sure how to account for the discrepancy.

Were sauropods similarly much fatter than their mounted skeletons suggest? Either because we’re mounting their skeletons wrongly with the ribs too vertical, or because they had a lot of superficial body mass?

Consider this mounted Camarasaurus skeleton in the Dinosaur Hall at the Arizona Museum of Natural History (photo by N. Neenan Photography, CC-BY-SA):

Camarasaurus_skeleton

Compare the breadth of its ribcage with that of the elephant above, and then think about how much body bulk should be added.

This should encourage palaeoartists involved in the All Yesterdays movement to dramatically bulk up at least some of their sauropod restorations.

It should also make us think twice about our mass estimates.

Stegotetrabelodon making tracks at Mleisa, © Mauricio Antón

Sweet new paper out today by Bibi et al. in Biology Letters, on some awesome elephant tracks from the United Arab Emirates. I’ve known this was coming for a while, because the second author on the study, Brian Kraatz, has his office about 30 feet down the hall from mine. And I just ran into the lead author, Faysal Bibi, at the Museum fur Naturkunde in Berlin when I was there in December. I knew Faysal when he was an undergrad at Berkeley, and now he’s Dr. Bibi and doing a postdoc in Berlin–how time flies. Congratulations to Faysal, Brian, and the rest of the team on a really cool discovery.

The study is nothing to do with sauropods, but it has a lot of weird connections to SV-POW! Most importantly, the paper is open access, which is both awesome and timely. The life restoration is by the wicked talented Mauricio Antón, who is best known for his paleomammals work but who also restored Brontomerus for National Geographic last year. And some comparative data used in the paper was supplied by SV-POW! favorite and sometime sci-fi author John Hutchinson.

Finally, the elephants that made the tracks were probably Stegotetrabelodon, and although they might not have been full-on Tolkien-by-way-of-Jackson Amphicoelias-sized war-beasts, they were still big four-tusked proboscideans, so I’m calling them oliphaunts. Bibi et al. didn’t find any evidence that the trackmakers were ridden by Haradrim, but they didn’t find any evidence that they weren’t, so that’s how I’m going to imagine them.

Probably not the Mleisa trackmakers. Dammit.

For more stuff, including the paper, the full-res version of the image at top, more sweet images, author bios, and so on, see the press page. There are also nice writeups at Not Exactly Rocket Science and Laelaps. Go check it out.

Reference

Bibi, F., Kraatz, B., Craig, N., Beech, M., Schuster, M., and Hill, A. 2012. Early evidence for complex social structure in Proboscidea from a late Miocene trackway site in the United Arab Emirates. Biology Letters. doi: 10.1098/rsbl.2011.1185

This  is the fifth in a series of posts reviewing the Apatosaurus maquette from Sideshow Collectibles. Other posts in the series are:

There are really only a couple of interesting points to discuss for posture: the neck and the feet.

The neck posture is fine. Easy to say, but since I’m one of the “sauropods held their necks erect” guys, it might need some unpacking.

On one hand, animals really do use stereotyped postures, especially for the neck and head (Vidal et al. 1986, Graf et al. 1995, van der Leeuw et al. 2001). The leading hypothesis about why animals do this is that the number of joints and muscle slips involved in the craniocervical system permits an almost limitless array of possible postures, and that having a handful of stereotyped postures cuts down on the amount of neural processing required to keep everything going. That doesn’t mean that animals only use stereotyped postures, just that they do so most of the time, when there’s no need to deviate.

This might work something like the central pattern generators in your nervous system. When you’re walking down the sidewalk thinking about other things or talking with a friend, a lot of the control of your walk cycle is handled by your spinal cord, not your brain. Your brain is providing a direction and a speed, but the individual muscles are being controlled from the spinal cord. Key quote from the Wikipedia article: “As early as 1911, it was recognized, by the experiments of T. Graham Brown, that the basic pattern of stepping can be produced by the spinal cord without the need of descending commands from the cortex.”

But then you see a puddle or some dog doo and have to place your foot just so, and your brain takes over for a bit to coordinate that complex, ad hoc action. After the special circumstance is past, you go back to thinking about whatever and your spinal cord is back in charge of putting one foot in front of the other. This is the biological basis of the proverbial chicken running around with its head cut off: thanks to the spinal cord, the chicken can still run, but without a brain it doesn’t have anywhere to go (I have witnessed this, by the way–one of the numerous benefits to the future biologist of growing up on a farm).

