An important paper is out today: Carpenter (2018) names Maraapunisaurus, a new genus to contain the species Amphicoelias fragillimus, on the basis that it’s actually a rebbachisaurid rather than being closely related to the type species Amphicoelias altus.

Carpenter 2018: Figure 5. Comparison of the neural spine of Maraapunisaurus fragillimus restored as a rebbachisaurid (A), and the dorsal vertebrae of Rebbachisaurus garasbae (B), and Histriasaurus boscarollii (C). Increments on scale bars = 10 cm.

And it’s a compelling idea, as the illustration above shows. The specimen (AMNH FR 5777) has the distinctive dorsolaterally inclined lateral processes of a rebbachisaur, as implied by the inclined laminae meeting at the base of the SPOLs, and famously has the very excavated and highly laminar structure found in rebbachisaurs — hence the species name fragillimus.

Ken’s paper gives us more historical detail than we’ve ever had before on this enigmatic and controversial specimen, including extensive background to the excavations. The basics of that history will be familiar to long-time readers, but in a nutshell, E. D. Cope excavated the partial neural arch of single stupendous dorsal vertebra, very briefly described it and illustrated it (Cope 1878), and then … somehow lost it. No-one knows how or where it went missing, though Carpenter offers some informed speculation. Most likely, given the primitive stabilisation methods of the day, it simply crumbled to dust on the journey east.

Carpenter 2018: Frontispiece. E. D. Cope, the discoverer of AMNH FR 5777, drawn to scale with the specimen itself.

Cope himself referred the vertebra to his own existing sauropod genus Amphicoelias — basically because that was the only diplodocoid he’d named — and there it has stayed, more or less unchallenged ever since. Because everyone knows Amphicoelias (based on the type species A. altus) is sort of like Diplodocus(*), everyone who’s tried to reconstruct the size of the AMNH FR 5777 animal has done so by analogy with Diplodocus — including Carpenter himself in 2006, Woodruff and Foster (2014) and of course my own blog-post (Taylor 2010).

(*) Actually, it’s not; but that’s been conventional wisdom.

Ken argues, convincingly to my mind, that Woodruff and Foster (2014) were mistaken in attributing the great size of the specimen to a typo in Cope’s description, and that it really was as big as described. And he argues for a rebbachisaurid identity based on the fragility of the construction, the lamination of the neural spine, the extensive pneumaticity, the sheetlike SDL, the height of the postzygapophyses above the centrum, the dorsolateral orientation of the transverse processes, and other features of the laminae. Again, I find this persuasive (and said so in my peer-review of the manuscript).

Carpenter 2018: Figure 3. Drawing made by E.D. Cope of the holotype of Maraapunisaurus fragillimus (Cope, 1878f) with parts labeled. “Pneumatic chambers*” indicate the pneumatic cavities dorsolateral of the neural canal, a feature also seen in several rebbachisaurids. Terminology from Wilson (1999, 2011) and Wilson and others (2011).

If AMNH FR 5777 is indeed a rebbachisaur, then it can’t be a species of Amphicoelias, whose type species is not part of that clade. Accordingly, Ken gives it a new generic name in this paper, Maraapunisaurus, meaning “huge reptile” based on Maraapuni, the Southern Ute for “huge” — a name arrived at in consultation with the Southern Ute Cultural Department, Ignacio, Colorado.

How surprising is this?

On one level, not very: Amphicoelias is generally thought to be a basal diplodocoid, and Rebbachisauridae was the first major clade to diverge within Diplodocoidae. In fact, if Maraapunisaurus is basal within Rebbachisauridae, it may be only a few nodes away from where everyone previously assumed it sat.

On the other hand, a Morrison Formation rebbachisaurid would be a big deal for two reasons. First, because it would be the only known North American rebbachisaur — all the others we know are from South America, Africa and Europe. And second, because it would be, by some ten million years, the oldest known rebbachisaur — irritatingly, knocking out my own baby Xenoposeidon (Taylor 2018), but that can’t be helped.

Finally, what would this new identity mean for AMNH FR 5777’s size?

Carpenter 2018: Figure 7. Body comparisons of Maraapunisaurus as a 30.3-m-long rebbachisaurid (green) compared with previous version as a 58-m-long diplodocid (black). Lines within the silhouettes approximate the distal end of the diapophyses (i.e., top of the ribcage). Rebbachisaurid version based on Limaysaurus by Paul (2016), with outline of dorsal based on Rebbachisaurus; diplodocid version modified from Carpenter (2006).

Because dorsal vertebrae in rebbachisaurids are proportionally taller than in diplodocids, the length reconstructed from a given dorsal height is much less for rebbachisaurs: so much so that Ken brings in the new version, based on the well-represented rebbachisaur Limaysaurus tessonei, at a mere 30.3 m, only a little over half of the 58 m he previously calculated for a diplodocine version. That’s disappointing for those of us who like our sauropods stupidly huge. But the good news is, it makes virtually no difference to the height of the animal, which remains prodigious — 8 m at the hips, twice the height of a giraffe’s raised head. So not wholly contemptible.

Exciting times!

