Last night, I submitted a paper for publication — for the first time since April 2013. I’d almost forgotten what it felt like. But, because we’re living in the Shiny Digital Future, you don’t have to wait till it’s been through review and formal publication to read it. I submitted to PeerJ, and at the same time, made it available as a preprint (Taylor 2014).

It’s called “Quantifying the effect of intervertebral cartilage on neutral posture in the necks of sauropod dinosaurs”, and frankly the results are weird. Here’s a taste:

Taylor (2014:figure 3). Effect of adding cartilage to the neutral pose of the neck of Apatosaurus louisae CM 3018. Images of vertebra from Gilmore (1936:plate XXIV). At the bottom, the vertebrae are composed in a horizontal posture. Superimposed, the same vertebrae are shown inclined by the additional extension angles indicated in Table 1. If the slightly sub-horizontal osteological neutral pose of Stevens and Parrish (1999) is correct, then the cartilaginous neutral pose would be correspondingly slightly lower than depicted here, but still much closer to the elevated posture than to horizontal. (Note that the posture shown here would not have been the habitual posture in life: see discussion.)

Taylor (2014:figure 3). Effect of adding cartilage to the neutral pose of the neck of Apatosaurus louisae CM 3018. Images of vertebra from Gilmore (1936:plate XXIV). At the bottom, the vertebrae are composed in a horizontal posture. Superimposed, the same vertebrae are shown inclined by the additional extension angles indicated in Table 1. If the slightly sub-horizontal osteological neutral pose of Stevens and Parrish (1999) is correct, then the cartilaginous neutral pose would be correspondingly slightly lower than depicted here, but still much closer to the elevated posture than to horizontal. (Note that the posture shown here would not have been the habitual posture in life: see discussion.)

A year back, as I was composing a blog-post about our neck-cartilage paper in PLOS ONE (Taylor and Wedel 2013c), I found myself writing down the rather trivial formula for the additional angle of extension at an intervertebral joint once the cartilage is taken into account. In that post, I finished with the promise “I guess that will have to go in a followup now”. Amazingly it’s taken me a year to get that one-pager written and submitted. (Although in the usual way of things, the manuscript ended up being 13 pages long.)

To summarise the main point of the paper: when you insert cartilage of thickness t between two vertebrae whose zygapophyses articulate at height h above the centra, the more anterior vertebra is forced upwards by t/h radians. Our best guess for how much cartilage is between the adjacent vertebrae in an Apatosaurus neck is about 10% of centrum length: the image above shows the effect of inserting that much cartilage at each joint.

And yes, it’s weird. But it’s where the data leads me, so I think it would be dishonest not to publish it.

I’ll be interested to see what the reviewers make of this. You are all of course welcome to leave comments on the preprint itself; but because this is going through conventional peer-review straight away (unlike our Barosaurus preprint), there’s no need to offer the kind of detailed and comprehensive comment that several people did with the previous one. Of course feel free if you wish, but I’m not depending on it.

References

Gilmore Charles W. 1936. Osteology of Apatosaurus, with special reference to specimens in the Carnegie Museum. Memoirs of the Carnegie Museum 11:175–300 and plates XXI–XXXIV.

Stevens, Kent A., and J. Michael Parrish. 1999. Neck posture and feeding habits of two Jurassic sauropod dinosaurs. Science 284(5415):798–800. doi:10.1126/science.284.5415.798

Taylor, Michael P. 2014. Quantifying the effect of intervertebral cartilage on neutral posture in the necks of sauropod dinosaurs. PeerJ PrePrints 2:e588v1 doi:10.7287/peerj.preprints.588v1

Taylor, Michael P., and Mathew J. Wedel. 2013c. The effect of intervertebral cartilage on neutral posture and range of motion in the necks of sauropod dinosaurs. PLOS ONE 8(10):e78214. 17 pages. doi:10.1371/journal.pone.0078214

Just a quick post to link to all five (so far) installments of the “necks lie” series. I need this because I want to cite all the “necks lie” posts in a paper that I’ll shortly submit, and it seems better to cite a single page than four of them.

