Snoozing brontosaur by Bakker

From The Dinosaur Heresies.

Part 1.

Yesterday I was in Oxford for the Rigour and Openness in 21st Century Science conference (web-site here, tweets here though they also include newer ones from Day 2 which is happening as I write this).

There was a lot to enjoy about the day, including meeting Cameron Neylon of PLOS and Jason Hoyt of PeerJ, both for the first time. The highlight for me, unsurprisingly, was the debate at the Oxford Union in the evening, of which more in following posts.

Another highlight was meeting my anti-particle — the pro-Elsevier Mike Taylor. There are quite a few odd coincidences linking him and me, and he has been using us both as a motivating example of the need for ORCID: skip to 5:50 in this video for an example.

?????????????????????

 

There was some speculation that if we ever met, we’d both be annihilated in a burst of pure energy, but happily there were no fireworks.

Apart from a brief fist-fight.

WP_000940

 

Other Mike Taylor (hereafter OMT) had some interesting things to say about Elsevier, but I won’t pass them on without his permission. Maybe he’ll drop by here and comment.

By the way, I think this was the second time I have worn a tie in the last decade or so.

 

I was cruising the monographs the other night, looking for new ideas, when the humerus of Opisthocoelicaudia stopped me dead in my tracks. I think you’ll agree it is an arresting sight:

Opisthocoelicaudia right humerus in lateral, anterior, medial, and posterior views, from Borsuk-Bialynicka (1977: figure 7)

Opisthocoelicaudia right humerus in medial, anterior, lateral, and posterior views, from Borsuk-Bialynicka (1977: figure 7)

I’d seen it before, but somehow I had never grokked its grotesque fatness. I mean, damn, Opisthocoelicaudia, you really let yourself go. Especially compared to the slenderness and grace of this juvenile Alamosaurus humerus:

Alamosaurus left humerus in anterior and posterior views, from Lehman and Coulson (2002: figure 7).

Alamosaurus left humerus in anterior and posterior views, from Lehman and Coulson (2002: figure 7).

Now, I realize that part of the slenderness of this Alamosaurus humerus might be because it’s a juvenile–other alamosaur humeri are a bit more robust–but it’s still a striking contrast. I couldn’t help but superimpose them, scaled to the same midshaft width:

Alamosaurus and Opisthocoelicaudia humeri superimposed

I flipped the Alamosaurus humerus left-to-right to match that astonishing lump of Opisthocoelicaudia. The result reminds me of one of Abrell and Thompson’s Actual Facts:

If you put Woodrow Wilson inside William Howard Taft, he would have stuck out by a good 18 inches.

None of that probably signifies anything more than that I am easily amused. And also,  Opisthocoelicaudia is Just Plain Wrong. You hear me, Opisthocoelicaudia? Don’t make me make you cry mayonnaise!

References

Brontomerus cartoon - John Trotter - paintmonkeystudios-dot-com

One of our army of field correspondents, Seth Segal, sent us a scan of this cartoon from the spring 2011 issue (#97) of Prehistoric Times (yes, we’re a bit late to the party on this one). Shifty little weasels that we are, we were entertained by it, so we tracked down John Trotter at Paintmonkey Studios. He kindly sent the nice version you see above, and gave us permission to post.

I really like the idea of undescribed dinosaurs just going about their business, and then being surprised by having new names sprung on them. I can well imagine some of them being disappointed, too.

Argentina…saurus. Lizard. From. Argentina. Seriously? You know, there’s a million dinosaurs from Argentina. Why do I get stuck with the generic name that is actually generic? Nothing about how big I am? Really? I mean, I weigh, like, two Supersauruses. What’s the Latin for double-Supersaurus-rex? And here I am with Antarctosaurus–that poser’s got a whole continent in his name, and he’s not even from there! And what about that so-called “earthquake lizard”? I heard they found him wandering around all delusionsal, claiming to be 150 feet long and the biggest thing ever, and the cops had to remind him he’s just an old-ass Diplodocus. Play some more Brain Age, grandpa! Forget it. I’m gonna go hunt up Brazilsaurus and Uruguaysaurus and get a football game together… What do you mean, they haven’t been named yet? Aw, man!”