References

 

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Foster and Wedel 2014 fig 3 - dorsals

Fig. 3. MWC 8028, Haplocanthosaurus dorsal vertebrae. A. Lateral view of dorsal centrum with bottom edge of lateral pneumatic fossa preserved. B. Dorsal view of same centrum as in A, showing the median septum between the paired lateral fossae. C. Lateral view of dorsal centrum with smaller segment of the lateral pneumatic fossa margin preserved. D. Dorsal view of same centrum as in C, again showing the median septum and paired lateral fossae. E. Lateral view of dorsal centrum with partial pleurocoel preserved. F. Cross-sectional (posterior) view of same dorsal as in E. G. Dorsal neural spines in lateral (top) and anterior or posterior (center, bottom) views. Scale bars = 10 cm.

Right on the heels of Aquilops last week, my paper with John Foster on the new specimen of Haplocanthosaurus from Snowmass, Colorado, was just published in Volumina Jurassica. I’ll have more to say about it later, but right now I am up against a deadline on a big project and I need to go work on that. I’m only popping up here to note two quick things.

First, if you’re not familiar with Volumina Jurassica – and I wasn’t, before this project – it’s a free-to-access* journal that publishes papers on all aspects of the Jurassic. The current issue is specifically dedicated to the Jurassic formations of the American West. There’s a lot of interesting stuff in there, but of special interest to SV-POW! readers will be the paper by Cary Woodruff and John Foster on the legendary and possibly apocryphal Amphicoelias fragillimus.

* But not truly open access since the journal claims to retain exclusive rights to distribute the papers. That seems like a quaint affectation now that the internet is here, but whatever – at least they let anyone download the PDF for free, which is primarily what I care about.

Foster and Wedel 2014 fig 4 - sacrum

Fig. 4. Sacra of Haplocanthosaurus.  A. MWC 8028, sacrum in right lateral view. B. MWC 8028, close-up of S4 and S5 centra highlighting pneumatic fossae. C. MWC 8028 with divisions between the vertebrae overlaid. D. CM 879, sacrum in right lateral view with divisions between the vertebrae overlaid. E. CM 572 in right lateral view, after Hatcher (1903c: plate 4). B–E are not shown at the same scale, scale bar for A = 20 cm. Note that the neural arches in CM 572 were restored during preparation, and the sacral neural spines as shown here are probably lower than they would have been in life.

Second, the figure resolution in the PDF of the Haplocanthosaurus paper is not stellar, so as is the case with almost all of our papers, the full-color, high-resolution figures are available at the paper’s page on the sidebar.

Gotta run.

For our previous posts on Haplocanthosaurus, go here; for those on Amphicoelias, including Mike’s very popular, “How big was Amphicoelias fragillimus? I mean, really?”, go here.

References

Although it would be nice to think that our site views have octupled in the last day because of Mike’s fine and funny posts about what search terms bring people to SV-POW!, the real reason is that we were blessed by incoming links from both pages of this Cracked.com article.

Now, as any person who has ever accomplished anything whatsoever knows, it is super-important to avoid Cracked.com or you’ll still be up 23 hours from now reading, “6 Mind-Blowing Ways that Comedy Writers are Secretly Destroying Your Productivity”. (I’m kidding, that article doesn’t really exist–but if it did, I’m sure it would consist entirely of descriptions and links to six other Cracked articles). But that’s only true because most of the articles there hit the sweet spot at the intersection of funny, surprisingly informative, mercifully short, and well-written. Crack.com would be a more honest URL, but I assume it was taken.

Anyway, I’d like to return the favor, so here’s a list of the 6 SV-POW! Posts Most Likely to Blow the Minds of Cracked.com Readers. If I missed some goodies or recommended some stinkers, let me know–the comment thread is open.

Amphicoelias vert reconstruction by Mike

1.How big was Amphicoelias fragillimus? I mean, really?

Who doesn’t want to read about the bizarre real-world mystery surrounding what might have been the world’s largest dinosaur? If you’re not sold, consider that the picture above shows a single vertebra that was–or at least might have been–seven and a half feet tall.

long nerves of sauropods

2. Oblivious sauropods being eaten

The mercifully short version of this much longer post, in which I consider the consequences of the world’s largest animals having the world’s longest cells.

krayt-cervicals

3. The sauropods of Star Wars

Weapons-grade anatomical pedantry.

Umbaran starfighters

4. CONFIRMED: the Umbaran Starfighter is an Apatosaurus cervical

Yes, there is a ship in Star Wars: The Clone Wars that is basically a flying dinosaur vertebra. It took us about five weeks to unravel that story–the post linked above has links to the rest of the saga.

blue-whale-and-brachiosaurus

5. SV-POW! showdown: sauropods vs whales

Our original linkbait post. Don’t miss the shorter follow-up with more critters.

Is that your flexor tubercle, Saurophaganax, or are you just hungry to see me?

Is that your flexor tubercle, Saurophaganax, or are you just hungry to see me?

6. Friday phalanges: Megaraptor vs Saurophaganax

A deliberately goofy post in which I wax poetic about the largest predatory dinosaur claws ever discovered.

