I’m pleased to announce that Darren has a new paper out (Naish and Sweetman 2011) in which he and fellow Portsmouth researcher Steve Sweetman describe a maniraptoran theropod from the Wealden Supergroup of southern England.  It’s represented only by a single cervical vertebra:

Indeterminate maniraptoran theropod BEXHM 2008.14.1, posterior cervical vertebra, in right lateral view. Sauroposeidon cervical vertebra 8 for scale.

This vertebra is described in seven and a bit pages, which means that it’s had nearly three times as much total coverage as Jobaria (Cf. Sereno et al. 1999).

Still, we can hope that Darren and Steve will return to their specimen some time and monograph it properly.

In the mean time, read all about it over on Tetrapod Zoology.

References

  • Naish, Darren, and Steven C. Sweetman.  2011.  A tiny maniraptoran dinosaur in the Lower Cretaceous Hastings Group: evidence from a new vertebrate-bearing locality in south-east England.  Cretaceous Research 32:464:471.  doi:10.1016/j.cretres.2011.03.001
  • Sereno, Paul C., Allison L. Beck, Didier. B. Dutheil, Hans C. E. Larsson, Gabrielle. H. Lyon, Bourahima Moussa, Rudyard W. Sadleir, Christian A. Sidor, David J. Varricchio, Gregory P. Wilson and Jeffrey A. Wilson.  1999.  Cretaceous Sauropods from the Sahara and the Uneven Rate of Skeletal Evolution Among Dinosaurs.  Science 282:1342-1347.

Meanwhile, elsewhere on the Internet …

On Tuesday morning, a rather nice article about our recent sauropod-necks-were-not-sexually-selected paper appeared on the BBC web-site.  At the time of writing, it’s just topped 100 comments (athough fifteen of those are by me — I wanted to respond to the questions that people were asking).

Here it is, for those who are interested (maybe more in the Q-and-A’s than in the actual article): Evolution, sex and dinosaur necks

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 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 this 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

In our recent paper on how the long necks of sauropods did not evolve primarily due to sexual selection (Taylor et al. 2011), one of the ideas we discussed is that sexual dimorphism between the necks of male and female sauropods would be an indicator of sexual selection.  And, rather despairingly, we wrote (page 4):

As Senter himself recognized, available samples of sauropod taxa are unfortunately not large enough to demonstrate bimodal distribution of morphological features within any sauropod species.

But I wonder if we realise just how true this is, and how blind we are flying?  How very far short we are of being able to do any kind of statistical analysis on sauropod necks.

How many complete necks of a given sauropod would we need in order to demonstrate a bimodal distribution of, say, length?  (That is, to show that the necks mostly fall into two separate buckets, a short-necked group and a long-necked group of which one is presumably male and the other female.)  I don’t know enough about stats, but this article at least suggests that you’d need thirty or so before you could be confident that you were seeing something statistically significant.

And how many sauropod species do we have thirty complete necks for?

Correct: none.

All right, then how many do we have ten complete necks for?

Five complete necks?

OK, how about just two necks?

ONE neck?

The answer is: not many species.  Off the top of my head, I think complete necks are known for Camarasaurus lentus (Gilmore 1925, one specimen), Mamenchisaurus hochuanensis (Young and Zhao 1972, one specimen), Shunosaurus lii (e.g. Zhang et al. 1984; probably multiple specimens but the paper is in Chinese so I don’t know for sure) Mamenchisaurus youngi (Ouyang and Ye 2002, one specimen, I think), and Spinophorosaurus nigerensis (Remes et al. 2009, one specimen).

No doubt I have missed some, but the point is that the total number of sauropods for which even one complete neck is known is a tiny, tiny proportion of all the sauropods that have been named.  I have listed five species here, and of those only one is known from more than a single complete neck.  And those multiple specimens have not been described (have they?)  So while in theory it might be possible to determine whether there is a bimodal distribution in the length of Shunosaurus lii necks, the data doesn’t exist to do this work.  (If there really are enough complete necks then someone ought to get out to China and measure those babies.)

So anyway.  We have very, very few complete sauropod necks.

Diplodocus carnegii

“But Mike!”, I hear you cry; “What about Diplodocus carnegii?  We’ve all seen its skeleton in a half-dozen different museums!”