—————————————–

Pre-emptive note to the etymology mafia: yes, I know that Antarctosaurus means “southern lizard”, not “lizard from Antarctica”. But in this joke, Argentinosaurus is not so well-informed.

This imaginary interlude was brought to you by Becky Crew’s habit of putting words in animals’ mouths, and by Mike’s proposed moratorium on “place-saurus” names, and by the number 11.

being eaten 600

My friend, colleague, and sometime coauthor Dave Hone sent the above cartoon, knowing about my more-than-passing interest in sauropod neurology. It was drawn by Ed McLachlan in the early 1980s for Punch! magazine in the UK (you can buy prints starting at £18.99 here).

I know that this isn’t the only image in the “oblivious sauropods getting eaten” genre. There’s a satirical drawing in Bakker’s The Dinosaur Heresies showing a sleeping brontosaur getting its tail gnawed on by some pesky mammals. I’ll scan that and post it when I get time. I’m sure there must be others in a similar vein–point me to them in the comments or email me and I’ll post as many as I can get my hands on.

I wouldn’t post stuff like this if I didn’t think it was funny. But if you want the real scoop on whether sauropods could have responded quickly to injuries to their distant extremities, here’s the deal:

First of all, sauropods really did have individual sensory nerve cells that ran from their extremities (tip of tail, soles of feet)–and from the rest of their skin–to their brainstems. In the longest sauropods, these cells were probably something like 150 feet long, and may have been the longest cells in the history of life. We haven’t found any fossils of these nerves and almost certainly never will, but we can be sure that sauropods had them because all vertebrates do, from hagfish on up. That’s just how we’re built. (This is all rehash for regular readers–see this post and the linked paper.)

Wedel RLN fig2 480

So how long does it take to send a nerve impulse 150 feet? The fastest nerve conduction velocities are in the neighborhood of 120 meters per second, so a signal from the very tip of the tail in a 150-foot sauropod would take about half a second to reach the brain.

Is it possible that sauropods had accelerated nerve conduction velocities, to bring in those distant signals faster? To the brain, probably not. The only ways to speed up a nerve impulse are to increase the diameter of the axon itself, which some invertebrates do, and to increase the thickness of the myelin sheath around the axon, which is what vertebrates tend to do (some invertebrates have myelin-like tissues that apparently help accelerate their nerve impulses, too). Fatter axons mean fatter nerves, and for at least half the trip to the brain, the axons in question are part of the spinal cord. And we know that sauropod spinal cords were pretty small, relative to their body size, because the neural canals of their vertebrae, through which their spinal cords passed, are themselves small–Hatcher wrote about this more than a century ago. So there’s a tradeoff–sauropods could have had very fast, very fat axons, but not very many of them, and therefore poor “coverage” at their extremities, with nerve endings widely spaced, or better coverage with more axons, but those axons would be skinnier and therefore slower. We don’t know which way they went.

Incidentally, you can experiment with the density of sensory nerve endings in your own body. Close your eyes or blindfold yourself, and have a friend poke you in various places with chopsticks. Seriously–start with the two chopsticks right together, and gradually spread them out until you can feel two distinct points (or, if you want to get really tricky, have your friend mix up the close and widely spread touches so there’s no direction for you to anticipate). The least sensitive part of your body is your back–over your back and shoulders, you’ll probably have a hard time distinguishing points of touch that are less than an inch apart. On your hands and face, you’ll probably be able to distinguish points only a few millimeters apart; in fact, for fingertips you’ll probably need finer instruments than chopsticks–maybe toothpicks or pins, but I take no responsibility for any accidental acupuncture!