Similarly, if the craniocervical system has a handful of regular postures–alert, feeding, drinking, locomoting, and so on–it lightens the load on the brain, which doesn’t have to figure out how to fire every muscle slip inserting on every cervical vertebra and on the skull to orient the head just so in three-dimensional space. That doesn’t mean that the brain doesn’t occasionally step in and do that, just like it takes over for the spinal cord when you place your feet carefully. But it doesn’t have to do it all the time.

van der Leeuw et al. (2001) took this a step further and showed that birds not only hold their heads and necks in stereotyped postures, they move between stereotyped postures in very predictable ways, and those movement patterns differ among clades (fig. 7 from that paper is above). There is a lot of stuff worth thinking about in that paper, and I highly recommend it, along with Vidal et al. (1986) and Graf et al. (1995), to anyone who is interested in how animals hold their heads and necks, and why.

So, on one hand, its wrong to argue that stereotyped postures are meaningless. But it’s also wrong to infer that animals only use stereotyped postures–a point we were careful to make in Taylor et al. (2009). And it’s especially wrong to infer that paleoartists only show animals doing familiar, usual things–I wrote the last post partly so I could make that point in this one.

For example, I think it would be a mistake to look at Brian Engh’s inflatable Sauroposeidon duo and infer that he accepts a raised alert neck posture for sauropods. He might or might not–the point is that the sauropods in the picture aren’t doing alert, they’re doing “I’m going to make myself maximally impressive so I can save myself the wear and tear of kicking this guy’s arse”. The only way the posture part of that painting can be inaccurate is if you think Sauroposeidon was physically incapable of raising its neck that high, even briefly (the inflatable throat sacs and vibrant colors obviously involve another level of speculation).

Similarly, the Sideshow Apatosaurus has its neck in the near-horizontal pose that is more or less standard for depictions of diplodocids (at least prior to 2009, and not without periodic dissenters). But it doesn’t come with a certificate that says that it is in an alert posture or that it couldn’t raise its neck higher–and even if it did, we would be free to ignore it. Would it have been cool to see a more erect-necked apatosaur? Sure, but that’s not a new idea, either, and there are other restorations out there that do that, and in putting this apatosaur in any one particular pose the artists were forced to exclude an almost limitless array of alternatives, and they had to do something. (Also, more practically, a more erect neck would have meant a larger box and heftier shipping charges.)

So the neck posture is fine. Cool, even, in that the slight ribbing along the neck created by the big cervical ribs (previously discussed here) gives you a sense of how the posture is achieved. Visible anatomy is fun to look at, which I suspect is one of the drivers behind shrink-wrapped dinosaur syndrome–even though it’s usually incorrect, and this maquette doesn’t suffer from it anyway.

Next item: the famous–or perhaps infamous–flipped-back forefoot. I have no idea who first introduced this in skeletal reconstructions and life restorations of sauropods, but it was certainly popularized by Greg Paul. It’s a pretty straightforward idea: elephants do this, why not sauropods?

Turns out there are good reasons to suspect that sauropods couldn’t do this–and also good reasons to think that they could. This already got some air-time in the comments thread on the previous review post, and I’m going to start here by just copying and pasting the relevant bits from that discussion, so you can see four sauropod paleobiologists politely disagreeing about it. I interspersed the images where they’re appropriate, not because there were any in the original thread.

Mike Taylor: the GSP-compliant strong flexion of the wrist always look wrong to me. Yes, I know elephants do this — see Muybridge’s sequence [above] — but as John H. keeps reminding us all, sauropods were not elephants, and one might think that in a clade optimsied for size above all else, wrist flexibility would not be retained without a very good reason.

Adam Yates: Yes I agree with Mike here, the Paulian, elephant-mimicking hyperflexion of the wrist is something that bugs me. Sauropod wrist elements are rather simple flat structures that show no special adaptation to achieve this degree of flexion. [Lourina sauropod right manus below, borrowed from here.]

Heinrich Mallison: Hm, I am not too sure what I think of wrist flexion. Sure it looks odd, but if you think it through the very reasons elephant have it is likely true in sauropods. And given the huge amount of cartilage mossing on the bones AND the missing (thus shape unknown) carpals I can well imagine that sauropods were capable of large excursions in the wrist.

 Mike: What are those reasons?