So, that was a big pile of superlatives and Star Wars. If you’re hungry for more substantial fare, you might start with our Tutorials page or our Things to Make and Do series on dissecting and skeletonizing modern animals. We also blog a lot about the evils of obstructive publishers and the need for open access to the scientific literature–you can find those posts on our Shiny Digital Future page.

flaming-vagabond-in-firefly-niska-station

A parting shot in my desperate quest for attention: this Star Wars ship flying around in the background in Firefly and Serenity is at least partly my fault–full story here. Oh, and my co-blogger Mike Taylor has written an insightful and affordable book about Doctor Who; read about it here.

From Jensen (1987, page 604):

“In 1985 I found the proximal third of an extremely large sauropod femur (Figs. 8A, 12A) in a uranium miner’s front yard in southern Utah.  The head of this femur is 1.67 m (5’6″) in circumference and was collected from the Recapture Creek Member of the the Morrison Formation in Utah near the Arizona border.  It is the largest bone I have ever seen.”

Jensen included not one but two figures of this immense shard of excellence. Here they are:

Jensen 1987 figure 8

Jensen 1987, Figure 8

Jensen 1987, Figure 12

Jensen 1987, Figure 12

The specimen was heavily reconstructed, as you can see from the big wodge of unusually smooth and light-colored material in the photo. So we can’t put much stock in that part of the specimen.

Unfortunately, the only measurement of the specimen that Jensen gives in the paper is that circumference; there are no straight-line linear measurements, and the figures both have the dreaded scale bars. Why dreaded? Check this out:

Recapture Creek figs 8 and 12 comparedAs you can see, when the scale bars are set to the same size, the bones are way off (the scale bar in the drawing is 50 cm). This is not an uncommon problem. I make the Fig 8 version 30% bigger in max mediolateral width of the entire proximal end, and still 17% bigger in minimum diameter across the femoral head, as measured from the slight notch on the dorsal surface (on the right in this view).

Can we figure out which is more accurate based  on the internal evidence of the paper? For starters, the Fig 12 version is a drawing (1), that does not match the outline from the photo (2), and the hand-drawn scale bar (3) does not actually coincide with any landmarks (4), and that’s plenty of reasons for me not to trust it.

What about that circumference Jensen mentioned? Unfortunately, he didn’t say exactly where he took it, just that the head of the femur had a circumference of 1.67 meters. Is that for the entire proximal end, or for the anatomical head that fits in the acetabulum, er wot? I’m afraid the one measurement given in the paper is no help in determining which of the figures is more accurately scaled.

The obvious thing to do would be to see if this bone is in the BYU collections, and just measure the damn thing. More on that at the end of the post.

In the meantime, Jensen said that the shape of the Recapture Creek femur was most similar to the femur of Alamosaurus, or to that of Brachiosaurus among Morrison taxa, and he referred it to Brachiosauridae. So how does this thing–in either version–compare with the complete femur of FMNH P25107, the holotype of Brachiosaurus altithorax?

The Recapture Creek brachiosaur femur fragment compared to the complete femur of the Brachiosaurus altithorax holotype FMNH P25107

The Recapture Creek femur fragment compared to the complete femur of the Brachiosaurus altithorax holotype FMNH P25107

The first thing to notice is that the drawn outline from Figure 12 is a much better match for the Brachiosaurus altithorax femur–enough so that I wonder if Jensen drew it from the Recapture Creek specimen, or just traced the B.a. proximal femur and scaled it accordingly (or maybe not accordingly, since the scale bars don’t match).

But let’s get down to business: how long would the complete femur have been?

Using the scale bar in the photograph from Figure 8 (on the left in above image), I get a total femur length of 2.36 meters. Which is long, but only 7.7% longer than the 2.19-meter femur of FMNH P25107, and therefore only 25% more massive. So, 35 tonnes to Mike’s 28-tonne B.a., or maybe 45 tonnes to a more liberal 36-tonne B.a. Big, yeah, but not world-shattering. Update 2014-05-19: I don’t know where I got the 2.19-meter femur length for Brachiosaurus altithorax, but it’s a mistake. So the rest of that paragraph should read: Which is 16% longer than 2.03-meter femur of FMNH P25107, and therefore 57% more massive. So, 44 tonnes to Mike’s 28-tonne B.a., or maybe 57 tonnes to a more liberal 36-tonne B.a. That’s nowhere near the 2.5-meter femur and estimated 70-tonne mass of the largest Argentinosaurus, but it’s pretty darned good for a brachiosaur.

Using the scale bar in the drawing from Figure 12 (on the right in the above image)–which, remember, is 50 cm, not 1 meter–I get a total femur length of about 1.9 meters, which is considerably smaller than the B.a. holotype. That is very much at odds with Jensen’s description of it as “the largest bone I have ever seen”, and given that we have many reasons for not trusting the scale bar in the drawing, it is tempting to just throw it out as erroneous.

So it would seem that unless Jensen got both scale bars too big, the Recapture Creek brachiosaur was at most only a shade bigger than the holotype specimen of Brachiosaurus altithorax.

But wait–is the Recapture Creek brachiosaur a brachiosaur at all? Jensen didn’t list any characters that pushed him toward a brachiosaurid ID, and I don’t know of any proximal femur characters preserved in the specimen that would separate Brachiosaurus from, say, Camarasaurus. And in fact a camarasaur ID has a lot to recommend it, in that Camarasaurus femora have very offset heads (the ball- or cylinder-like articular surface at the top end sticks out a big more to engage with the hip socket–see Figure 12 up near the top of the post), moreso than in many other Morrison sauropods, and that would make them better matches for the Recapture Creek femur photo. Here’s what the comparo looks like:

Recapture Creek - Camarasaurus comparo

The Recapture Creek femur fragment compared with a complete femur of Camarasaurus.

