If you check out the Shiny Digital Future page on this site, where we write about scholarly publishing, open access, open data and other such matters, you will see the following:

  • 2009: 9 posts
  • 2010: 5 posts
  • 2011: 9 posts
  • 2012: 116 posts! Woah!
  • 2013: 75 posts
  • 2014: 34 posts
  • 2015: 31 posts
  • 2016, up until the end of June: 34 posts
  • 2016, July onwards: 8 posts
  • 2017: 12 posts
  • 2018: 6 posts
  • 2019: 4 posts
  • 2020: nothing yet.

In four and a half years up to the end of June, Matt and I (but mostly I) posted 290 times in the Shiny Digital Future, for an average of 64.4 posts a year (one every 5.6 days). Since then we’ve posted 30 times in a bit more than three and a half years, for an average of 8.6 posts a year (one every 42.6 days).

Shiny Digital Future posts by year (2016 split into halves)

Something happened half way through 2016 that cut my Shiny Digital Future productivity to 13% of what it was before. (And, no, I wasn’t bought off by Elsevier.)

Here’s another funny thing. My eldest son was taking his A-levels in the summer of 2016. He had got so good at the Core 4 paper in maths that he was reliably scoring 95–100% on every past paper. He took the actual exam on the morning of 24th June, and scored 65% — a mark so low that it prevented him getting an A* grade.

Well, we all know what happened on the 23rd of June 2016: the Brexit referendum. I know that opinions differ on the desirability of Brexit, but for our family it was emotionally devastating. It’s the reason Dan was so knocked sideways that he botched his Core 4 paper. It’s hung over us all to a greater or lesser extent ever since, and it’s only with the recent triumph of the “Conservative” Party1 in the 2019 General Election that I’ve finally attained the ability to think of it as Somebody Else’s Problem. There is something gloriously liberating about being so comprehensively beaten that you can just give up.

I’m not going to rehearse all the reasons why Brexit is awful — not now, not ever again. (If you have a taste for that kind of thing, I recommend Chris Grey’s Brexit Blog, which is dispassionate, informed and forensic.) I’m not going to follow Brexit commentators on Twitter, and read all the desperately depressing analysis they highlight. I’m certainly not going to blog about it myself any more. More importantly, I’m not going to let the ongoing disintegration of my country dominate my mind or my emotions. I’m walking away: because obviously absolutely nothing I say or do about it can make the slightest bit of difference.

But there is an area of policy where I can hope to make some small difference, and that is of course open science — including but not limited to open access, open data, open reviewing and how research is evaluated. That’s where my political energy should have been going for the last three years, and it’s where that energy will be going from now on.

Because so much is happening in this space right now, and we need to be thinking about it and writing about it! Ludicrously, we’ve never even written anything about Plan S even though it’s nearly eighteen months old. But so much more is going on:

Each of these developments merits its own post and discussion, and I’m sorry I don’t have the energy to do that right now.

What I offer instead is an apology for letting my energy by stolen for so long by such a stupid issue; and a promise to refocus in 2020. I’ll start shortly by writing up the R2R debate that I was involved in on Monday, on the proposition “The venue of its publication tells us nothing useful about the quality of a paper”.

 


1The more right-wing of the two large political parties in the UK is called the Conservative party, and traditionally it has adhered to small-c conservative ideals. But at the moment, it’s the exact opposite of what it says on the tin: it’s been hijacked by a radical movement that, contra Chesterton’s Fence, wants to smash everything up in the hope that whatever emerges from the chaos will be better than what we have now. It may be exciting; it may even (who knows?) prove to be right, in the end2. What it ain’t, is conversative.

2Spoiler: it won’t.

 

In the last post, we looked at some sauropod vertebrae exposed in cross-section at our field sites in the Salt Wash member of the Morrison Formation. This time, we’re going to do it again! Let’s look at another of my faves from the field, with Thuat Tran’s hand for scale. And, er, a scale bar for scale:

And let’s pull the interesting bits out of the background:

Now, confession time. When I first saw this specimen, I interpeted it as-is, right-side up. The round thing in the middle with the honeycomb of internal spaces is obviously the condyle of a vertebra, and the bits sticking out above and below on the sides frame a cervical rib loop. I figured the rounded bit at the upper right was the ramus of bone heading for the prezyg, curved over as I’ve seen it in some taxa, including the YPM Barosaurus. And the two bits below the centrum would then be the cervical ribs. And with such big cervical rib loops and massive, low-hanging cervical ribs, it had to an apatosaurine, either Apatosaurus or Brontosaurus.

Then I got my own personal Cope-getting-Elasmosaurus-backwards moment, courtesy of my friend and fellow field adventurer, Brian Engh, who proposed this:

Gotta say, this makes a lot more sense. For one, the cervical ribs would be lateral to the prezygs, just as in, oh, pretty much all sauropods. And the oddly flat inward-tilted surfaces on what are now the more dorsal bones makes sense: they’re either prezyg facets, or the flat parts of the rami right behind the prezyg facets. The missing thing on what is now the right even makes sense: it’s the other cervical rib, still buried in a projecting bit of sandstone. That made no sense with the vert the other way ’round, because prezygs always stick out farther in front than do the cervical ribs. And we know that we’re looking at the vert from the front, otherwise the backwards-projecting cervical rib would be sticking through that lump of sandstone, coming out of the plane of the photo toward us.