Oh yes.  Here is its “complete” neck, from Hatcher (1901:plate 8):

Let’s, for now, ignore the fact that the scapula seems to articulate with the base of the neck rather than the torso.  We can all see that there are fifteen cervical vertebrae, right?

Right?

Well, let’s see what Uncle J. Bell had to say (Hatcher 1901:4):

[Diplodocus carnegii holotype CM 84] has been entirely freed from the matrix and is found to consist of the right femur and pelvis complete except for the left ilium, which is for the most part wanting, right scapula and coracoid, two sternals, eighteen ribs and forty-one vertebrae divided as follows: fourteen cervicals including the axis, eleven dorsals, four sacrals, and twelve caudals.  These vertebrae are for the most part fairly complete, though unfortunately the sacrals and anterior cervicals are more or less injured.  This series of forty-one vertebrae are believed to pertain to one individual and to form an unbroken series from the axis to the twelfth caudal, although as was shown in a previous paper, there is some evidence that there are perhaps one or more interruptions in the series and that one or more vertebrae are missing.  On the other hand, as will appear later, it is not entirely impossible that at least one vertebra of this supposed series pertains to a second individual belonging perhaps to a distinct genus.

Oh and there’s this, from page 10:

Unfortunately no diagram of the quarry was made, at the time of exhuming the remains, showing the relative position of each of the several vertebrae and other bones as they lay in the rock.  [Plate 1 is a map of the quarry as remembered by W. H. Reed.]

Hey!  That’s not what it said in the brochure!  So, as it turns out, our conclusion is: Diplodocus carnegii had fifteen cervicals, or more, or maybe less.

Giraffatitan brancai

“Well, then, Mike, how about that awesome mounted Giraffatitan skeleton in the Berlin museum?”

Well, the presacral vertebrae of that mount are not real bone, nor even casts, but they are very good sculptures based on real bones.  However, the real bones that they’re based on are those of two specimens — the lectotype SI and paralectotype SII.  The former includes cervicals 2-7, and we can be confident about that because C2 in sauropods is very distinctive, having a completely different anterior articular surface from all the other cervicals; and the latter includes cervicals 3-13 (although many of them are damaged).

But but but.  SI and SII were smushed up and mixed in together, with little articulation.  Any reconstruction — or even assignment of individual vertebrae to one specimen or the other — has to be considered provisional.  Take a look at this quarry map, from Heinrich (1999:fig. 16):

Yeesh, what a mess.  I’ve previously suggested (Taylor 2009:800-801) that the distinctively high-spined dorsal vertebra usually considered the fourth of SII may not actually belong to that specimen, or even that taxon — that it may belong to a more Archbishop-like animal (which may be what SI is).  Janensch (1950:33) says that things are not so bad for the cervical vertebrae, but still not good:

The vertebrae from the 3rd to 15th presacrals [of SII] lay in articulation in a consolidated lime sandstone lens; of them, the 3rd to 5th vertebrae are tolerably complete, the remaining 10 vertebrae were articulated with one another, with one interruption that arose when the 8th presacral vertebra rotated out of the series and was displaced.

So might there have been other displaced cervicals, before and/or after the “8th”, that were not recovered?  And can we be confident that the anteriormost cervical of SII really is C3?  Why?  Because of the overlap with vertebrae of SI?  But we’re not even certain that SI is the same species as SII.  Maybe the anteriormost preserved cervical is really C4?  Maybe some of the “SII” cervicals really belong to SI?

So all in all, our conclusion is: Giraffatitan brancai had thirteen cervicals, or more, or maybe less.

What does it all mean?

Only this: we don’t know as much as we think we do.  We don’t know how many cervical vertebrae Diplodocus and Giraffatitan had, even.  We don’t have complete necks for either of these sauropods, nor for almost any others.  Even those we do have are in some cases badly crushed (e.g. Mamenchisaurus hochuanensis, which I must post about properly some time).  To summarise: we are woefully short of sauropod necks.

We need to get out of the habit of blithely asserting, “oh, Diplodocus had 15 cervicals and Giraffatitan only 13″.  Because we really don’t know this.  We think it’s true: these numbers are certainly the best guesses for the taxa in question.  But they are, in the end, only guesses.

References

Thanks to everyone who joined in the discussion last time on why sauropods had such long necks.  I’ve discussed this a little with Matt, and we are both amazed that so many different hypotheses have been advanced (even if some of them are tongue-in-cheek).  We’ll probably come back to all these ideas later.

