Matt’s last post contained a nice overview of the occurrence of epipophyses in sauropodomorphs: that is, bony insertion points for epaxial ligaments and muscles above the postzygapophyseal facets. What we’ve not mentioned so far is that these structures are not limited to sauropods. Back when we were preparing one of the earlier drafts of the paper that eventually became Why sauropods had long necks; and why giraffes have short necks (Taylor and Wedel 2013a), I explored their occurrence in related groups. But that section never got written up for the manuscript, and now seems as good a time as any to fix that.

Theropods (including birds)

Most obviously, epipophyses occur in theropods, the sister group of sauropodomorphs.

Taylor and Wedel (2013a: figure 11). Archosaur cervical vertebrae in posterior view, Showing muscle attachment points in phylogenetic context. Blue arrows indicate epaxial muscles attaching to neural spines, red arrows indicate epaxial muscles attaching to epipophyses, and green arrows indicate hypaxial muscles attaching to cervical ribs. While hypaxial musculature anchors consistently on the cervical ribs, the principle epaxial muscle migrate from the neural spine in crocodilians to the epipophyses in non-avial theropods and modern birds, with either or both sets of muscles being significant in sauropods. 1, fifth cervical vertebra of Alligator mississippiensis, MCZ 81457, traced from 3D scans by Leon Claessens, courtesy of MCZ. Epipophyses are absent. 2, eighth cervical vertebra of Giraffatitan brancai paralectotype HMN SII, traced from Janensch (1950, figures 43 and 46). 3, eleventh cervical vertebra of Camarasaurus supremus, reconstruction within AMNH 5761/X, “cervical series I”, modified from Osborn and Mook (1921, plate LXVII). 4, fifth cervical vertebra of the abelisaurid theropod Majungasaurus crenatissimus,UA 8678, traced from O’Connor (2007, figures 8 and 20). 5, seventh cervical vertebra of a turkey, Meleagris gallopavo, traced from photographs by MPT.

Taylor and Wedel (2013a: figure 11). Archosaur cervical vertebrae in posterior view, Showing muscle attachment points in phylogenetic context. Blue arrows indicate epaxial muscles attaching to neural spines, red arrows indicate epaxial muscles attaching to epipophyses, and green arrows indicate hypaxial muscles attaching to cervical ribs. While hypaxial musculature anchors consistently on the cervical ribs, the principle epaxial muscle migrate from the neural spine in crocodilians to the epipophyses in non-avial theropods and modern birds, with either or both sets of muscles being significant in sauropods. 1, fifth cervical vertebra of Alligator mississippiensis, MCZ 81457, traced from 3D scans by Leon Claessens, courtesy of MCZ. Epipophyses are absent. 2, eighth cervical vertebra of Giraffatitan brancai paralectotype HMN SII, traced from Janensch (1950, figures 43 and 46). 3, eleventh cervical vertebra of Camarasaurus supremus, reconstruction within AMNH 5761/X, “cervical series I”, modified from Osborn and Mook (1921, plate LXVII). 4, fifth cervical vertebra of the abelisaurid theropod Majungasaurus crenatissimus,UA 8678, traced from O’Connor (2007, figures 8 and 20). 5, seventh cervical vertebra of a turkey, Meleagris gallopavo, traced from photographs by MPT.

In this figure from the 2013 paper, the rightmost images show cervical vertebrae of Majungasaurus (an abelisaurid theropod) and a turkey, both in posterior view. The red arrows indicate epaxial musculature pulling on the epipophyses. They are particularly prominent in Majungasaurus, rising almost a full centrum’s height above the postzygapophyseal facets.

The epipophyses are very prominent in the anterior cervicals of Tyrannosaurus, but much less so in its posterior cervicals — presumably because its flesh-tearing moves involved pulling upwards more strongly on the anterior part of the neck. Here’s a photo of the AMNH mount, from our post T. rex‘s neck is pathetic:

amnh-tyrannosaurus-is-pathetic

You can see something similar in the neck of Allosaurus, and the trend generally seems to be widespread among theropods.

Ornithischians

Note the very prominent epipophyses protruding above the postzygs in the anterior cervicals of this Heterodontosaurus in the AMNH public gallery:

Cast of AMNH 28471, Heterodontosaurus tucki, collected from the Early Jurassic Voisana, Herschel district, South Africa. Anterior to the left.