Here’s what I now think is going on:

I’m still convinced that the bits of bone on what is now the left side of the image are framing a cervical rib loop. And as we discussed in the last post, the only Morrison sauropods with such widely-set cervical ribs are Camarasaurus and the apatosaurines. So what makes this an apatosaurine rather than a camarasaur? I find several persuasive clues:

  • If we have this thing the right way up, those prezygs are waaay up above the condyle, at a proportional distance I’ve only seen in diplodocids. See, for example, this famous cervical from CM 3018, the holotype of A. louisae (link).
  • The complexity of the pneumatic honeycombing inside the condyle is a much better fit for an apatosaurine than for Camarasaurus–I’ve never seen that level of complexity in a camarasaur vert.
  • The bump on what we’re now interpreting as the cervical rib looks suspiciously like one of the ventrolateral processes that Kent Sanders and I identified in apatosaurine cervicals back in our 2002 paper. I’ve never seen them, or seen them reported, in Camarasaurus–and I’ve been looking.
  • Crucially, the zygs are not set very far forward of the cervical ribs. By some rare chance, this is pretty darned close to a pure transverse cut, and the prezygs, condyle (at its posterior extent, anyway), and the one visible cervical rib are all in roughly the same plane. In Camarasaurus, the zygs strongly overhang the front end of the centrum in the cervicals (see this and this).

But wait–aren’t the cervical ribs awfully high for this to be an apatosaurine? We-ell, not necessarily. This isn’t a very big vert; max centrum width here is 175mm, only about a third the diameter of a mid-cervical from something like CM 3018. So possibly this is from the front of the neck, around the C3 or C4 position, where the cervical ribs are wide but not yet very deep. You can see something similar in this C2-C5 series on display at BYU:

Or, maybe it’s just one of the weird apatosaurine verts that has cervical rib loops that are wide, but not very deep. Check out this lumpen atrocity at Dinosaur Journey–and more importantly, the apatosaur cervical he’s freaking out over:

UPDATE just a few minutes later: Mike reminded me in the comments about the Tokyo apatosaurine, NSMT-PV 20375, which has wide-but-not-deep cervical ribs. In fact, C7 (the vertebra on the right in this figure) is a pretty good match for the Salt Wash specimen:

UpchurchEtAl2005-apatosaurus-plate2-C3-6-7

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

UPDATE the 2nd: After looking at it for a few minutes, I decided that C7 of the Tokyo apatosaurine was such a good match for the Salt Wash specimen that I wanted to know what it would look like if it was similarly sectioned by erosion. In the Salt Wash specimen, the prezygs are sticking out a little farther than the condyle and cervical rib sections. The red line in this figure is my best attempt at mimicking that erosional surface on the Tokyo C7, and the black outlines on the right are my best guess as to what would be exposed by such a cut (or pair of cuts). I’ve never seen NSMT-PV 20375 in person, so this is just an estimate, but I don’t think it can be too inaccurate, and it is a pretty good match for the Salt Wash specimen.

Another way to put it: if this is an apatosaurine, everything fits. Even the wide-but-not-low-hanging cervical ribs are reasonable in light of some other apatosaurines. If we think this is Camarasaurus just because the cervical ribs aren’t low-hanging, then the pneumatic complexity, the height of the prezygs, and the ventrolateral process on the cervical rib are all anomalous. The balance of the evidence says that this is an apatosaurine, either a small, anterior vert from a big one, or possibly something farther back from a small one. And that’s pretty satisfying.

One more thing: can we take a moment to stand in awe of this freaking thumb-sized cobble that presumably got inside the vertebra through one its pneumatic foramina and rattled around until it got up inside the condyle? Where, I’ll note, the internal structure looks pretty intact despite being filled with just, like, gravel. As someone who spends an inordinate amount of time thinking about how pneumatic vertebrae get buried and fossilized, I am blown away by this. Gaze upon its majesty, people!

This is another “Road to Jurassic Reimagined, Part 2″ post. As before, Part 1 is here, Part 2 will be going up here in the near future. As always, stay tuned.

References

Since I posted my preprint “Almost all known sauropod necks are incomplete and distorted” and asked in the comments for people to let me know if I missed any good necks, the candidates have been absolutely rolling in:

I will be investigating the completeness of all of these and mentioning them as appropriate when I submit the revision of this paper. (In retrospect, I should have waited a week after posting the preprint before submitting for formal review; but I was so scared of letting it brew for years, as we’re still doing with the Barosaurus preprint to our shame, that I submitted it immediately.)

So we probably have a larger number of complete or near-complete sauropod necks than the current draft of this paper suggests. But still very few in the scheme of things, and essentially none that aren’t distorted.

So I want to consider why we have such a poor fossil record of sauropod necks. All of the problems with sauropod neck preservation arise from the nature of the animals.