But today, we want to draw your attention to a new contribution to this discussion — a paper in the Journal of Zoology, with the tell-it-like-it-is title “The long necks of sauropods did not evolve primarily through sexual selection”, written by the three of us SV-POW!er rangers together with our buddy Dave “Archosaur Musings” Hone (Taylor et al. 2011).

Taylor et al. (2011), fig. 1: Sauropod necks, showing relationships for a selection of species, and the range of necks lengths and morphologies that they encompass. Phylogeny based on that of Upchurch et al. (2004: fig. 13.18). Mamenchisaurus hochuanensis (neck 9.5 m long) modified from Young & Zhao (1972: fig. 4); Dicraeosaurus hansemanni (2.7 m) modified from Janensch (1936: plate XVI); Diplodocus carnegii (6.5 m) modified from Hatcher (1903: plate VI); Apatosaurus louisae (6 m) modified from Lovelace, Hartman & Wahl (2008: fig. 7); Camarasaurus supremus (5.25 m) modified from Osborn & Mook (1921: plate 84); Giraffatitan brancai (8.75 m) modified from Janensch (1950: plate VIII); giraffe (1.8 m) modified from Lydekker (1894:332). Alternating grey and white vertical bars mark 1 m increments.

This is one of those papers that has been literally years in the making, which is why it’s a rather belated response to the paper that we were responding to — Phil Senter’s (2006) argument that sexual selection was the primary driver of neck elongation in sauropods.

Senter supported his hypothesis by laying out six predictions which he argued should be true for sexually selected necks; then showing that, while the first two could not be assessed, the last four all supported sexual selection.  In our paper, we do three things.  First, we make the point that sexual selection and feeding advantage are not mutually exclusive.  Second, we revisit all six predictions and show that they do not in fact support sexual selection — in fact, most of them provide support for feeding advantage.  Finally, we show that no tetrapod clade comparable with Sauropoda has consistently selected for a single sexual signal.

My email records show that Darren, Matt and I were discussing this as early as 22 September 2006, just six weeks after Senter’s paper was published, and that we started working on a response only a couple of days later.  But as so often happens, it got crowded out by a hundred other things.  Then in November 2007 Dave Hone mentioned that he was independently thinking of writing a response, and we decided to join forces.  And then … we all went back to working on other things again, touching on the necks-for-sex issue every now and then.  It’s mostly due to Dave’s repeated prods that this project wasn’t allowed to wither away, and has now, finally, made it across the finish line.

Like the neck-posture paper (Taylor et al. 2009), this was a true collaboration — one of those where, for many parts of the text, none of us is sure which of us originally wrote it.  It went through the wringer many times before reaching its final form, and most of the text must have been rewritten two or three times along the way.  We hope all the shuffling and polishing has resulted in a paper that reads straightforwardly and even seems obvious.  “When something can be read without effort, great effort has gone into its writing” — Enrique Jardiel Poncela.  That’s the goal, anyway.

The paper itself is available at the link below, so take a look and see whether you find our argument convincing.  As always, comments are open!

Update (the next morning)

Co-author Dave Hone discusses this paper on his own blog.

References

Why did sauropods have such long necks?

Mamenchisaus hochuanensis skeletal reconstruction (Young and Zhao 1972:fig. 4), based on the holotype

It’s the single most obvious and important question about sauropods, so it’s a bit surprising to think that we’ve never really addressed this question directly.

Maybe sauropod necks are so obvious and familiar that we just take them for granted, and move straight on to questions of how they were able to grow so long and remain workable.

Well, let’s fix that.  Let’s think about why they had such long necks.  What were they for?  What were sauropods doing with their necks that was valuable enough to justify all that investment?

Back in the good old days, everyone assumed that sauropod necks were all about high browsing.  If you have a 9.5m neck, then of course you will use it to browse high up in trees — it’s intuitively obvious.  But of course “intuitively obvious” is not the same thing as “true”.

Then John Martin (1987) proposed that the long necks were used for low browsing — not raised above shoulder level, but swept back and forth to allow food to be gathered across a wide area without all that tedious mucking about with locomotion.  This interpretation was of course endorsed by Stevens and Parrish (1999) in their DinoMorph work.