Heinrich: Mike, long humeri, very straight posture – try getting up from resting with weak flexion at the wrist. Or clearing an obstacle when walking. I can’t say too much, since this afternoon this has become a paper-to-be.

Mike: OK, Heinrich, but the Muybridge photos (and many others, including one on John H.’s homepage) show that elephants habitually flex the wrist in normal locomotion, not just when gwetting up from resting or when avoiding obstacles. Why?

The interesting thing here is that this is evidence of how flawed our (or maybe just my) intuition is: looking at an elephant skeleton, I don’t think I would ever have guessed that it would walk that way. (That said, the sauropod wrist skeleton does look much less flexible than that of the elephant.)

Matt: (why elephants flex their wrists) Possibly for simple energetics. If the limb is not to hit the ground during the swing phase, it has to be shortened relative to the stance limb. So it has to be bent. Bending the limb at the more proximal joints means lifting more weight against gravity. Flexing the wrist more might be a way to flex the elbow less.

(sauropod wrists look less flexible) Right, but from the texture of the ends of the bones we already suspect that sauropods had thicker articular cartilage caps than do mammals. And remember the Dread Olecranon of Kentrosaurus (i.e., Mallison 2010:fig. 3).

Mike: No doubt, but that doesn’t change the fact that elephant wrists have about half a dozen more discrete segments.

Matt: Most of which are very tightly bound together. The major flexion happens between the radius and ulna, on one hand, and the carpal block on the other, just as in humans. Elephants may have more mobile wrists than sauropods did–although that is far from demonstrated–but if so, it’s nothing to do with the number of bony elements. [Loxodonta skeleton below from Wikipedia, discovered here, arrow added by me.]

(Aside: check out the hump-backed profile of the Asian Elephas skeleton shown previously with the sway-backed profile of the African Loxodonta just above–even though the thoracic vertebrae have similar, gentle dorsal arches in both mounts. I remember learning about this from the wonderful How to Draw Animals, by Jack Hamm, when I was about 10. That book has loads of great mammal anatomy, and is happily still in print.)

And that’s as far as the discussion has gotten. The Dread Olecranon of Kentrosaurus is something Heinrich pointed out in the second of his excellent Plateosaurus papers (Mallison 2010: fig. 3).

Heinrich’s thoughts on articular cartilage in dinosaurs are well worth reading, so once again I’m going to quote extensively (Mallison 2010: p. 439):

Cartilaginous tissues are rarely preserved on fossils, so the thickness of cartilage caps in dinosaurs is unclear. Often, it is claimed that even large dinosaurs had only thin layers of articular cartilage, as seen in extant large mammals, because layers proportional to extant birds would have been too thick to be effectively supplied with nutrients from the synovial fluid. This argument is fallacious, because it assumes that a thick cartilage cap on a dinosaur long bone would have the same internal composition as the thin cap on a mammalian long bone. Mammals have a thin layer of hyaline cartilage only, but in birds the structure is more complex, with the hyaline cartilage underlain by thicker fibrous cartilage pervaded by numerous blood vessels (Graf et al. 1993: 114, fig. 2), so that nutrient transport is effected through blood vessels, not diffusion. This tissue can be scaled up to a thickness of several centimeters without problems.

An impressive example for the size of cartilaginous structures in dinosaurs is the olecranon process in the stegosaur Kentrosaurus aethiopicus Hennig, 1915. In the original description a left ulna (MB.R.4800.33, field number St 461) is figured (Hennig 1915: fig. 5) that shows a large proximal process. However, other ulnae of the same species lack this process, and are thus far less distinct from other dinosaurian ulnae (Fig. 3B, C). The process on MB.R.4800.33 and other parts of its surface have a surface texture that can also be found on other bones of the same individual, and may indicate some form of hyperostosis or another condition that leads to ossification of cartilaginous tissues. Fig. 3B–D compares MB.R.4800.33 and two other ulnae of K. aethiopicus from the IFGT skeletal mount. It is immediately obvious that the normally not fossilized cartilaginous process has a significant influence on the ability to hyperextend the elbow, because it forms a stop to extension. Similarly large cartilaginous structures may have been present on a plethora of bones in any number of dinosaur taxa, so that range of motion analyses like the one presented here are at best cautious approximations.