First, sauropods are big. This is a recipe for incompleteness of preservation. (It’s no accident that the most completely preserved specimens are of small individuals such as CM 11338, the cow-sized juvenile Camarasaurus lentus described by Gilmore, 1925). For an organism to be fossilised, the carcass has to be swiftly buried in mud, ash or some other substrate. This can happen relatively easily to small animals, such as the many finely preserved stinkin’ theropods from the Yixian Formation in China, but it’s virtually impossible with a large animal. Except in truly exceptional circumstances, sediments simply don’t get deposited quickly enough to cover a 25 meter, 20 tonne animal before it is broken apart by scavenging, decay and water transport.

Taylor 2015: Figure 5. Quarry map of Tendaguru Site S, Tanzania, showing incomplete and jumbled skeletons of Giraffatitan brancai specimens MB.R.2180 (the lectotype, formerly HMN SI) and MB.R.2181 (the paralectotype, formerly HMN SII). Anatomical identifications of SII are underlined. Elements of SI could not be identified with certainty. From Heinrich (1999: figure 16), redrawn from an original field sketch by Werner Janensch.

Taylor 2015: Figure 5. Quarry map of Tendaguru Site S, Tanzania, showing incomplete and jumbled skeletons of Giraffatitan brancai specimens MB.R.2180 (the lectotype, formerly HMN SI) and MB.R.2181 (the paralectotype, formerly HMN SII). Anatomical identifications of SII are underlined. Elements of SI could not be identified with certainty. From Heinrich (1999: figure 16), redrawn from an original field sketch by Werner Janensch.

Secondly, even when complete sauropods are preserved, or at least complete necks, distortion of the preserved cervical vertebrae is almost inevitable because of their uniquely fragile construction. As in modern birds, the cervical vertebrae were lightened by extensive pneumatisation, so that they were more air than bone, with the air-space proportion typically in the region of 60–70% and sometimes reaching as high as 89%. While this construction enabled the vertebrae to withstand great stresses for a given mass of bone, it nevertheless left them prone to crushing, shearing and torsion when removed from their protective layer of soft tissue. For large cervicals in particular, the chance of the shape surviving through taphonomy, fossilisation and subsequent deformation would be tiny.

So I think we’re basically doomed never to have a really good sauropod neck skeleton.

Well, I’m a moron again. In the new preprint that I just published, I briefly discussed the six species of sauropod for which complete necks are known — Camarasaurus lentus (but it’s a juvenile), Apatosaurus louisae (but the last three and maybe C5 are badly damaged), Mamenchisaurus hochuanensis (but all the vertebrae are broken and distorted), Shunosaurus lii, Mamenchisaurus youngi and Spinophorosaurus nigerensis.

I did have the wit to say, in the Author Comment:

Although I am submitting this article for formal peer-review at the same time as publishing it as a preprint, I also solicit comments from readers. In particular I am very keen to know if I have missed any complete sauropod necks that have been described in the literature. In the final version of the manuscript, I will acknowledge those who have offered helpful comments.

Happily, several people have taken me up on this (see the comments on the preprint), but one suggestion in particular was a real D’oh! moment for me. Oliver Demuth reminded me about Kaatedocus — a sauropod that we SV-POW!sketeers love so much that it has its own category on our site and we’ve held it up as an example of how to illustrate a sauropod specimen. More than that: we have included several illustrations of its vertebrae in one of our own papers.

Aaanyway … the purpose of this post is just to get all the beautiful Kaatedocus multiview images up in one convenient place. They were freely available as supplementary information to the paper, but now seem to have vanished from the publisher’s web-site. I kept copies, and now present them in the conveniently viewable JPEG format (rather the download-only TIFF format of the originals) and with each image labelled with its position in the column.

Please note, these images are the work of Tschopp and Mateus (2012) — they’re not mine!

Atlas and axis (C1-2)

Atlas and axis (C1-2)

C3

C3

C4

C4

C5

C5

C6

C6

C7

C7

C8

C8

C9

C9

C10

C10

C11

C11

C12

C12

C13

C13

C14

C14

C15 (and the rest of the skeleton) is missing, which makes this a very nearly, but not quite, complete sauropod neck.

Reference

  • Tschopp, Emanuel, and Octávio Mateus. 2012. The skull and neck of a new flagellicaudatan sauropod from the Morrison Formation and its implication for the evolution and ontogeny of diplodocid dinosaurs. Journal of Systematic Palaeontology 11(7):853-888. doi:10.1080/14772019.2012.746589

Here I am at SVPCA in 2015. I am haunted by the fact that ten years ago at SVPCA 2005, I gave a talk about the NHM’s Tendaguru brachiosaurid, NHMUK R5937. And the description is still not done and submitted a full decade later. Even though it’s objectively one of the most beautiful specimens in the world:

dorsals-ab-composite

So here is my pledge to the world:

By this time next year (i.e. the start of SVPCA 2016 in Liverpool), I will have written and submitted this description. If I fail, I give you all permission — no, I beg you — to mock me mercilessly. Leave mocking comments on this blog, yes; but more than that, those of you at SVPCA are invited to spend the week pointing contemptuously at me and saying “Ha!”

Let’s hope it doesn’t come to that.

Update (6 September): see also.

Last October, Mike posted a tutorial on how to choose a paper title, then followed it up by evaluating the titles of his own papers. He invited me to do the same for my papers. I waited a few days to allow myself to forget Mike’s comments on our joint papers – not too hard during my fall anatomy teaching – and then wrote down my thoughts.

And then did nothing with them for three and a half months.