Cast of AMNH 28471, Heterodontosaurus tucki, collected from the Early Jurassic Voisana, Herschel district, South Africa. Neck in left lateral view.

Here’s the hadrosaur Corythosaurus:

AMNH 5338, Corythosaurus casuarius, from the Campanian of the Red Deer River, Alberta, Canada. Collected by Barnum Brown and P. C. Kaisen, 1914. Cervicals 1-4 in right lateral view.

AMNH 5338, Corythosaurus casuarius, from the Campanian of the Red Deer River, Alberta, Canada. Collected by Barnum Brown and P. C. Kaisen, 1914. Cervicals 1-4 in right lateral view.

The prominent vertebra is C2: note that is has both a modest blade-like neural spine and prominent epipophyses — but that already by C3 the epipophyses are gone. Here is that C2 postzyg/epipophyses complex is close-up, clearly showing anteroposteriorly directed striations on the epipophysis, presumably representing the orientation of the attaching ligaments and muscles:

As previous image: close-up of posterior part of C2.

As previous image: close-up of posterior part of C2.

Here’s a close-up of the neck of the boring ornithopod Tenontosaurus, also in the AMNH gallery. (I’m not sure of the specimen number — if anyone can clarify, please leave a comment).

AMNH ?3554, Tenontosaurus tilletti, cervcials 2-4 in right lateral view.

AMNH ?3554, Tenontosaurus tilletti, cervicals 2-4 in right lateral view.

The interesting thing here is that it its axis (C2) seems to lack epipophyses (unlike C3), and to have a tall blade-like neural spine, as seen in mammals. We don’t really see C2 spines this big in other dinosaurs — compare with the much more modest spine in Corythosaurus, above. The texture of this part of the Tenontosaurus specimen looks suspicious, and I wonder whether that neural spine is a fabrication, created back in the day by AMNH staff who were so used to mammals that they “knew” what a C2 should look like? Anyway, the epipophysis above the postzyg of C3 is very distinct and definitely real bone.

Pterosaurs

Things get much more difficult with pterosaurs, because their cervicals are so fragile and easily crushed (like the rest of their skeleton, to be fair). While it’s easy to find nice, well-preserved ornithischian necks on display, you don’t ever really see anything similar for pterosaurs.

As a result, we have to rely on specimen photographs from collections, or more often on interpretive drawings. Even high-resolution photos, such as the one in Frey and Tischlinger (2012: fig 2) tend not to show the kind of detail we need. Usually, the only usable information comes from drawings made by people who have worked on the specimens.

Here, for example, is Rhamphorhynchus, well known as the most difficult pterosaur to spell, in figure 7 from Bonde and Christiansen’s (2003) paper on its axial pneumaticity:

BondeChristiansen2003-axial-pneumaticity-of-rhamphorhynchus-fig7It’s not the main point of the illustration, but you can make out clear epipophyses extending posteriorly past the postzygapophyseal facets in at least C3 and C5 — in C4, the relevant area is obscured by a rib. (Note that the vertebrae are upside down in this illustration, so you need to be looking towards the bottom of the picture.)

I’m pretty sure I’ve seen a better illustration of Rhamphorhynchus epipophyses, but as I get older my memory for Rhamphorhynchus epipophyses is no longer what it used to be and I can’t remember where. Can anyone help?

But also of interest is the azhdarchid pterosaur Phosphatodraco, here illustrated by Pereda Suberbiola et al. (2003):

Pereda Suberbiola et al. (2003: fig. 3). Phosphatodraco mauritanicus gen. et sp. nov, OCP DEK/GE 111, Late Cretaceous (Maastrichtian), Morocco: (a) cervical five in two fragments, ventral and left lateral views; (b) cervical six in ventrolateral view; (c) cervical seven in ventral view; (d) cervical eight in left lateral view; (e) cervical nine in posterior view; (f) cervical six in anterior view. c, centrum; co, condyle; ct, cotyle; hyp, hypapophysis; nc, neural canal; ns, neural spine; poe, postexapophysis; poz, postzygapophysis; prz, prezygapophysis; su, sulcus; tp, transverse process.