Back to sauropods. Could predators have taken advantage of the comparatively long nerve conduction velocities in sauropods? I seriously doubt it, for several reasons:

  • Simple reflex arcs are governed by interneurons in the spinal cord. The tail-tip-to-spinal-cord distance was a lot shorter than the tail-tip-to-brain route. Even over the round trip of “sensory impulse in, motor impulse out”, it would have been at worst equal, and that’s assuming the nerve impulse had to go all the way to the base of the tail.* Call it half a second, max.
  • It gets worse: the peripheral nerves outside the spinal cord are not limited by the size of the neural canal, so they can be more heavily myelinated, with faster conduction times. For example, each of the sciatic nerves running down the backs of your thighs is much larger in cross-section than your entire spinal cord. If sauropod peripheral nerves were selected for fast conduction, they might have been bigger and faster than anything around today.
  • Half a second is not much time for a theropod to formulate a plan, especially if Step 1 of the plan is “grab 150-foot sauropod by the tail”.
  • This assumes that said theropod was able to sneak right up to the sauropod without being detected. You go try that with a big wild herbivore and let me know how it works out. (Also, a big animal tolerating your presence, because you are pathetically small and weak, is not the same as it being unaware of your presence.)
  • All of this assumes the theropod only went for the bony whip-lash at the tip of the tail–the fastest-moving extremity, and the least-nourishing single bite anywhere on the target. If the theropod went for a meatier bite closer to the base of the tail, it would have to sneak closer to the sauropod’s head (better chance of being spotted), and the nerve conduction delay would be shortened.
  • A 150-foot sauropod would probably mass somewhere between 50 and 100 tons, and would be capable of dealing incredible damage to even the largest theropods, which maxed out around 15 tons. There’s a good reason predators go after the young, sick, and weak. Smaller sauropods would be less dangerous, but they’d also have faster tail-to-central-nervous-system-and-back reaction times.
  • A theropod big enough to go after a 150-foot sauropod would also be subject to fairly long nerve-conduction delays, which would limit whatever trifling advantage it might have gotten from such delays in the sauropod.

So, although I have no doubt that in their long history together, giant theropods did occasionally tackle full-grown giant sauropods–because real animals do all kinds of weird things if you watch them long enough, and lions will take on elephants when they get desperate–I am extremely skeptical that the theropods enjoyed any advantage based on the “slow” nervous systems of those sauropods.

* Some relevant hard-core anatomy for the curious: sauropods have neural canals in their tail vertebrae, and usually far down their tails, too. But that doesn’t mean much–you have neural canals to the bottom half of your sacrum, but your spinal cord stops around your first or second lumbar vertebra. From there on down, you just have nerve roots. So the shortest reflex arc from your big toe has to go up to your lower back and return. Why is your spinal cord so short? Basically because your central nervous system stops growing when you’re still a child–it will add new connections after that, and a few new cells in your olfactory bulbs and hippocampus, but it won’t get appreciably bigger or longer. After mid-childhood, your body keeps growing but your spinal cord stays the same length, so you end up with this freaky little-kid spinal cord tucked up inside your grown-up vertebral column. Weird, huh?

So did sauropod spinal cords stop at mid-back or go all the way into the tail? We have several conflicting lines of evidence. In extant reptiles, the spinal cord does extend into the tail in at least some taxa (I haven’t done anything like a complete survey, just read a couple of papers). Birds are no help because their tails are extremely short, but their spinal cords do extend into the synsacrum (and expand there, thanks to the glycogen body, which was probably also present in sauropods and responsible for the inaccurate “second brain” meme). But then birds grow up very fast, with even the largest reaching full size in a year or two, so they don’t share our problem of the body outgrowing the nervous system. We know that sauropods grew pretty quickly, but they also took a while to mature–somewhere between one and three decades, probably. Did that protracted growth period give their vertebral columns the time to outgrow their spinal cords? I have no idea, because the division of the spinal cord into roots happens inside the dura mater and doesn’t leave any skeletal traces that I know of. Someone should go figure it out–or at least figure out if it can be figured out!