The other day I rediscovered that draft and thought, hey, I don’t remember anything I wrote back then, I should redo the experiment and see if my evaluations will be consistent. And this time without looking at Mike’s post at all, so the risk of contamination would be even lower.

BUT FIRST I thought I should write down what I admire in paper titles, so I could see whether my titles actually lived up to my ideals. So now we can compare:

  • what I say I like in paper titles;
  • what I actually titled my papers;
  • what I had to say about my titles last October;
  • what I have to say about them now;
  • and, for some of my papers, what Mike had to say about them.

What I Admire In Paper Titles

Brevity. I first became consciously aware of the value of concise titles when I read Knut Schmidt-Nielsen’s autobiography, The Camel’s Nose, in 2004 or 2005. (Short-short review: most of the book is a narrative about scientific questions and it’s great, the self-congratulatory chapters near the end are much less interesting. Totally worth reading, especially since used copies can be had for next to nothing.) Schmidt-Nielsen said he always preferred short, simple titles. Short titles are usually punchy and hard to misunderstand. And I like titles that people can remember, and a short title is easier to recall than a long one.

Impact. In short, maximum information transfer using the minimum number of words. This is a separate point from sheer brevity; a paper can have a short title that doesn’t actually tell you very much. But brevity helps, because it’s difficult to compose a long title that really hits hard. Whatever impact a title might have, it will be diluted by every extraneous word.

Full sentences as titles. This is taking the information-transfer aspect of the last admirable quality to its logical extreme, although often at the expense of brevity. I was heavily influenced here by two things that happened while I was at Berkeley. First, I taught for a year in an NSF GK-12 program, where graduate students went out into local elementary, middle, and high schools and taught biology enrichment classes. One thing that was drilled into us during that experience is that we were teaching concepts, which ideally would be expressed as complete sentences. Also about that same time I read James Valentine’s book On the Origin of Phyla. The table of contents of that book is several pages long, because every chapter title, heading, and subheading is a complete sentence. This has a lovely effect: once you’ve read the table of contents of the book or any of its parts, you’ve gotten the TL;DR version of the argument. Sort of like a distributed abstract. I’d like to do that more.

How Did I Do?

Time to see if my actions match my words. Full bibliographic details and PDFs are available on my publications page. I stuck with Mike’s red-blue-green color scheme for the verdicts. My October 2014 and February 2015 thoughts are labeled. For joint papers with Mike, I’ve copied his assessment in as well. Any comments in brackets are my editorializing now, comparing what I said in October to what I said a few days ago before I’d looked back at my old comments or Mike’s.

* * * * * * * * * * * *

Sauroposeidon proteles, a new sauropod from the Early Cretaceous of Oklahoma. (11 words)

GOOD
Oct 2014: Like it. Short, to the point, includes the taxon name.
Feb 2015: Good, gets the job done with a minimum of fuss

Osteology, paleobiology, and relationships of the sauropod dinosaur Sauroposeidon. (9 words)

OK
Oct 2014: This title was inspired by the papers from the early 20th century
Feb 2015: It gets the job done, I suppose. I can’t help but wonder if there might have been a more elegant solution. Part of my unease is that this title is an example of the same attitude that produced the next monstrosity.

Osteological correlates of cervical musculature in Aves and Sauropoda (Dinosauria: Saurischia), with comments on the cervical ribs of Apatosaurus. (19 words)

BAD
Oct 2014: Ugh. It gets the job done, I suppose, but it’s waaaay long and just kind of ugly.
Feb 2015: Ugh. Waaay too wordy. I had a (fortunately brief) fascination with long titles, and especially the phrase, “with comments on”. Now I would cut it down to “Bony correlates of neck muscles in birds and sauropod dinosaurs” (10 words)

Vertebral pneumaticity, air sacs, and the physiology of sauropod dinosaurs. (10 words)

OK
Oct 2014: Like it. Would be better made into a sentence, like, “Vertebral pneumaticity is evidence for air sacs in sauropod dinosaurs.”
Feb 2015: Fairly clean. Does what it says on the tin. I’m having a hard time seeing how it could be turned into a sentence and still convey so much of what the paper is about in so few words.

[Heh. As we will see again later on, I was evidently smarter last fall than I am now.]

The evolution of vertebral pneumaticity in sauropod dinosaurs. (8 words)

GREAT

Oct 2014: Like it. It couldn’t really be any shorter without losing crucial information. Happy to have a decent title on my second-most-cited paper!
Feb 2015: Short, clean, probably my best title ever.

First occurrence of Brachiosaurus (Dinosauria: Sauropoda) from the Upper Jurassic Morrison Formation of Oklahoma. (14 words)

OK
Oct 2014: Yep. once you’ve read the title, you barely need to read the paper. Even better would have been, “A metacarpal of Brachiosaurus from the Upper Jurassic Morrison Formation of Oklahoma.” (12 words)
Feb 2015: Does what it says, but like my other PaleoBios pub, it’s a long title for a short paper. Now I would title it, “First record of the sauropod dinosaur Brachiosaurus from Oklahoma” (9 words)

[my October title was better!]