There has been plenty written about habitual sauropod posture — including by us (Taylor et al.2009).  But actually the high-browsing and low-browsing explanations of sauropod neck elongation have much in common.  Most crucially, they both relate to enlarging the feeding envelope; more broadly they are both explanations that rely on the neck having a survival benefit.  But Senter (2006) proposed a completely different explanation — that sauropod necks were sexual signals, selected not for survival advantage but for reproductive success.  The idea is that female sauropods, being very shallow, would go for the males with the biggest protuberances.

Are there other candidate explanations that I’ve missed?

Or is it between high browsing, low browsing and sexual selection?

Comments are open!

References

People who’ve been paying especially close attention may have noted than on four separate occasions in the last eighteen months, I’ve casually referred to our old buddy HMN SII as the paralectotype specimen of Giraffatitan brancai.  (Butchering a wallaby, photographing big bones, How fat was Camarasaurus, and baby giraffe neck, in case you were wondering.)

Giraffatitan brancai paralectotype HMN SII in the justly underrated left posteroventrolateral view, slightly obscured by a bit of Boring Old Diplodocus neck

But in my Big Brachiosaur Bonanza (Taylor 2009:788), I nominated HMN SII as the lectotype of this species.  So why all this paralectotype stuff?  Well, what I wrote in the paper was:

The original type specimen, “Skelett S” (Janensch, 1914:86) was subsequently found (e.g., Janensch, 1929:8) to consist of two individuals, which were designated SI (the smaller) and SII (the larger and more complete). Janensch never explicitly designated these two specimens as a syntype series or nominated either specimen as a lectotype; I therefore propose HMN SII as the lectotype specimen of Brachiosaurus brancai.

But in May last year, I got an email from Mark Konings, a dinosaur enthusiast from the Netherlands, pointing out (more politely than I deserved) that I’d got this wrong.  In fact, Janensch did nominate a lectotype — the wrong one, SI, but we’re stuck with it.  He did this in a paper on skulls (Janensch 1935-1936:151), which is why I overlooked it.  (Well, that and the fact that he rather inconsiderately wrote in German.)

Once I’d been shown my mistake, I realised that the only thing to do was formally correct it in JVP, where the original article had been, so I sent them the shortest and most boring manuscript I’ve ever written (and it is up against some pretty stiff competition in the “most boring” category).  And that manuscript was published today (Taylor 2011), fixing my dumb mistake.

Many thanks to Mark for spotting this!

References

It’s been a couple of months since Brontomerus came out, but new coverage continues to trickle in. For anyone who’s still following, I thought I’d draw attention to a few that I particularly like.

A favourite is One Hip Dino in The Scientist.  It’s told largely from Matt’s perspective, and includes quotes by Mike D’Emic, Susie Maidment and Ray Wilhite.  (Although D’Emic’s statement that “The ilium projects forward by 55 percent, while in other species it’s 52 percent” could do with some substantiation — I think we’ve shown pretty convincingly how different the ilium is from anything else out there.)

The most recent of the new articles is The biggest, baddest dinos still rule, in Macleans, which describes itself as “Canada’s only national weekly current affairs magazine”.  I guess that makes it Canada’s Time or Newsweek, and it has 2.4 million readers.  Despite the rather unpromising title, the article is good, and touches on some of the potential downsides of palaeo publicity.

But one of the best things about publicising Brontomerus has been hearing about how it’s been used in education.  (As one example, it was the lever that got me an opportunity to give a talk about palaeontology and evolution at my eldest son’s school a few weeks ago.)  One article describing Brontomerus‘s involvement in engaging kids’ interest is Dinosaur teaching topics – how to name a dinosaur at Everything Dinosaur.  The author tells me “we chose Brontomerus as the focus for our teaching session and I introduced concepts such as ontogeny and used the children’s knowledge of how farmyard animals grow and change, relating this to the fossil evidence of the adult and juvenile of the Brontomerus genus.”

Another benefit of letting the world know about Brontomerus was that it opened the door to my writing an article for the Guardian‘s science blog: How I got to know thunder thighs, the dinosaur with a fearsome kick.  They chose the title, sadly: I’d suggested something more like “How we know what we know”, and that is indeed that main topic of the article.  It was a rare opportunity to talk in a mainstream media outlet about how we actually do palaeontology, and the varying levels of certainty in which we hold different conclusions.