I make that a 2.07-meter femur using the photo on the left, and a 1.66-meter femur using the drawing on the right. The one decent femur in the AMNH 5761 Camarasaurus supremus collection is 1.8 meters long, so these results are surprisingly similar to those for the B. althithorax comparison–the drawing gives a femur length shorter than the largest known specimens, and the photo gives a length only slightly longer. A camarasaur with a 2.07 meter femur would be 15% larger than the AMNH C. supremus in linear terms, and  assuming isometric scaling, 1.5 times as massive–maybe 38 tonnes to AMNH 5761’s estimated 25. A big sauropod to be sure, but not as big as the largest apatosaurs, and not nearly as big as the largest titanosaurs.

I have always been surprised that the Recapture Creek femur frag has attracted so little attention, given that “Dinosaur Jim” himself called it the biggest bone he had ever seen. But it appears that the lack of attention is justified–whether it was a brachiosaur or a camarasaur, and using the most liberal estimates the scale bars allow, it simply wasn’t that big.

Update about half an hour later: Okay, maybe I was a little harsh here. IF the photo scale bar is right, the Recapture Creek femur might still represent the largest and most massive macronarian from the Morrison Formation (Edit: only if it’s a brachiosaur and not a camarasaur; see this comment), which is something. I suppose I was particularly underwhelmed because I was expecting something up in OMNH 1670-to-Argentinosaurus territory, and so far, this ain’t it. I’ll be interested to see what the actual measurements say (read on).

The Moral of This Story

So, if it wasn’t that big after all, and if no-one has made a stink about it being big before now, why go to all this trouble? Well, mostly just to satisfy my own curiosity. If there was a truly gigantic brachiosaur from the Morrison, it would be relevant to my interests, and it was past time I crunched the numbers to find out.

But along the way something occurred to me: this should be a cautionary tale for anyone who gets all wound up about the possible max size of Amphicoelias fragillimus. As with A. fragillimus, for the Recapture Creek critter we have part of one bone, and at least for this exercise I was working only from published illustrations with scale bars. And as with A. fragillimus, the choice of a reference taxon is not obvious, and the size estimates are all over the place, and some of them just aren’t that big.

It always amuses me when A. fragillimus comes up and people (well, trolls) accuse us of being big ole’ wet blankets that just don’t want to believe in 200-tonne sauropods. It amuses me because it’s wrong on so many levels. Believe me, when we have our sauropod fanboy hats on, we most definitely do want to believe in 200-tonne sauropods. That would rock. But when we put our scientist hats on, wanting and belief go right out the window. We have to take a cold, hard look at the data, and especially at its limitations.

Oh, the other moral is to go buy a tape measure, and use it. Sheesh!

Coda

As I said above, the obvious thing to do would be to just track down the bone and measure it. It does still exist, it’s in the BYU collections, and Brooks Britt has kindly offered to send along some measurements when he gets time. So we should have some real answers before long (and here they are). But I wanted to work through this example without them, to illustrate how much uncertainty creeps in when trying to estimate the size of a big sauropod from published images of a single partial bone.

Reference

Jensen, J.A. 1987. What I did on my holidaysNew brachiosaur material from the Late Jurassic of Utah and Colorado. Great Basin Naturalist 47(4): 592-608.

In our PeerJ neck-anatomy paper, we speculated on how long individual cervical vertebrae might have grown. Here is the relevant section:

Mere isometric scaling would of course suffice for larger animals to have longer necks, but Parrish (2006, p. 213) found a stronger result: that neck length is positively allometric with respect to body size in sauropods, varying with torso length to the power 1.35. This suggests that the necks of super-giant sauropods may have been even longer than imagined: Carpenter (2006, p. 133) estimated the neck length of the apocryphal giant Amphicoelias fragillimus Cope, 1878 as 16.75 m, 2.21 times the length of 7.5 m used for Diplodocus, but if Parrish’s allometric curve pertained then the true value would have been 2.21^1.35 = 2.92 times as long as the neck of Diplodocus, or 21.9 m; and the longest single vertebra would have been 187 cm long.

Now this speculation is shot through with uncertainty. As we’ve discussed before, at length, all estimates of Amphicoelias fragillimus length and mass are wildly speculative; and Parrish’s allometry result was extrapolated from an unconvincingly small data set. But still, these numbers are probably the best we can do with what we have.

In Diplodocus carnegii, C14 is the longest individual vertebra at 642 mm long (Hatcher 1901, p. 38). The Amphicoelias:Diplodocus size ratio of 2.21 from Carpenter and the neck allometry constant of 1.35 from Parrish suggest that the corresponding vertebra in the big boy would have been 2.92 times as long as that 642 mm, hence the 187 cm that we reported.

So what does a 187-cm long cervical vertebra look like? Scaling up from the Diplodocus carnegii C14 in Hatcher (1901: plate III) and using my good self as a scalebar, here it is:

amphicoelias-fragillimus-c14-whiteBG

I find that just a little bit frightening. In more ways than one.