Postcranial skeletal pneumaticity in sauropods and its implications for mass estimates. (11 words)

OK
Oct 2014: It’s not elegant but it gets the job done. I wanted that paper to be one-stop shopping for sauropod PSP, but of course the real payoff there is the ASP/mass-estimate stuff, so I’m happy to have punched that up in the title.
Feb 2015: Good enough. I like it. It’s a little long–I could reasonably have just titled this, “Postcranial skeletal pneumaticity in sauropods”, but I wanted to draw attention to the implications for mass estimates.

Sauroposeidon: Oklahoma’s native giant (4 words)

OK
Feb 2015: Nice and short. Not terribly informative, but since this was a narrative about the discovery and description of Sauroposeidon aimed mostly at an Oklahoma audience, it’s not obvious how it could be improved.
[Note sure how missed this one last October, but I did.]

Origin of postcranial skeletal pneumaticity in dinosaurs. (7 words)

GOOD
Oct 2014: About all I would change now would be to add the word “early” at the beginning of the title.
Feb 2015: Great. Could not be shortened further without losing information.

What pneumaticity tells us about ‘prosauropods’, and vice versa. (9 words)

GOOD

Oct 2014: Love this title. I used it for the abstract of the SVP talk that the paper was derived from, too.
Feb 2015: Kind of a gimmick title, but it’s accurate–the SVP abstract this paper was based on was built around a bullet list. And it’s still nice and short.

Evidence for bird-like air sacs in saurischian dinosaurs. (9 words)

GOOD

Oct 2014: Along with Wedel (2003b) and Wedel (2006), this has a short (7-9 words apiece) title that tells you what’s in the paper, simply and directly. For once, I’m glad I didn’t turn it into a sentence. I think a declarative statement like “Saurischian dinosaurs had air sacs like those of birds” would have been less informative and come off as advertising. I wanted this paper to do what the title said: run down the evidence for air sacs in saurischians.
Feb 2015: I like it and wouldn’t change it. The “evidence for” part is key – I didn’t want to write a paper primarily about the air sacs themselves. Instead I wanted to lay out the evidence explaining why we think sauropods had air sacs.

Head and neck posture in sauropod dinosaurs inferred from extant animals. (8 words)

OK
Oct 2014: It’s not horrible but it would be better as a declarative statement like, “Sauropod dinosaurs held their necks and heads elevated like most other tetrapods.” (12 words)
Feb 2015: Good. Reads almost telegraphically brief as it is. Does what it says on the tin.

Mike: RUBBISH

[October Matt wins again!]

A new sauropod dinosaur from the Lower Cretaceous Cedar Mountain Formation, Utah, USA. (13 words)

OK
Oct 2014: Two things about this one. First, I wish we’d been able to include the taxon name in the title, as we were allowed to do back in the day for Sauroposeidon. Second, I know some people whinge about us using the CMF in the title and in the paper instead of the Burro Canyon Fm, which is what the CMF is technically called east of the Colorado River. But srsly, how many people search for Burro Canyon Fm versus CMF? All of the relevant faunal comparisons are to be made with the CMF, so I don’t feel the least bit bad about this.
Feb 2015: Fine. About as short as it could be and still be informative.

Mike: RUBBISH

The long necks of sauropods did not evolve primarily through sexual selection. Journal of Zoology. (12 words)

GOOD
Oct 2014: Perfect. The abstract and the paper expand on the title, but if all you read is the title, you know what we found. That’s a worthy goal.
Feb 2015: My first sentence title. Every word does work, so even though this is one of my longer titles, I like it. The length relative to my other titles is not a knock against this one; rather, it emphasizes how well I did at keeping my early titles short and to the point (with a couple of regrettable exceptions as noted above).

Mike: SWEET

The early evolution of postcranial skeletal pneumaticity in sauropodomorph dinosaurs. (10 words)

GOOD
Oct 2014: Not bad. I wonder if something like, “Widespread vertebral fossae show that pulmonary pneumaticity evolved early in sauropodomorphs” might be better. It’s hard, though, to put so many long, polysyllabic words in a title that doesn’t sound like a train wreck. At a minimum, this paper does what it says on the tin.
Feb 2015: Short and to the point. Another one that couldn’t be any shorter without losing valuable information.

A monument of inefficiency: the presumed course of the recurrent laryngeal nerve in sauropod dinosaurs. (15 words)

Objectively: BAD to OK
Subjectively: GOOD to FREAKIN’ AWESOME
Oct 2014: I readily admit that I could have fashioned a more informative title, but I dearly love this one. It’s derived from a TV commercial for cheeseburgers (true story), and it warms my heart every time I read it.
Feb 2015: This is definitely a gimmick title that is longer than it has to be (it would be a concise 11 words without the unnecessary intro clause) BUT I love it and I’d do it exactly the same if I could do it again. So there!

Why sauropods had long necks; and why giraffes have short necks. (11 words)

GOOD
Oct 2014: This is one of those ‘draw the reader in’ titles. I like it.
Feb 2015: We both liked the even shorter, “Why giraffes have short necks” but we really felt that a paper about sauropod necks needed sauropod necks in the title. I feel about this one like I feel about my 2007 prosauropod paper: it’s a gimmick title, but it’s short, so no harm done.