Pereda Suberbiola et al. (2003: fig. 3). Phosphatodraco mauritanicus gen. et sp. nov, OCP DEK/GE 111, Late Cretaceous (Maastrichtian), Morocco: (a) cervical five in two fragments, ventral and left lateral views; (b) cervical six in ventrolateral view; (c) cervical seven in ventral view; (d) cervical eight in left lateral view; (e) cervical nine in posterior view; (f) cervical six in anterior view. c, centrum; co, condyle; ct, cotyle; hyp, hypapophysis; nc, neural canal; ns, neural spine; poe, postexapophysis; poz, postzygapophysis; prz, prezygapophysis; su, sulcus; tp, transverse process.

The cervicals of Phosphatodraco seem to have no epipophyses. So they were not ubiquitous in pterosaurs.

What does it all mean? This post has become a bit of a monster already so I’ll save the conclusion for another time. Stay tuned for more hot epipophyseal action!

References

  • Bonde, Niels and Per Christiansen. 2003. The detailed anatomy of Rhamphorhynchus: axial pneumaticity and its implications. pp 217-232 in: E. Buffetaut and J-M Mazin (eds), Evolution and Palaeobiology of Pterosaurs. Geological Society, London, Special Publications 217. doi:10.1144/GSL.SP.2003.217.01.13
  • Frey Eberhard and Helmut Tischlinger. 2012. The Late Jurassic Pterosaur Rhamphorhynchus, a Frequent Victim of the Ganoid Fish Aspidorhynchus? PLoS ONE 7(3):e31945. doi:10.1371/journal.pone.0031945
  • Janensch, Werner. 1950. Die Wirbelsaule von Brachiosaurus brancai. Palaeontographica, Supplement 7 3:27-93.
  • O’Connor Patrick M. 2007. The postcranial axial skeleton of Majungasaurus crenatissimus (Theropoda: Abelisauridae) from the Late Cretaceous of Madagascar. pp 127-162 in: S. D. Sampson., D. W. Krause (eds), Majungasaurus crenatissimus (Theropoda: Abelisauridae) from the Late Cretaceous of Madagascar. Society of Vertebrate Paleontology Memoir 8.
  • Osborn, Henry F., and Charles C. Mook. 1921. Camarasaurus, Amphicoelias and other sauropods of Cope. Memoirs of the American Museum of Natural History, New Series 3:247-387.
  • Pereda Suberbiola, Xabier, Nathalie Bardet, Stéphane Jouve, Mohamed Iarochène, Baadi Bouya and Mbarek Amaghzaz. 2003. A new azhdarchid pterosaur from the Late Cretaceous phosphates of Morocco. pp 79-90 in: E. Buffetaut and J-M Mazin (eds), Evolution and Palaeobiology of Pterosaurs. Geological Society, London, Special Publications 217. doi:10.1144/GSL.SP.2003.217.01.08
  • 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

metaphor

…is not actually about scholarly publication. It’s Steve Albini’s keynote address at Melbourne’s Face the Music conference. It’s about the music industry, and how the internet transformed it from a restrictive, top-down oligarchy that mostly benefited middlemen into a more open, level, vibrant ecosystem where artists can get worldwide exposure for free, and yet are often compensated better than they were under the old system. Go read it, and then think about this:

Once the music world met the internet, the problem of getting information from musicians (authors) to listeners (readers) didn’t require any central planning to solve. What little building needed to happen was taken care of by people who were just happy to let the internet work the way it was designed to, and the way it works the most naturally: it makes sharing information almost effortless. Publishers (record labels) still exist, because they offer certain conveniences, but few people are under the delusion that they are necessary.

dead horse

Over here in academia, we’ve already spent more than a decade wringing our hands over how to manage the shift from a barrier-based publishing world to one based on OA. We’ve put so much time and effort and thought into the problem of how to “save” or “transform” scholarly publishing. Why do we do that? Why not just walk away? Publishing is a button, and anything that we do to lend it any more importance–anything we feed it, in terms of time, effort, energy, or regard–is wasted. Wasted because we deliberately ignore the new reality in favor of propping up a system that performed a job that no-one needs done anymore. I keep wondering when the hell we’re all going to wake up, and start sharing our work the way that musicians and listeners share digital music.

And yet even out here on the crazy-eyed, axe-wielding fringe of the OA movement, we are still conservative. Zen Faulkes published a paper on his blog, and he did it 26 months ago, which is a near eternity in the Shiny Digital Future (it’s 13.4% of the lifespan to date of Google). Mike and I have admired that move, and talk about it, but we haven’t done it. Why not? We could even solicit peer reviews from people we know to be tough but fair reviewers. We all do unpaid editorial and review work for publishers, why not for each other directly? It’s like we’re thinking, “Okay, okay, I’ll review this paper, but only if there’s a publisher somewhere that will benefit from my unpaid labor!”