Just sayin’:

gallery15[1]

vs.

apato-growth-series

(From here.)

Update

The rest of the Umbaran Starfighter Saga:

Yesterday, Matt showed you this starship from the Star Wars universe:

UmbaranStarfighter-SWE

And asked whether it’s based on a cervical vertebra of Apatosaurus.

Absolutely it is. It can’t be just a coincidence. Matt showed a lot of useful orthogonal views of various Apatosaurus cervicals in the last post, but here’s a a nice informative oblique view which is similar (though not identical) to that of the Umbaran ship:

apatosaurus-ajax-holotype--YPM-1860--left-anterolateral-DSCN5946

Apatosaurus ajax holotype YPM 1860, cervical vertebra of unspecified position but probably from around the middle of the neck, in left anterolateral view. This is the same vertebra that appears in the last three photos in Matt’s post.

(Because the vertebra photo was taken from higher up than the starfighter image, the condyle/cockpit appears lower on the vertebra. That is basically an effect of perspective rather than a difference in proportions.)

The questions for me are twofold: which Apatosaurus vertebra is it based on, and who did it?

What vertebra is it based on?

In some ways, the cervical that it most resembles is this classic: C?8 of the Apatosaurus excelsus holotype YPM 1980:

Ostrom and McIntosh (1966:plate 12) -- Brontosaurus excelsus YPM 1980 cervical 8

This one resembles the starfighter in the very deep cervical rob loops — deep even for Apatosaurus — and in the small, high condyle. It also resembled the ship in the absence of neural-spine metapophyses (due to breakage, not taxonomically significant variation, alas). The result of their absence is that the “upper wings” (i.e. postzygapophyseal rami) are swept up, out and back, as in the ship.

But in other respects it’s very different — notably the very elongate prezyg rami (an effect exaggerated by the breakage) and the more or less parallel trajectories of the top and bottom margins of the loop.

Another candidate would be the one that appears in the top left part of figure 7 (“the freak gallery”) from our recent neck-anatomy paper on arXiv:

This on is rather bulky for a model for the ship, but does have a less wrong shape of the cervical rib loops. And the damage that blew off both the prezygs and the metapophyses leaves the isolated “wings” on the top, just as in the ship.

I think the best model I can find for the starfighter is probably C8 of the Apatosaurus louisae holotype CM 3018, which Gilmore illustrated beautifully in his 1936 monograph but which unfortunately I only have as this bad scan:

Gilmore1936-plateXXIV-C8

It has good, deep cervical-rib loops; a definite bend from the fairly lateral upper part to the more ventrally inclined lateral part; a high, fairly small condyle; and a definite bulges where the parapophysis fuses with the cervical ribs, corresponding to the weapons pods of the starfighter.

And yet, and yet …

I can’t shake the feeling that I’ve seen another Apatosaurus vertebra somewhere that is a more or less perfect fit for the ship. But I can’t remember where I saw it. Come to that, I can’t think what specimen it could be from, if not one of those that Matt and I have shown in these posts.

Anyone?

Whose work is it?

The other mystery is — whose work is this design, and where did he or she get the shape from? In a comment on the last post, I said to Matt that “one can hardly help but suspect that Jarrod did it on your instruction”. (Jarrod is an old friend of Matt’s who works in digital effects for film and TV.) But  Matt insists it’s none of his doing — and I must say that if it had been in any way his work, he would have been shouting about it long before now.

So how did it happen?

Anyone know?

Update

The Umbaran Starfighter is an Apatosaurus vertebra–check out the rest of the saga:

UmbaranStarfighter-SWE

Update, January 21, 2013: YES, it was! Scroll down for links to the entire saga.

Because it’s doing a hell of an impression of one, if not. It’s got the huge cervical rib loops (wings), bifurcated neural spine (top fins), and even a condyle on the front of the centrum (cockpit pod). About all it’s missing are the zygapophyses and the cervical ribs themselves.