Mike: EXCELLENT

Neural spine bifurcation in sauropod dinosaurs of the Morrison Formation: ontogenetic and phylogenetic implications. (14 words)

OK
Oct 2014: Blah. It’s okay, not great. Maybe better as, “No evidence for increasing neural spine bifurcation through ontogeny in diplodocid sauropods of the Morrison Formation”, or something along those lines.
Feb 2015: This one is long but I think the length is necessary. It’s also kinda boring, but it was addressing a fairly dry point. I think any attempt to shorten it or sexy it up would come off as gratuitous.

Mike: WEAK

The effect of intervertebral cartilage on neutral posture and range of motion in the necks of sauropod dinosaurs. (18 words)

OK
Oct 2014: Probably better along the lines of, “Intervertebral spacing suggests a high neutral posture and broad range of motion in the necks of sauropod dinosaurs” or something like that.
Feb 2015: My second-longest title ever! Looking at it now, I think we could have titled it, “Effects of intervertebral cartilage on neck posture and range of motion in sauropod dinosaurs” and gotten it down to 14 words, but the word ‘neutral’ is doing real work in the original so maybe that’s a bust.

Mike: UGH, rubbish.

[October Matt is up by three points at least]

Caudal pneumaticity and pneumatic hiatuses in the sauropod dinosaurs Giraffatitan and Apatosaurus. (12 words)

OK
Oct 2014: Along the same lines as the previous: “Caudal pneumaticity and pneumatic hiatuses show that pulmonary diverticula in the tails of sauropod dinosaurs were pervasive and complex” or something.
Feb 2015: Good. Long only by comparison with some of my earlier titles. Does what it says.

Mike: NOT GOOD ENOUGH

The neck of Barosaurus was not only longer but also wider than those of Diplodocus and other diplodocines. (18 words)

GOOD
Feb 2015: My second sentence-as-title, and another entry in the run of mostly long titles from 2012 onward. I like how precise it is, despite the length.

Mike: GOOD

A ceratopsian dinosaur from the Lower Cretaceous of Western North America, and the biogeography of Neoceratopsia. (16 words)

GOOD
Feb 2015: I had no say in this one (by choice, I’m sure Andy et al. would have listened if I had had any suggestions about the title, but I didn’t). If I could rewrite it, I’d probably make it even longer by adding in the word ‘new’ between A and ceratopsian

Haplocanthosaurus (Saurischia: Sauropoda) from the lower Morrison Formation (Upper Jurassic) near Snowmass, Colorado. (13 words)

OK
Feb 2015: Feels a lot longer than its 13 words, mostly because so many of the words are polysyllabic. Normally I like pulling the words in parentheses out, but in this case I can’t see that doing that would actually improve the title. Sometimes descriptive papers need plain titles. It’s okay.

* * * * * * * * * * * *

Lessons

First, Mike graded harder than I did. In fact, I only rated one of my titles as BAD, which seems a bit feeble. I think we were using different criteria. If a title was boring but serviceable, I gave it an OK, whereas Mike tended to flag any suboptimal title as RUBBISH. But I didn’t remember that about his post, and I deliberately avoided looking at it until I’d made my evaluations.

Second, except for the two PaleoBios papers, all of the titles from the first half of my career (2000-2007) are 12 words or fewer, including a substantial bundle from before I’d read either The Camel’s Nose or Strunk & White. I’m sure that being a Cifelli student and then a Padian student had something to do with that; Rich and Kevin made me into the word choice and grammar pedant that I am today (my rhetorical excrescences on this site are my fault, not theirs).

Third, much to my surprise and consternation, my titles have gotten longer over time, not shorter. Partly that’s because my little corner of the science ecosystem is getting increasingly subdivided, so it’s hard for me to write a paper now with a title as broad as, “The evolution of vertebral pneumaticity in sauropod dinosaurs.” (Possibly a prod to keep seeking out new, more open horizons?) And I suppose there is some tension between brevity, informativeness, and precision. For example, saying in the title of a descriptive paper than a specimen is “from the Upper Jurassic Morrison Formation of [Location], [State or Country]” adds 11 words, but the title really does need those words. That could be a segue into a whole other discussion about descriptive versus analytical work, but that will be a topic for another time.

Ultimately, this has been a fun exercise and it’s made me more aware of how I title my papers. This is useful because I have some manuscripts in the works that deal with really detailed anatomy, and I need to figure out how to give them titles that are precise and informative but still punchy. It’s not easy.

Parting thought: after I posted the slides from my photography and illustration talk, Mike and I talked about posting some of our figures and dissecting them to see how they could be improved (it’s axiomatic that almost all figures could be improved in one way or another). We should really get started on that.

In light of yesterday’s tutorial on choosing titles, here are the titles of all my own published papers (including co-authored ones), in chronological order, with my own sense of whether I’m happy with them now I look back. All the full references are on my publications page (along with the PDFs). I’ll mark the good ones in green, the bad ones in red and the merely OK in blue.

The Phylogenetic Taxonomy of Diplodocoidea (Dinosauria: Sauropoda).

OK, I suppose. It does at least clearly state what the paper is about. I’ll give myself a pass on this since it was my very first paper.

Dinosaur diversity analysed by clade, age, place and year of description.

NOT BAD, since the paper was basically a list of many, many results that could hardly have been summarised in the title. I give myself some points for listing the ways I analysed the data, rather than just saying “An analysis of dinosaur diversity” or something equally uninformative.