One of the crucial points to take away from all of this is that thick cartilage caps did not only expand or only limit the ranges of motions of different joints. The mistake is to think that soft tissues always do one or the other. The big olecranon in Kentrosaurus probably limited the ROM of the elbow, by banging into the humerus in extension. In contrast, thick articular cartilage at the wrist probably expanded the ROM and may have allowed the strong wrist flexion that some artists have restored for sauropods. I’m not arguing that it must have done so, just that I don’t think we can rule out the possibility that it may have. And so the flipped-back wrist in the Sideshow Apatosaurus does not bother me–but not everyone is convinced. Welcome to science!

I can’t finish without quoting a comment Mike left on Matt Bonnan’s blog a little over a year ago:

Ever since I saw Jensen’s (1987) paper about how mammals are so much better than dinosaurs because their limb-bones articulate properly, I’ve been fuming on and off about this — the notion that the clearly unfinished ends we see are what was operating in life. No.

This is a pretty fair summary of Jensen’s position. Of course, thanks to Heinrich, now we know why dinosaurs had such crap distal limb articulations: they weren’t mammals (part 1part 2part 3).

Finally, interest in articular cartilage is booming right now, as Mike blogged about here. In addition to the Dread Olecranon of Kentrosaurus, see the Dread Elbow Condyle-Thingy of Alligator from Casey Holliday’s 2001 SVP talk, and of course the culmination of that project in Holliday et al. (2010), and, for a more optimistic take on inferring the shapes of articular surfaces from bare bones, read Bonnan et al. (2010).

Next time: texture and color.

References

Photo copyright Derek Bromhall, borrowed from ARKive.

Let’s say you want to paint an elephant. Where will you locate your elephant, and what will it be doing?

If you depict an elephant standing on a glacier at 14,000 feet, your depiction is accurate, because elephants have been caught doing that. Elephant, standing in a dunescape with no water or vegation in sight: accurate, for the same reason. Elephant, swimming in the ocean out of sight of land: accurate. Elephant, scraping salt out of the wall of a cave: accurate. Elephant, rearing to pull down otherwise unreachable vegetation: accurate. Elephant beating the hell out of a monitor lizard for no apparent reason: accurate. Depictions of elephants doing these things might not be familiar–at least to those of us who don’t live around elephants and therefore don’t get to see them doing all the wacky stuff that real animals do–but they are all accurate, in that elephants actually do these things. A lot, apparently, given that all of the above behaviors were documented in the space of just a few decades. Who knows what you might see if you could watch all the elephants, all the time, for a million years or so.

Is there any reason to think that extinct animals were any less versatile?

On the other hand, just because elephants occasionally go for strolls on glaciers or voluntarily rear up on their hind legs to reach higher does not mean that glaciers are their usual habitat or that rearing is a big part of their behavioral repertoire. So these things are accurate, in that they do happen, unfamiliar, in that they are not widely known by most laypeople*, and unusual, in that they are in the long tail of elephant behavior.

* Before you flood the comment section with, “I knew that about elephants!”, consider the implicit possibility that you are not most laypeople. Does your grandmother know that elephants do all this weird stuff?

So we’ve got three potentially orthogonal axes: accuracy, familiarity, usualness. If this was xkcd, at this point I’d draw a Venn diagram. But it’s not and I’m lazy, so I’m just going to pick three possibilities that illustrate an ascending scale of weirdness. First, the most vanilla (by behavioral weirdness, not artistic achievement) wildlife art depicts animals doing things that they actually do (accurate), frequently (usual), that are known to most people (familiar): giraffes eating out of trees, lions with bloody faces crowded around a dead zebra. Second, art that depicts animals doing things that they actually do (accurate), frequently (usual), that are not known to most people (unfamiliar): hummingbirds eating dirt, mud turtles (kinosternids) climbing trees. Third, art that depicts animals doing things that they actually do (accurate), infrequently (unusual), that are not known to most people (unfamiliar): mammals raising the adopted offspring of other species that are their typical predators or prey, grey whales in the Mediterranean Sea.

The question is, what expectations do we have for paleoart or wildlife art in general? Do paleoartists have a responsibility to only depict extinct animals doing things that are accurate, usual, and familiar? Maybe, if an art director for a book or documentary requested a vanilla dinosaur doing vanilla stuff, but outside of that situation?

Tree-climbing Protoceratops by John Conway, inspired by tree-climbing goats, borrowed from Tetrapod Zoology.