References

  • Carpenter, Kenneth. 2006 Biggest of the big: a critical re-evaluation of the mega-sauropod Amphicoelias fragillimus (Cope, 1878). New Mexico Museum of Natural History and Science Bulletin 36:131.
  • Cope, Edward D. 1878. Geology and paleontology: a new species of Amphicoelias. The American Naturalist 12:563.
  • Hatcher, Jonathan B. 1901. Diplodocus (Marsh): its osteology, taxonomy and probable habits, with a restoration of the skeleton. Memoirs of the Carnegie Museum 1:1-63 and plates I-XIII.
  • Parrish, J. Michael. 2006. The origins of high browsing and the effects of phylogeny and scaling on neck length in sauropodomorphs. pp 201-224 in: Amniote paleobiology, University of Chicago Press, Chicago.

I have a new paper out:

Wedel, M.J. 2012. A monument of inefficiency: the presumed course of the recurrent laryngeal nerve in sauropod dinosaurs. Acta Palaeontologica Polonica 57(2):251-256.

Update June 6, 2012: the final version was formally published yesterday, so the rest of this paragraph is of historical interest only. Like Yates et al. on prosauropod pneumaticity, it is “out” in the sense that the manuscript has been through peer review, has been accepted for publication, and is freely available online at Acta Palaeontologica Polonica. Technically it is “in press” and not published yet, but all that formal publication will change is to make a prettier version of the paper available. All of the content is available now, and the paper doesn’t include any of those pesky nomenclatural acts, and so, as with the prosauropod pneumaticity paper, I don’t see any reason to pretend it doesn’t exist. Think of the accepted manuscript as the caterpillar to the published version’s butterfly: different look, but same genome.

This one came about because last summer I read a review of Richard Dawkins’s book, The Greatest Show on Earth: The Evidence for Evolution. The review mentioned that the book includes a lengthy discussion of the recurrent laryngeal nerve (RLN) in the giraffe, which is a spectacularly dumb piece of engineering and therefore great evidence against intelligent design creationism. It wasn’t the first time I’d heard of the RLN, of course. It’s one of the touchstones of both human anatomy and evolutionary biology; anatomy because of its clinical importance in thyroid surgery, known for more than two millennia, and evolutionary biology because it is such a great example of a developmental constraint. (Dawkins’s coverage of all of this is great, BTW, and you should read the book.)

No, the reason the book review inspired me to write the paper was not because the RLN was new to me, but because it was overly familiar. It is a cool piece of anatomy, and its fame is justly deserved–but I am sick and tired of seeing the stinkin’ giraffe trotted out as the ultimate example of dumb design. My beloved sauropods were way dumber, and it’s time they got some credit.

But first, let’s talk about that nerve, and how it got to be there.

No necks for sex? How about no necks for anybody!

Embryos are weird. When you were just a month old (counting from fertilization), you had a set of pharyngeal arches that didn’t look radically different from those of a primitive fish. These started out quite small, tucked up underneath your comparatively immense brain, and each pharyngeal arch was served by a loop of artery called an aortic arch. What we call the arch of the aorta in an adult human is a remnant of just one of these embryonic aortic arches, and as you’ve no doubt noticed, it’s down in your chest, not tucked up next to your brain. When you were in the embryonic stages I’m talking about, you didn’t yet have a neck, so your brain, pharyngeal arches, aortic arches, and the upper parts of your digestive system were all smooshed together at your front end.

One thing you did have at that stage was a reasonably complete peripheral nervous system. The nerve cell bodies in and near your central nervous system sent out axons into the rest of your body, including your extremities. Many of these axons did not persist; they failed to find innervation targets and their parent neurons died. Imagine your embryonic central nervous system sending out a starburst of axons in all directions, and some of those axons finding targets and persisting, and others failing and dying back. So the architecture of your nervous system is the result of a process of selection in which only some cells were successful.

Crucially, this radiation and die-off of axons happened very early in development, when a lot of what would become your guts was still hanging under your proportionally immense brain like the gondola on a blimp. This brings us to the recurrent laryngeal nerve.

Going back the way we came

The fates of your embryonic pharyngeal arches are complex and I’m not going to do a comprehensive review here (go here for more information). Suffice it to say that the first three arches give rise to your jaws and hyoid apparatus, the fourth and sixth form your larynx (voicebox), and fifth is entirely resorbed during development. Update: I made a pharyngeal arch cheat sheet.

There are two major nerves to the larynx, each of which is bilaterally paired. The nerve of the fourth pharyngeal arch becomes the superior laryngeal nerve, and it passes cranial to the fourth aortic arch. The nerve of the sixth pharyngeal arch becomes the inferior or recurrent laryngeal nerve, and it passes caudal to the sixth aortic arch. At the time that they form, both of these nerves take essentially straight courses from the brainstem to their targets, because you’re still in the blimp-gondola stage.

If you were a shark, the story would be over. The more posterior pharyngeal arches would persist as arches instead of forming a larynx, each arch would hold on to its artery, and the nerves would all maintain their direct courses to their targets.