I hesitate to mention it, but the New York Times did a piece on, well, mostly me: Dinosaur-hunting hobbyist makes fresh tracks for paleontology.  I’m mostly really happy with it, except that an unfortunate bit of abridgement gives the impression that I described Jack McIntosh as “a minor paleontologist”.  Let the record show, that is not what I said: it’s actually how I described myself.

Finally, I’d like to draw attention to a very cheerful interview that Australian science blogger Bec Crew did for ABC Radio’s Triple J channel, in a program called The Doctor on 8th March.  Bec is best known for her truly unique blog Save your breath for running ponies, (I can’t help inserting the missing comma in the title), and my only regret regarding Brontomerus is that it’s never been given the SYBFRP treatment.

That’s all for now.

Atacamatitan chilensis gen. et sp. nov., caudal centrum SGO.PV.961c in ventral (A) and ventrolateral views (B); caudal vertebrae SGO-PV-961h in lateral (C) and dorsal (D) views. Scale bars: 50 mm. (Kellner et al. 2011:fig. 2)

Although we like to stay sauropod-o-centric on SV-POW!, I just want to take a moment to acknowledge the most astounding publication I have ever seen, Sterling Nesbitt’s new basal archosaur phylogeny (Nesbitt 2011).  Thanks to the wonder of open access publishing, it is freely available, and I urge everyone to check it out, if only to gaze in open-mouthed astonishment at the scale of the thing.

In 292 packed pages, Nesbitt provides a new phylogenetic analysis of basal archosaurs, using 80 species and 412 characters.  But if that doesn’t sound like the hugest matrix you’ve ever heard of, what sets this contribution apart is the incredibly detailed work in describing and illustrating those characters.  In those terms, I can only compare it with Wilson and Sereno’s (1988) JVP monograph — but that described 109 characters, and even then not in such exhaustive detail as in the new work.  And everything else about this paper is also super-comprehensive: the discussion of earlier work, the description of the mechanics of the analysis, the extensive sections talking through the expected and unexpected results of that analysis.  To give just a tiny flavour, here’s a figure showing a bunch of basal archosaur braincases:

Braincases of basal archosaurs in lateral view (Nesbitt 2011:fig. 23)

Knowing nothing about basal archosaurs myself, I have nothing intelligent to say about the content of the paper — I will leave that to others, and I don’t doubt that Bill and Jeff will have plenty to say on their respective blogs.  I just want to marvel at the sheer scale of the undertaking.  My Ph.D dissertation was 285 pages long — by coincidence, almost exactly the same length of Nesbitt’s epic.  But dissertations are much less dense than papers: they are double-spaced (or 1.5x spaced in my case, since that was an option and I hate wide spacing with a passion), and figures each take up a whole page — or even two if the caption is separate.  All in all, I’d say that two pages of dissertation are worth one page of publication, near enough.  Which means that Nesbitt has poured twice as much work into a single paper as most of us do into our entire Ph.Ds.

Dude, pls.  You’re making the rest of us look bad.

(By the way, since a decent dissertation contains four or five non-trivial papers, it follows that there’s enough work in the new Nesbitt tome to have been equivalent to maybe ten papers.  but because it’s all in one package, he’ll only get 1/10 as many citations as he would have, had he written ten papers instead.  This just shows what a stupid way counting citations is for assessing the importance of someone’s work.)

The final thing that should be said about this is that by all accounts, Nesbitt is an uncommonly nice guy.  (I’ve only met him once myself, briefly, which is why I don’t feel justified in using his first name in this article.)  And I have found, almost without exception, that the most impressive palaeontologists are also the ones who are most helpful and generous.  I could mention Randy Irmis, for example, who seems to churn out half a dozen top-class papers for every publication I manage to get out the door, and who would be terrifying to be around if he wasn’t such a good guy.  Steve Brusatte is another one whose rate and quality of work is astonishing, yet who is always ready to help out other people.  (I am going to stop mentioning people by name now, otherwise those who don’t get a mention might feel slighted.  There are plenty of other examples, and you probably know who some of them are.)  I don’t know why it should be that quality × quantity of work correlates so well with niceness, but that’s how it seems to be, and I like it that way.

Anyway, go and look at — I won’t say read, not all the way though — Nesbitt’s giant analysis.  It sets the bar higher for us all.

References

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