As will probably come as no surprise, I skew pretty hard in the other direction. Paleoartists are vastly more important to paleontology than wildlife artists are to zoology, because they have to do everything that artists of extant wildlife do–and one more crucial thing. If, say, a mammalogist needs to be reminded of the complexity and sheer otherness of her study animals, she can usually go out and observe them for a while, and see herbivores eating meat and carnivores eating plants and interspecies sex and all kinds of crazy stuff that real animals do. Paleontologists do not have the same luxury. It is all too easy to slip into the trap of thinking that we know what our animals were like in life. Consider, for example, the difference in temperament between black and white rhinos, or African and Asian elephants, and then consider Morrison sauropods or Two Medicine ceratopsians, and tell me you know anything about the behavioral differences between Apatosaurus and Diplodocus and their ecological ramifications. We need to be periodically shaken out of our comfortable assumptions and creeping anthropomorphizing (sensu Witton–not just attributing human traits to animals, but casting them in standard roles). We need to be confronted with the essential weirdness–and indeed unknowability–of our study animals. And we need paleoartists to do at least some of this shaking and confronting.

I’m not saying that paleoartists have a responsibility to deliver the unfamiliar or unusual in their art, any more than they have a responsibility to only draw vanilla stuff. I don’t think that paleoartists have a responsibility to anything but accuracy, and I mean accuracy in the inclusive, “not directly contradicted by the fossil record” sense* instead of the exclusive, “only what the evidence will support” sense. I’m saying that we–paleontologists, dino enthusiasts, science writers, museum docents, interested citizens–need the unfamiliar and unusual in paleoart as much or more than we need the comfortable and familiar, and we can only ask for it and be grateful when it appears.

* Hat tip to John Conway for this very useful turn of phrase.

Now, on the flip side, just because there is a huge amount that we will never know about extinct animals does not mean that we should give up trying, or that we should play down the reasonable inferences that we can make. Triceratops probably fought each other more than Centrosaurus, for example, or at least inflicted more damage on the squamosals of their conspecifics (evidence, discussion, link to paper). Would a painting showing two Centrosaurus beating the hell out of each other with their horns and doing all kinds of gnarly damage to each others’ heads therefore be inaccurate? Of course not–I am certain that at some point in the multi-million-year history of centrosaurs, two of them did in fact beat the hell out of each other in just that way. But neither would that painting show their usual mode of settling differences, so far as we can tell from our current interpretation of the available fossils (count the caveats there). That’s what the usualness axis is all about–getting comfortable with the  distinction between what animals occasionally do and what they commonly do.

Scavenging Styracosaurus by Mark Witton–go here for the full-size version and Mark’s thoughts on ceratopsian carnivory.

There is a lot that we simply won’t ever know. Which is why I advise caution in assessing accuracy. As long as whatever the animal is doing doesn’t violate the laws of physics, I think it’s hard to rule out that it could have happened, somewhere, at least once. So the interesting discussions will probably center not around accuracy but around usualness. It’s hard to argue that a styracosaur never scavenged a carcass, but do we think that scavenging and even predation were common behaviors for ceratopsians? Given that squirrels are notorious for killing and eating chipmunks, and that deer apparently eat the eggs and nestlings of ground-nesting birds as often as they can get them, the possibility that carnivory was a usual feature of ceratopsian behavior is worthy of serious consideration. At least, we can say that (1) it is consistent with the behavior of many extant herbivores, and (2) it is something that ceratopsians were  well-equipped to carry out. And given those antecedents, it is a difficult hypothesis to falsify. Then again, “difficult to falsify” does not mean “true”–so there is room for interesting discussions.

And that’s really what this post is all about: fostering productive conversations. I have seen and been part of many paleobiology conversations that went nowhere because accuracy, familiarity, and usualness were all scrambled up–often in my own mind. I’m not saying that this particular parsing of the issues is the best possible–indeed, I hope that it inspires someone else to come up with something better. But I also think that it is better than nothing, and that couching things in these terms might help us zero in on our points of genuine disagreement, and thereby make some progress, whether we’re talking about paleobiology, paleoart, or both.

What do you think?

UPDATE: Dave Hone has blogged on this sort of “what if” stuff, at least thrice: here, here, and here. That last post includes more of John Conway’s art from his “All Yesterdays” slideshow at the SVPCA 2011 icebreaker, which was awesome.

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