The normal fate of the aortic arches in humans. From http://education.yahoo.com/reference/gray/subjects/subject/135

But you’re not a shark, you’re a tetrapod. Which means that you have, among other things, a neck separating your head and your body. And the formation of your neck shoved your heart and its associated great vessels down into your chest, away from the pharyngeal arches. This was no problem for the superior laryngeal nerve, which passed in front of the fourth aortic arch and could therefore stay put. But the inferior laryngeal nerve passed behind the sixth aortic arch, so when the heart and the fourth and sixth aortic arches descended into the chest, the inferior laryngeal nerve went with them. Because it was still hooked up to the brainstem and the larynx, it had to grow in length to compensate.

As you sit reading this, your inferior laryngeal nerves run down your neck into your chest, loop around the vessels derived from the fourth and sixth aortic arches (the subclavian artery on the right, and the arch of the aorta and ductus arteriosus on the left) and run back up your neck to your larynx. Because they do this U-turn in your chest and go back the way they came, the inferior laryngeal nerves are said to ‘recur’ to the larynx and are therefore more commonly referred to as the recurrent laryngeal nerves (RLNs).

An enlightening diversion

The RLN is the poster child for “unintelligent design” because it is pretty dumb. Your RLNs travel a heck of a lot farther to reach your larynx than they ought to, if they’d been designed. Surely an intelligent designer would have them take the same direct course as the superior laryngeal nerve. But evolution didn’t have that option. Tetrapod embryos could not be built from the ground up but had to be modified from the existing “sharkitecture” of ancestral vertebrates. The rules of development could not be rewritten to accommodate a shorter RLN. Hence Dawkins’s love affair with the RLN, which gets 7 pages in The Greatest Show on Earth. He also appeared on the giraffe episode of Inside Nature’s Giants, in which the RLN was dug out of the neck and the continuity of its ridiculous path was demonstrated–probably the most smack-you-in-the-face evidence for evolution that has ever been shown on television (said the rabid fan of large-tetrapod dissections).

Incidentally, the existence and importance of the RLN has been known since classical times. The RLN innervates the muscles responsible for speech, and on either side it passes right behind the thyroid gland, which is subject to goiters and tumors and other grotesque maladies. So a careless thyroidectomy can damage one or both of the RLNs; if one gets snipped, the subject will be hoarse for the rest of his or her life; if both are cut, the subject will be rendered mute. The Roman physician Galen memorably demonstrated this by dissecting the neck of an immobilized but unanesthetized pig and isolating the RLNs (Kaplan et al. 2009). One moment the poor pig was squealing its head off–as any of us would be if someone dug out our RLNs without anesthesia–and the next moment Galen severed the RLNs and the animal abruptly fell silent, still in unbelievable pain but now without a mechanism to vocally express its discomfort.

Galen versus pig. Figure 2 from Kaplan et al. 2009.

The name of the nerve also goes back to Galen, who wrote:

I call these two nerves the recurrent nerves (or reversivi) and those that come upward and backward on account of a special characteristic of theirs which is not shared by any of the other nerves that descend from the brain.

Like at least some modern surgeons, Galen does not seem to have been overly burdened by humility:

All these wonderful things, which have now become common property, I was the first of all to discover, no anatomist before me ever saw one of these nerves, and so all of them before me missed the mark in their anatomical description of the larynx.

Both of those quotes are from Kaplan et al. (2009), which is a fascinating paper that traces the knowledge of the recurrent laryngeal nerve from classical times to the early 20th century. If you’d like a copy and can’t get hold of one any other way, let me know and I’ll hook you up.

Share and share alike

By now you can see where this is going: all tetrapods have larynges, all tetrapods have necks, and all tetrapods have recurrent laryngeal nerves. Including giraffes, much to the delight of Richard Dawkins. And also including sauropods, much to the delight of yours truly.

Now, I cannot show you the RLN in a living sauropod, nor can I imagine a scenario in which such a delicate structure would be recognizably preserved as a fossil. But as tetrapods, sauropods were bound to the same unbreakable rules of development as everything else. The inference that sauropods had really long, really dumb RLNs is as secure as the inference that they had brainstems, hearts, and larynges.

Wedel (2012) Fig. 1. Course of the left vagus nerve and left recurrent laryngeal nerve in a human, a giraffe, and Supersaurus. The right recurrent laryngeal nerve passes caudal to the right subclavian artery rather than the aorta and ductus arteriosus, but otherwise its course is identical to that of the left.

Giraffes have necks up to 2.4 meters long (Toon and Toon 2003), so the neurons that make up their RLNs approach 5 meters in the largest indiividuals. But the longest-necked sauropods had necks 14 meters long, or maybe even longer, so they must have had individual neurons at least 28 meters long. The larynx of even the largest sauropod was probably less than 1 meter away from the brainstem, so the “extra” length imposed on the RLN by its recurrent course was something like 27 meters in a large individual of Supersaurus. Take that, Giraffa.

Inadequate giraffe is inadequate.

One way or another

It is possible to have a nonrecurrent laryngeal nerve–on one side, anyway. If you haven’t had the opportunity to dissect many cadavers, it may come as a surprise to learn that muscles, nerves, and blood vessels are fairly variable. Every fall in Gross Anatomy at WesternU, we have about 40 cadavers, and out of those 40 people we usually have two or three with variant muscles, a handful with unusual branching patterns of nerves, and usually half a dozen or so with some wackiness in their major blood vessels. Variations of this sort are common enough that the better anatomy atlases illustrate not just one layout for, say, the branching of the femoral artery, but 6-10 of the most common patterns. Also, these variations are almost always asymptomatic, meaning that they never cause any problems and the people who have them usually never know (ask Mike about his lonely kidney sometime). You–yes, you, gentle reader!–could be a serious weirdo and have no idea.