I suppose that for us, one answer is that PeerJ has given us other options that are just as easy as blogging, like posting preprints. So I am a bit torn: I like PeerJ, I support it, I have several papers in the pipeline that I’m planning on sending there. It offers certain conveniences, like sticking DOIs on everything for us, and tracking all of our metrics. But do we need PeerJ? I wonder if it is just the methadone that will help ease us out of our sad addiction to publishers.

Okay we get it already

Bonus observation: don’t just translate Albini’s thoughts on music to scholarly publishing, also try doing the reverse. It becomes pretty clear that the central theme of The Scholarly Kitchen is, “How will poor, helpless music listeners survive without all the middlemen to tell them what to listen to? They’ll be so lost.” Keep polishing that brass, guys, and thanks for the patronization!

The photos are of the dodo skeleton in the Yorkshire Museum, which I saw at SVPCA back in September. If you’re a dodo-phile like me, you should consider supporting Leon Claessens’s, Kenneth Rijsdijk’s, and Hanneke Meijer’s quest to better understand the skull and feeding mechanics of dodos. Their crowdfunding campaign runs through the end of the year–please go check it out.

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.

 

In the last post I pointed out some similarities between Davide Bonadonna’s new Spinosaurus painting and Brian Engh’s Spinosaurus painting from 2010. I also suggested that Davide might have borrowed from Brian and might have crossed a line in doing so. I was mistaken about that, as this post will show, and I’m sorry. 

I woke up this morning to find that Mike and Davide had a very fruitful and collegial discussion going in email, which they had kindly copied me on. Davide had offered to send his in-progress sketches, Mike had offered to put them up here as a guest post, “because it’ll be a fascinating post — NOT as any kind of defense” (his words, with which I fully agree), and Davide had kindly assented (Brian’s post on how his Spinosaurus came to be is on his own blog). Davide and I corresponded directly this morning and he’s been very gracious and generous with his time, thoughts, and art.

We are always thrilled when we have the opportunity to show how awesome paleoart came into being (like this and this), and this case is no exception. Best now if I just get out of the way, so — over to Davide!

— Matt

——————-

About the illustration:

In early November 2013, I was commissioned by NGMag, via Nizar Ibrahim of the University of Chicago, to create an illustration for a page in the October 2014 issue.

Working for about six years with Simone Maganuco, co-author of the study, on the Spinosaurus (I made the digital model from which the model exhibited in Washington was printed, Nizar left us carte blanche.

Some key points were essential, however: showing the Spinosaurus while swimming, his webbed feet, show its prey in the environment of Kemkem, possibly including all the major players in the scene, Mawsonia, Alanqa and Carcharodontosaurus.

Problems: the Spinosaurus is very long, the subjects to be represented too many. It was decided first of all to exclude the Carcharodontosaurus and then framing a foreshortened Spinosaurus, which would allow us to make room for the actors. Given the size and shape of Spinosaurus we knew that we would inevitably get what I call the “Luis Rey-effect” style. So, after gathering plenty of references, I made my sketches, suggesting a frontal dynamic sight (4) and a back view (1-2-3), presenting both solutions to Nizar at last SVP in L.A.

1

1

2-Spinobozza-1

2

3-SpinoNG_bozza-1

3

4-SpinoNG_bozza-2

4

Meanwhile the size of the final art had to be changed because from the mag they asked for a double opening page of the article. And in the same time, thanks to a friend suggestion, I drew a third version (5), with the Idea to put all them together (8).

5

5

8

8

But the scene was too crowded and we decided to use just two animals, so I tried different combinations (6).

6

6

And the best one was to put both frontal versions together, one close to the other (7).

7

7

And again the two-pages image had to be changed because NG decided to turn it in a three-pages wide illustration, something that helped me to put Mawsonia in the background (9).

9

9

When finished, before approval, the NG editorial staff asked me to put an animal familiar to the modern public, which could help the reader to feel how big was the Spinosaurus, and a turtle was the chosen one (10).

10

10

Brian Engh’s illustration:

I vaguely remember I once had seen Brian’s illustration before today and I did not put it in my archive as a reference. All my main references are these: crocodile photos, patchworks made with my 3D digital model and Dinoraul one (11).