I’ll update this post as and when we write more about lying necks.

X-ray of the neck of a seal, from Irish Seal Sanctuary. Note that the vertebral column becomes much more vertical than the fleshy envelope suggests.

X-ray of the neck of a seal, from Irish Seal Sanctuary. Note that the vertebral column becomes much more vertical than the fleshy envelope suggests.

 

A couple of times now, I’ve pitched in an abstract for a Masters project looking at neck cartilage, hoping someone at Bristol will work on it with me co-supervising, but so far no-one’s bitten. Here’s how I’ve been describing it:

Understanding posture and motion in the necks of sauropods: the crucial role of cartilage in intervertebral joints

The sauropod dinosaurs were an order of magnitude bigger than any other terrestrial animal. Much sauropod research has concentrated on their long necks, which were crucial to their success (e.g. Sander et al. 2010). One approach to understanding neck function tries to determine neutral posture and range of motion by modelling the cervical vertebrae as a mechanical system (e.g. Stevens and Parrish 1999).

The raw material of such studies is fossilised vertebrae, but these are problematic for several reasons. The invariable incompleteness and distortion of sauropod neck fossils causes fundamental difficulties; but even given perfect fossils, the lack of preserved cartilage means that the bones are not shaped or sized as they were in life.

Ignoring cartilage has dramatic consequences for neutral posture, range of motion and even length of necks: pilot studies (Cobley 2011, Taylor 2011) found that intact bird necks are 8–12% longer than articulated sequences of their dry bones, and that figure is as high as 24% for a juvenile giraffe neck. A turkey neck postzygapophysis was 26% longer when cartilage was included than after being stripped down to naked bone.

We do not yet know how much articular cartilage sauropods had in their necks, nor even what kind of intervertebral joints they had: crocodilians have fibrocartilaginous discs like those of mammals, while birds have synovial joints, so the extant phylogenetic bracket is uninformative.

The project will involve dissection and measurement of bird and crocodilian necks, documenting the extent and shape of articular cartilage, identifying osteological correlates of fibrocartilaginous and synovial joints, and applying this data to sauropods to determine the nature of their neck joints and length of their necks, to reconstruct the lost cartilage, and to determine its effect on neutral pose and range of motion.

Following completion, we anticipate publication of the project.

References

Cobley, Matthew J. 2011. The flexibility and musculature of the ostrich neck: implications for the feeding ecology and reconstruction of the Sauropoda (Dinosauria: Saurischia). MSc Thesis, Department of Earth Sciences, University of Bristol. vi+64 pages.

Sander, P. Martin, Andreas Christian, Marcus Clauss, Regina Fechner, Carole T. Gee, Eva-Maria Griebeler, Hanns-Christian Gunga, Jürgen Hummel, Heinrich Mallison, Steven F. Perry, Holger Preuschoft, Oliver W. M. Rauhut, Kristian Remes, Thomas Tütken, Oliver Wings and Ulrich Witzel. 2010. Biology of the sauropod dinosaurs: the evolution of gigantism. Biological Reviews 86:117–155. doi:10.1111/j.1469-185X.2010.00137.x

Stevens, Kent A., and J. Michael Parrish. 1999. Neck Posture and Feeding Habits of Two Jurassic Sauropod Dinosaurs. Science 284:798–800. doi:10.1126/science.284.5415.798

Taylor, Michael P., and Mathew J. Wedel. 2011. Sauropod necks: how much do we really know?. p. 20 in Richard Forrest (ed.), Abstracts of Presentations, 59th Annual Symposium of Vertebrae Palaeontology and Comparative Anatomy, Lyme Regis, Dorset, UK, September 12th–17th 2011. 37 pp. http://www.miketaylor.org.uk/dino/pubs/svpca2011/TaylorWedel2011-what-do-we-really-know.ppt

(Obviously some part of this have since been covered by my and Matt’s first cartilage paper, but plenty has not.)