Variations in the blood vessels seem to be particularly common, possibly because the vessels develop in situ with apparently very little in the way of genetic control. Most parts of the body are served by more than one artery and vein, so if the usual vessel isn’t there or takes an unusual course, it’s often no big deal, as long as the blood gets there somehow. To wit: occasionally a person does not have a right subclavian artery. This does not mean that their right shoulder and arm receive no blood and wither away; usually it means that one of the segmental arteries branching off the descending aorta–which normally serve the ribs and their associated muscles and other soft tissues–is expanded and elongated to compensate, and looks for all the world like a normal subclavian artery with an abnormal connection to the aorta. But if the major artery that serves the forelimb comes from the descending aorta, and the 4th aortic arch on the right is completely resorbed during development, then there is nothing left on the right side to drag the inferior laryngeal nerve down into the torso. A person with this setup will have an inferior laryngeal nerve on the right that looks intelligently designed, and the usual dumb RLN on the left.

Can people have a nonrecurrent laryngeal nerve on the left? Sure, if they’ve got situs inversus, in which the normal bilateral asymmetry of the internal organs is swapped left to right. Situs inversus is pretty darned rare in the general population, occurring in fewer than 1 in 10,000 people. It is much more prevalent in television shows and movies, where the hero or villain may survive a seemingly mortal wound and then explain that he was born with his heart on the right side. (Pro tip: the heart crosses the midline in folks of both persuasions, so just shoot through the sternum and you’ll be fine. Or, if you’re worried about penetration, remember Rule #2 and put one on either side.) Anyway, take everything I wrote in the preceding paragraph, mirror-image it left to right, and you’ve got a nonrecurrent laryngeal nerve on the left. But just like the normally-sided person who still has an RLN on the left, a person with situs inversus and no remnant 4th aortic arch on the left (double variation alert!) still has an RLN looping around the aorta and ductus arteriosus on the right.

Bottom line: replumb the vessels to your arms, swap your organs around willy-nilly, you just can’t beat the aorta. If you have an aorta, you’ve got at least one RLN; if you don’t have an aorta, you’re dead, and no longer relevant to this discussion.

Nonrecurrent laryngeal nerves–a developmental Hail Mary?

But wait–how do we know that the inferior laryngeal nerve in embryonic sauropods didn’t get rerouted to travel in front of the fourth and sixth aortic arches, so it could be spared the indignity of being dragged into the chest later on?

First of all, such a course would require that the inferior laryngeal nerve take an equally dumb recurrent course in the embryo. Or maybe it should be called a procurrent course. Instead of simply radiating out from the central nervous system to its targets in the sixth pharyngeal arch, the axons that make up the RLN would have to run well forward of their normal course, loop around the fourth and sixth aortic arches from the front, and then run back down to the sixth pharyngeal arch. There is simply no known developmental mechanism that could make this happen.

Even if we postulated some hypothetical incentive that would draw those axons into the forward U-turn, other axons that took a more direct course from the central nervous system would get to the sixth pharyngeal arch first. By the time the forwardly-recurring axons finished their intelligently-routed course and finally arrived at the sixth pharyngeal arch, all of the innervation targets would be taken, and those axons would die off.

Also, at what point in the evolution of long necks would this forwardly-looping course supposedly be called into existence? Ostriches and giraffes have RLNs that take the same recurrent course as those of humans, pigs, and all other tetrapods. The retention of the recurrent course in extant long-necked animals is further evidence that the developmental constraint cannot be broken.

Finally, the idea that a non-recurrent laryngeal nerve would need to evolve in a long-necked animal is based on the perception that long nerve pathways are somehow physiologically taxing or otherwise bad for the animals in which they occur. But almost every tetrapod that has ever lived has had much longer neurons than those in the RLN, and we all get on just fine with them.

In dire extremity

Probably you have seen enough pictures of neurons to know what one looks like: round or star-shaped cell body with lots of short branches (dendrites) and one very long one (the axon), like some cross between an uprooted tree–or better yet, a crinoid–and the Crystalline Entity. When I was growing up, I always imagined these things lined up nose to tail (or, rather, axon to dendrite) all down my spinal cord, arms, and legs, like boxcars in a train. But it ain’t the case. Textbook cartoons of neurons are massively simplified, with stumpy little axons and only a few to a few dozen terminals. In reality, each neuron in your brain is wired up to 7000 other neurons, on average, and you have about a hundred billion neurons in your brain. (Ironically, 100 billion neurons is too many for your 100 billion neurons to visualize, so as a literal proposition, the ancient admonition to “know thyself” is a non-starter.)

Back to the axons. Forget the stumpy little twigs you’ve seen in books and online. Except for the ganglia of your autonomic nervous system (a semi-autonomous neural network that runs your guts), all of the cell bodies of your neurons are located in your central nervous system or in the dorsal root ganglia immediately adjacent to your spinal cord. The nerves that branch out into your arms and legs, across your face and scalp, and into your larynx are not made of daisy chains of neurons. Rather, they are bundles of axons, very long axons that connect muscles, glands, and all kinds of sensory receptors back to the nerve cell bodies in and around your brain and spinal cord.