Phylogenetic definitions in the pre-PhyloCode era; implications for naming clades under the PhyloCode.

NOT BAD again, I suppose, since it was a discussion paper that couldn’t be summarised in a short title. Could I have said what the alluded-to implications are? I think probably not, in a reasonably concise title.

An unusual new neosauropod dinosaur from the Lower Cretaceous Hastings Beds Group of East Sussex, England.

RUBBISH, since it doesn’t name the new dinosaur (which was of course Xenoposeidon). I was young and stupid back then, and just followed convention. In mitigation, it does at least say when and where the specimen is from.

Case 3472: Cetiosaurus Owen, 1841 (Dinosauria, Sauropoda): proposed conservation of usage by designation of Cetiosaurus oxoniensis Phillips, 1871 as the type species.

DOUBLE-PLUS UGLY. But I am going to blame the journal on this one — they have a very firmly defined format for petition titles.

Head and neck posture in sauropod dinosaurs inferred from extant animals.

RUBBISH. What was I thinking, and why did my SV-POW!sketeer co-authors let me choose such an uninformative title? We should of course have gone with a title that says what posture we inferred. The associated blog-post had a much better title: Sauropods held their necks erect … just like rabbits.

A re-evaluation of Brachiosaurus altithorax Riggs 1903 (Dinosauria, Sauropoda) and its generic separation from Giraffatitan brancai (Janensch 1914).

ADEQUATE, since the title strongly implies the conclusion (generic separation) even if doesn’t quite come out and say it.

Electronic publication of nomenclatural acts is inevitable, and will be accepted by the taxonomic community with or without the endorsement of the Code.

BRILLIANT. The best title in my CV. You hardly even need to read the paper once you’ve read the title. The only downside: it’s 12 characters too long to tweet.

Sharing: public databases combat mistrust and secrecy.

GOOD, but I can’t take the credit for that (A) because I was third author behind Andy Farke and Matt, and (B) because the journal chose the title.

The Open Dinosaur Project.

OK, but we should have done better. Something like “The Open Dinosaur Project recruits volunteer effort to analyse dinosaur evolution”. Or, if we were being honest (and prescient), “The Open Dinosaur Project will lie embarrassingly moribund for more than two years”.

Sauropod dinosaur research: a historical review.

OK, since it does say what the paper is. But this title is not as good as that of the talk it was based on, “The evolution of sauropod dinosaurs from 1841 to 2008”. (I notice that Mark Witton nicked my title for his talk at TetZooCon.)

Running a question-and-answer website for science education: first hand experiences.

UNOBJECTIONABLE, but not my choice anyway — lead author Dave Hone presumably picked it. Could have done better by stating what at least one of those experiences was.

A new sauropod dinosaur from the Lower Cretaceous Cedar Mountain Formation, Utah, USA.

RUBBISH. At least this time it wasn’t entirely my fault. When we submitted this to Acta Palaeontologica Polonica, it was called “Brontomerus mcintoshi, a new sauropod dinosaur from the Lower Cretaceous Cedar Mountain Formation, Utah, USA”, but the journal made us take the taxon name out of the title. Why? Why why WHY?

The long necks of sauropods did not evolve primarily through sexual selection.

SWEET.

Why sauropods had long necks; and why giraffes have short necks.

EXCELLENT. Short, appealing and (hopefully) funny. When I give talks based on this paper, I use the even better short version, just “Why giraffes have short necks”. But that seemed a bit too cute for an academic setting.

Neural spine bifurcation in sauropod dinosaurs of the Morrison Formation: ontogenetic and phylogenetic implications.

WEAK. We should have stated the conclusion: a title like “Neural spine bifurcation in sauropods of the Morrison Formation is not an ontogenetic feature, but is phylogenetically significant” would have been better.

The neck of Barosaurus was not only longer but also wider than those of Diplodocus and other diplodocines.

GOOD. Not particularly exciting, but explicit.

Caudal pneumaticity and pneumatic hiatuses in the sauropod dinosaurs Giraffatitan and Apatosaurus.

NOT GOOD ENOUGH. We should have stated the main finding: “Caudal pneumaticity and pneumatic hiatuses reveal cryptic diverticula in the sauropod dinosaurs Giraffatitan and Apatosaurus“.

The effect of intervertebral cartilage on neutral posture and range of motion in the necks of sauropod dinosaurs.

UGH, rubbish. What the heck was I thinking? I should have written this post a couple of years ago, and used it to make me choose a much better title. As it is, it just leaves the reader assuming intervertebral cartilage probably has some effect, but they have no idea what.

 —

I make that six good titles, seven bad ones and six indifferent. Awarding two points per good title and one per adequate title, I give myself 18 points out of a possible 38 — slightly less than half, at 47%. More worryingly, there’s no apparent trend towards choosing better titles.

Must do better.

 

[NOTE: see the updates at the bottom. In summary, there’s nothing to see here and I was mistaken in posting this in the first place.]

Elsevier’s War On Access was stepped up last year when they started contacting individual universities to prevent them from letting the world read their research. Today I got this message from a librarian at my university:

babys-first-takedown

The irony that this was sent from the Library’s “Open Access Team” is not lost on me. Added bonus irony: this takedown notification pertains to an article about how openness combats mistrust and secrecy. Well. You’d almost think NPG wants mistrust and secrecy, wouldn’t you?