I now think there are two reasons no-one’s taken up this project: first, because I wrote it as very focussed only on the question of what type of joint was present, whereas there are plenty of related issues to be investigated along the way; and second, because I wrote it as a quest to discover a specific treasure (an osteological correlate), with the implication that if there’s no treasure to be found then the project will have been a failure.

But I do think there is still plenty of important work to be done in this area, and that there’s lots of important information to be got out of comparative dissection of extant critters.

If anyone out there fancies working in this area, I’d be delighted. I’d also be happy to offer whatever advice and help I could.

Update (18 October 2014)

Somehow I’d forgotten, when I wrote this post, that I’d previously written a more detailed post about the discs-in-sauropod-necks problem. If you’re interested in the problem, you should read that.

Folks,

You may know that the inaugral TetZooCon is set to take place next Saturday (12 July) at the London Wetland Centre. It’s an informal convention that’s condensed around occasional SV-POW!sketeer Darren Naish’s absurdly informative blog Tetrapod Zoology, and features a day of talks, a palaeoart workshop and a quiz. At £40 for the day, it’s a bit of a bargain.

Among the speakers is my own good self, and I will be talking about why giraffes are rubbish.

Taylor and Wedel 2013a: Figure 3. Necks of long-necked sauropods, to scale. Diplodocus, modified from elements in Hatcher (1901, plate 3), represents a “typical” long-necked sauropod, familiar from many mounted skeletons in museums. Puertasaurus, Sauroposeidon, Mamenchisaurus and Supersaurus modified from Scott Hartman’s reconstructions of Futalognkosaurus, Cedarosaurus, Mamenchisaurus and Supersaurus respectively. Alternating pink and blue bars are one meter in width. Inset shows Fig. 1 to the same scale.

Taylor and Wedel 2013a: Figure 3. Necks of long-necked sauropods, to scale. Diplodocus, modified from elements in Hatcher (1901, plate 3), represents a “typical” long-necked sauropod, familiar from many mounted skeletons in museums. Puertasaurus, Sauroposeidon, Mamenchisaurus and Supersaurus modified from Scott Hartman’s reconstructions of Futalognkosaurus, Cedarosaurus, Mamenchisaurus and Supersaurus respectively. Alternating pink and blue bars are one meter in width. Inset shows Fig. 1 to the same scale.

If that sounds like your idea of a good time, then you need to move fast! Booking closes at 4pm this evening. Better get on it now!

 

Emeus crassus mount

In a back room at the Field Museum, from my visit in 2012.

I took a lot of photos of the neck, which nicely records the transition in neural spine shape from simple to bifurcated–a topic of interest to sauropodophiles.

Emeus crassus neural spines

Are you a lover of sauropod necks?

Do you long to demonstrate to your friends and family how much better[1] they are than the necks of other long-necked critters?

Are you crazy for the Taylor and Wedel (2013a) paper on why sauropods had long necks; and why giraffes have short necks, but disappointed that it’s not, until now, been obtainable in T-shirt form?

front

back

If so, it’s your lucky day! You can now buy a T-shirt featuring Figure 1 on the front (necks of a human, giraffe, ostrich, Paraceratherium[2], Therizinosaurus, Gigantoraptor, Arambourgiania and Tanystropheus) and Figure 3 on the back (necks of Diplodocus, Puertasaurus, Sauroposeidon, Mamenchisaurus and Supersaurus).

And here it is in real life — sorry I couldn’t get a more photogenic model at short notice.

DSCN0800-front

DSCN0796-back

And here are the original figures as they appeared in the paper. The full captions, as reproduced here, are also on the shirts — just in case you need to check details while you’re out and about.