Indulge me for a second and wiggle your toes. The cell bodies of the motor neurons that caused the toe-wiggling muscles to fire are located in your spinal cord, at the top of your lower back. Those motor neurons got orders transmitted down your spinal cord from your brain, and the signals were carried to the muscles of your feet on axons that are more than half as long as you are tall.

Some of your sensory neurons are even longer. Lift your big toe and then set it down gently, just hard enough to be sure that it’s touching down on the floor or the sole of your shoe, but not hard enough to exert any pressure. When you first felt the pad of your toe touch down, that sensation was carried to your brain by a single neuron (or, rather, by several neurons in parallel) with receptor terminals in the skin of your toe, axon terminals in your brainstem, and a nerve cell body somewhere in the middle (adjacent to your sacrum and just a bit to one side of your butt crack, if you want the gory details). That’s right: you have individual sensory neurons that span the distance from your brainstem to your most distal extremity. And so does every other vertebrate, from hagfish to herons to hippos. Including, presumably, sauropods.

I had you set your toe down gently instead of pushing down hard because the neurons responsible for sensing pressure do not travel all the way from toe-tip to brainstem; they synapse with other neurons in the spinal cord and those signals have been through a two-neuron relay by the time they reach your brainstem. Ditto for sensing temperature. But the neurons responsible for sensing vibration and fine touch go all the way.

If you want to experience everything I’ve discussed in this post in a single action, put your fingertips on your voicebox and hum. You are controlling the hum with signals sent from your brain to your larynx through your recurrent laryngeal nerves, and sensing the vibration through individual neurons that run from your fingertips to your brainstem. Not bad, eh?

Wedel (2012) Fig. 2. The longest cells in the bodies of sauropods were sensory neurons that connected receptors in the skin of the extremities with interneurons in the brainstem, a pattern of neural architecture that is present in all extant vertebrates. The nerve cell bodies would have been located in the dorsal root ganglia adjacent to the spinal cord. The diagram of the neuron is based on Butler and Hodos (1996: fig. 2–1B).

Getting back to big animals: the largest giraffes may have 5-meter neurons in their RLNs, but some of the sensory neurons to their hindfeet must be more like 8 meters long. I don’t think anyone’s ever dissected one out, but blue whales must have sensory neurons to the tips of their flukes that are almost 30 meters (98 feet) long (subtract the length of the skull, but add the lateral distance from body midline to fluke-tip). And Supersaurus, Amphicoelias, and the like must have had neurons that were approximately as long as they were, minus only the distance from the snout-tip to the back of the skull. I could be wrong, and if I am I’d love to be set straight, but I think these must have been the longest cells in the history of life.

Oh, one more thing: up above I said that almost every tetrapod that has ever lived has had much longer neurons than those in the RLN. The exceptions would be animals for which the distance from brainstem to base of neck was longer than the distance from base of neck to tip of limb or tail, so that twice the length of the neck would be longer than the distance from base of skull to most distal extremity. In that case, the neurons that contribute to the RLN would be longer than those running from brainstem to tail-tip or toe-tip. Tanystropheus and some of the elasmosaurs probably qualified; who else?

Parting Thoughts

In this post I’ve tried to explain the courses that these amazingly long cells take in humans and other vertebrates. I haven’t dealt at all with the functional implications of long nerves, for which please see the paper. The upshot is that big extant animals get along just fine with their crazy-long nerves, and there’s no reason to assume that sauropods were any more troubled. So why write the paper, then? Well, it was fun, I learned a lot (dude: axoplasmic streaming!), and most importantly I got to steal a little thunder from those silly poseurs, the giraffes.

Department of Frivolous Nonsense: yes, I titled the paper after those TV ads for Chili’s hamburgers from a few years back. If you never saw them, the ads compared mass-produced, machine-stamped fast-food burgers with restaurant burgers painstakingly built by hand, and concluded with, “Chili’s Big-Mouth Burgers: monuments of inefficiency!”

Update: All of the following is out of date now that the paper has been formally published. Department of Good Karma: since the paper is at the “accepted manuscript” stage, I still have the chance to make (hopefully minor) changes when I get the proofs. As is always, always, always the case, I caught a few dumb errors only after the manuscript had been accepted. Here’s what I’ve got so far, please feel free to add to the list:

  • Page 1, abstract, line 3: pharyngeal, not pharyngial
  • Page 1, abstract, line 8: substitute ‘made up’ for ‘comprised’
  • Page 6, line 12: substitute ‘make up’ for ‘comprise’
  • Page 9, line 5: citation should be of Carpenter (2006:fig. 3), not fig. 2
  • Page 10, line 7: “giant axons of squid are”, not ‘ares’
  • Page 12, entry for Butler and Hodos should have year (1996)
  • Page 12, entry for Carpenter has ‘re-evaluation misspelled
  • Page 16, entry for Woodburne has ‘mammalian’ misspelled

(Notes to self: stop trying to use ‘comprise’, lay off the ‘s’ key when typing bibliography.)

References

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.