Some actual Apatosaurus cervicals for comparison, from previous posts:

Apatosaurus ajax NSMT-PV 20375, cervical vertebrae 3, 6 and 7 in anterior and posterior views. Modified from Upchurch et al. (2005: plate 2)

Apatosaurus parvus CM 563/UWGM 15556 cervicals 7, 5, 4 and 3 in anterior and right lateral views, from Gilmore (1936:pl. 31)

Various Apatosaurus cervicals–see Wedel and Sanders (2002) for specimen numbers and sources.

And of course Mike’s magisterial work photographing the Apatosaurus ajax holotype YPM 1860 cervical:

More on the Umbaran Starfighter here.

The complete Umbaran Starfighter Saga–at least as told on SV-POW!:

For other Star Wars/paleontology crossovers, please see:

The sauropods of Star Wars

The sauropods of Star Wars: Special Edition

and–mostly as shameless self-promotion since the paleo link is pretty tenuous:

Tales of the Flaming Vagabond

References

One of our anatomy students this year, Tess MacFife, was inspired by the other Dr. Wedel’s skull lecture and produced this excellent anatomy-inspired jack-o-lantern:

Random passers-by probably thought this was some kind of bat/demon/Lovecraftian horror, but those in the know would recognize it as the human sphenoid bone in anterior view. Tess writes, “Full disclosure, I did print out a template and used toothpicks for the outline.” Here’s her template image, borrowed from here.

Any other anatomy- or paleontology-inspired Halloween geekery this year? Feel free to alert us in the comments. And well done, Tess!

Here’s a blast from the past:

This alleged Compsognathus is a card from the “Flesh” card-game that was printed across several progs (issues) of the comic 2000 AD in 1977. This one is from the back cover of Prog 9. (Click through the picture for the whole back cover.)

“Flesh” was one of the half-dozen or so stories that appeared each week in those early months of 2000 AD. It was the story of how cowboys of the future travelled back to the Mesozoic to harvest dinosaurs for their meat, and was the subject of Jeff Liston’s chapter in the recentish Geological Society volume on the history of dinosaur research.

Compsognathus made another pop-culture appearance in The Lost World: Jurassic Park, of course, as the cute little “compys” that tear one of the nastier human characters to pieces.

Why does the 2000 AD Compsognathus have actinopterygian-like fins for arms? According to Wikipedia, The idea comes from Bidar et al. (1972), who supposed that the French specimen had webbed forefeet, which would look like flippers in life — an idea illustrated as part of a larger scene by Halstead (1975):

John Ostrom’s (1978) Compsognathus monograph showed that this was nonsense, but of course that was too late for the early issues of 2000 AD.

References

Bidar, A.; Demay L., Thomel G. 1972. Compsognathus corallestris, une nouvelle espèce de dinosaurien théropode du Portlandien de Canjuers (Sud-Est de la France). Annales du Muséum d’Histoire Naturelle de Nice 1:9–40.

Halstead L.B. 1975. The evolution and ecology of the dinosaurs. Eurobook. ISBN 0-85654-018-8.

Ostrom, J.H. 1978. The osteology of Compsognathus longipesZitteliana 4:73–118.

Update 1 (the next day)

In a comment below, Andrea Cau points to this post on his blog Theropoda (“the most inclusive blog containing Allosaurus fragilis but not Saltasaurus loricatus) which contains two more flippered-Compsognathus illustrations. Here they are: one from David Lambert’s book Dinosaur! …

… and one from David Norman’s Illustrated Encyclopedia of Dinosaurs.

Update 2 (two days later)

Silly me, I should of course have posted Bidar et al.’s (1972) own life restoration of Compsognathus. It’s not great art, but it’s … actually, I’m not sure what it is. But anyway, here it is:

Attempted reconstruction of attitudes of Compsognathus corallestris nov. sp. A, erect stance (walking); B, sitting (inspired by O. Abel); C, Swimming; D, Diving. (Bidar et al. 1972:figure 21)

 

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