11

11

The water view comes from an NG poster about marine reptiles (12).

12-Spinonuoto_NG2_reference

12

Most of my illustrations have a fisheye distortion, this is not the first one I make (see on my website Scipionyx, Neptunidraco, Diplodocus-Allosaurus and others).

You can easily see from the sketches progress how a traditional vanishing point becomes gradually a curve.

Conclusion:

This is a case of illustrative convergence. ;-)

That’s all folks, I think. If you have any other doubt, just ask. I’m at your disposal.

Best,

Davide

http://www.davidebonadonna.it/

https://www.facebook.com/pages/Davide-Bonadonna/286308368137641?fref=ts

Spinosaurus fishiness, part n

September 15, 2014

UPDATE the next day: Since I published this post, it’s become clear that the similarities in the two images are in fact convergence. Davide Bonadonna got in touch with Mike and me, and he has been very gracious and conciliatory. In fact, he volunteered to let us post the making-of images for his painting, which I will do shortly. I’m sorry that my initial post was more inquisitorial than inquisitive, and implied wrongdoing on Davide’s part. Rather than edit it out of existence, I’m going to let it stand as a cautionary signal to my future self. Stand by for the new post as soon as I can get it assembled and published….aaaand here it is.

——-

Scott Hartman has already explainedtwice–that the super-short-legged, “Ambulocetus-grade” Spinosaurus from the new Ibrahim et al. (2014) paper has some major problems. Those are both good, careful, thought-provoking posts and you should go read them.

I’m writing about something else fishy with the “new” Spinosaurus and, in particular, National Geographic’s media push. Let’s check out this life restoration, newly prepared for the Spinosaurus story:

Spinosaurus - Nat Geo

And now let’s look at this one by Brian Engh from a couple of years ago, borrowed from Brian’s art page:

Spinosaurus KemKem - Brian Engh

And let’s count up the similarities:

  • Two spinosaurs, one in the foreground with its head mostly or entirely submerged as it bites a fish, and one further back on the right with its head complete out of the water;
  • Two turtles, one in the foreground with its head out of the water, and one further back on the right fully submerged;
  • A good diversity of fish swimming around in the foreground;
  • Pterosaurs flying way back in the background;

And finally, and most interestingly to me:

  • A curved-water-surface, fish-eye perspective to the whole scene.

All the bits are moved around a bit, but pretty much everything in Brian’s picture is in the new one. Is it all just a big coincidence–or rather, a fairly lengthy series of coincidences? Seems unlikely. Your thoughts are welcome.

I’m scrambling to get everything done before I leave for England and SVPCA this weekend, so no time for a substantive post. Instead, some goodies from old papers I’ve been reading. Explanations will have to come in the comments, if at all.

Streeter (1904: fig. 3). Compare to the next image down, and note that in birds and other reptiles the spinal cord runs the whole length of the vertebral column, in contrast to the situation in mammals.

Streeter (1904: fig. 3). Compare to the next image down, and note that in birds and other reptiles the spinal cord runs the whole length of the vertebral column, in contrast to the situation in mammals.

Nieuwenhuys (1964: fig. 1)

Nieuwenhuys (1964: fig. 1)

Butler and Hodos (1996: fig. 16.27)

Butler and Hodos (1996: fig. 16.27)

For more noodling about nerves, please see:

References

  • Butler, A.B., and Hodos, W. 1996. Comparative Vertebrate Neuroanatomy: Evolution and Adaptation. 514 pp. Wiley–Liss, New York.
  • Nieuwenhuys, R. (1964). Comparative anatomy of the spinal cord. Progress in Brain Research, 11, 1-57.
  • Streeter, G. L. (1904). The structure of the spinal cord of the ostrich. American Journal of Anatomy, 3(1), 1-27.

 

photo1

I was at the Natural History Museum of Los Angeles County yesterday to do some research in the ornithology collection. After lunch I was working on this pelican skeleton and I thought, “Geez, there is just no way to do this thing justice with still photos. I should make a video.” Here it is. You’ll want to see it full-screen–this being my first time out making a video, I didn’t realize that I was holding the phone the wrong way for efficient viewing on other devices.

The specimen is LACM Ornithology 86262. I’m posting this video with the knowledge and kind permission of the ornithology collection staff.

For previous things in this vein, please see:

If you like it that stuff like this exists, please support your local natural history museum, especially the LACM, which has some really fantastic education and outreach programs.

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