Figure 1. Necks of long-necked non-sauropods, to scale. The giraffe and Paraceratherium are the longest necked mammals; the ostrich is the longest necked extant bird; Therizinosaurus and Gigantoraptor are the largest representatives of two long-necked theropod clades; Arambourgiania is the longest necked pterosaur; and Tanystropheus has a uniquely long neck relative to torso length. Human head modified from Gray’s Anatomy (1918 edition, fig. 602). Giraffe modified from photograph by Kevin Ryder (CC BY, http://flic.kr/p/cRvCcQ). Ostrich modified from photograph by “kei51” (CC BY, http://flic.kr/p/cowoYW). Paraceratherium modified from Osborn (1923, figure 1). Therizinosaurus modified from Nothronychus reconstruction by Scott Hartman. Gigantoraptor modified from Heyuannia reconstruction by Scott Hartman. Arambourgiania modified from Zhejiangopterus reconstruction by Witton & Naish (2008, figure 1). Tanystropheus modified from reconstruction by David Peters. Alternating blue and pink bars are 1 m tall.

Figure 1. Necks of long-necked non-sauropods, to scale. The giraffe and Paraceratherium are the longest necked mammals; the ostrich is the longest necked extant bird; Therizinosaurus and Gigantoraptor are the largest representatives of two long-necked theropod clades; Arambourgiania is the longest necked pterosaur; and Tanystropheus has a uniquely long neck relative to torso length. Human head modified from Gray’s Anatomy (1918 edition, fig. 602). Giraffe modified from photograph by Kevin Ryder (CC BY, http://flic.kr/p/cRvCcQ). Ostrich modified from photograph by “kei51” (CC BY, http://flic.kr/p/cowoYW). Paraceratherium modified from Osborn (1923, figure 1). Therizinosaurus modified from Nothronychus reconstruction by Scott Hartman. Gigantoraptor modified from Heyuannia reconstruction by Scott Hartman. Arambourgiania modified from Zhejiangopterus reconstruction by Witton & Naish (2008, figure 1). Tanystropheus modified from reconstruction by David Peters. Alternating blue and pink bars are 1 m tall.

x

Figure 3. Necks of long-necked sauropods, to scale. Diplodocus, modified from elements in Hatcher (1901, plate 3), represents a “typical” long-necked sauropod, familiar from many mounted skeletons in museums. Puertasaurus, Sauroposeidon, Mamenchisaurus and Supersaurus modified from Scott Hartman’s reconstructions of Futalognkosaurus, Cedarosaurus, Mamenchisaurus and Supersaurus respectively. Alternating pink and blue bars are one meter in width. Inset shows Fig. 1 to the same scale.

No doubt these will be all the rage at SVPCA this year!

So get your T-shirts!

Update (the same evening)

As suggested by Kevin, I’ve now made the shirt available in a selection of eight versions: four men’s shirt, two women’s, and two kids. I don’t really understand what the differences are between them all, but they seemed to be the saner choices among those offered by Cafe Press. You can get any or all of them here. The shirt modelled above is the one called simple “White T-Shirt”. Please be aware that unlike all the others, the “Value T-Shirt” has no printing on the back — only Figure 1 on the front.

Notes

[1] i.e. bigger.

[2] Not to be confused with Paramecium.

References

Taylor, Michael P., and Mathew J. Wedel. 2013. Why sauropods had long necks; and why giraffes have short necks. PeerJ 1:e36. doi:10.7717/peerj.36

Bird vertebra diagrams

January 10, 2014

bird neck note sheet

I made these back in the day. The idea was that you could print them out and have them along while dissecting bird necks, so you could draw on the muscles.

bird neck note sheet - LEFT - all three views

It’s basically one drawing of an ostrich vertebra, morphed in GIMP and stacked to simulate articulation. All of the ones in this post show the vertebrae in left lateral view. If you need right views, flip ‘em in GIMP or heck, I think even Windows Explorer will do that for you. The one above has dorsal views in the top row, lateral view in the middle row, and ventral views in the bottom row.

bird neck note sheet - LEFT - double lateral

Here’s a sheet with two rows in lateral view, the idea being that you draw on the more superficial multi-segment muscles on one row, and the deeper single- or two-segment muscles on the other row.

bird neck note sheet - LEFT - 12 cervicals

A version with 12 vertebrae, so you can map out the often complicated patterns of origins and insertions in the really long muscles. How complicated? Well, check out this rhea neck with the M. longus colli dorsalis and M. longus colli ventralis fanned out.

Rhea neck muscles fanned - full

That’s all. Have fun!

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