It’s sometimes been noted that by talking so much about Elsevier on this blog, we can appear to be giving other barrier-based publishers a free ride. If we give that impression, it’s not deliberate. By initiating this takedown, Nature Publishing Group has self-identified itself as yet another so-called academic publisher that is in fact an enemy of science.

So what next? Anyone who wants a PDF of this (completely trivial) letter can still get one very easily from my own web-site, so in that sense no damage has been done. But it does leave me wondering what the point of the Institutional Repository is. In practice it seems to be a single point of weakness allowing “publishers” to do the maximum amount of damage with a single attack.

But part of me thinks the thing to do is take the accepted manuscript and format it myself in the exact same way as Nature did, and post that. Just because I can. Because the bottom line is that typesetting is the only actual service they offered Andy, Matt and me in exchange for our right to show our work to the world, and that is a trivial service.

The other outcome is that this hardens my determination never to send anything to Nature again. Now it’s not like my research program is likely to turn up tabloid-friendly results anyway, so this is a bit of a null resolution. But you never know: if I happen to stumble across sauropod feather impressions in an overlooked Wealden fossil, then that discovery is going straight to PeerJ, PLOS, BMC, F1000 Research, Frontiers or another open-access publisher, just like all my other work.

And that’s sheer self-interest at work there, just as much as it’s a statement. I will not let my best work be hidden from the world. Why would anyone?

Let’s finish with another outing for this meme-ready image.

Publishers ... You're doing it wrong

Update (four hours later)

David Mainwaring (on Twitter) and James Bisset (in the comment below) both pointed out that I’ve not seen an actual takedown request from NPG — just the takedown notification from my own library. I assumed that the library were doing this in response to hassle from NPG, but of course it’s possible that my own library’s Open Access Team is unilaterally trying to prevent access to the work of its university’s researchers.

I’ve emailed Lyn Duffy to ask for clarification. In the mean time, NPG’s Grace Baynes has tweeted:

So it looks like this may be even more bizarre than I’d realised.

Further bulletins as events warrant.

Update 2 (two more hours later)

OK, consensus is that I read this completely wrong. Matt’s comment below says it best:

I have always understood institutional repositories to be repositories for author’s accepted manuscripts, not for publisher’s formatted versions of record. By that understanding, if you upload the latter, you’re breaking the rules, and basically pitting the repository against the publisher.

Which is, at least, not a nice thing to do to the respository.

So the conclusion is: I was wrong, and there’s nothing to see here apart from me being embarrassed. That’s why I’ve struck through much of the text above. (We try not to actually delete things from this blog, to avoid giving a false history.)

My apologies to Lyn Duffy, who was just doing her job.

Update 3 (another hour later)

This just in from Lyn Duffy, confirming that, as David and James guessed, NPG did not send a takedown notice:

Dear Mike,

This PDF was removed as part of the standard validation work of the Open Access team and was not prompted by communication from Nature Publishing. We validate every full-text document that is uploaded to Pure to make sure that the publisher permits posting of that version in an institutional repository. Only after validation are full-text documents made publicly available.

In this case we were following the regulations as stated in the Nature Publishing policy about confidentiality and pre-publicity. The policy says, ‘The published version — copyedited and in Nature journal format — may not be posted on any website or preprint server’ (http://www.nature.com/authors/policies/confidentiality.html). In the information for authors about ‘Other material published in Nature’ it says, ‘All articles for all sections of Nature are considered according to our usual conditions of publication’ (http://www.nature.com/nature/authors/gta/others.html#correspondence). We took this to mean that material such as correspondence have the same posting restrictions as other material published by Nature Publishing.

If we have made the wrong decision in this case and you do have permission from Nature Publishing to make the PDF of your correspondence publicly available via an institutional repository, we can upload the PDF to the record.

Kind regards,
Open Access Team

Appendix

Here’s the text of the original notification email so search-engines can pick it up. (If you read the screen-grab above, you can ignore this.)

University of Bristol — Pure

Lyn Duffy has added a comment

Sharing: public databases combat mistrust and secrecy
Farke, A. A., Taylor, M. P. & Wedel, M. J. 22 Oct 2009 In : Nature. 461, 7267, p. 1053

Research output: Contribution to journal › Article

Lyn Duffy has added a comment 7/05/14 10:23

Dear Michael, Apologies for the delay in checking your record. It appears that the document you have uploaded alongside this record is the publishers own version/PDF and making this version openly accessible in Pure is prohibited by the publisher, as a result the document has been removed from the record. In this particular instance the publisher would allow you to make accessible the postprint version of the paper, i.e., the article in the form accepted for publication in the journal following the process of peer review. Please upload an acceptable version of the paper if you have one. If you have any questions about this please get back to us, or send an email directly to open-access@bristol.ac.uk Kind regards, Lyn Duffy Library Open Access Team.

Better

AMNH T. rex mount, photo by Mike Taylor.

In a recent comment, Doug wrote:

If I want to be a truly educated observer of Tyrannosaurus rex mounts, what 5 things should I look for in a reconstruction to assess if it is true to our current scientific understanding? I’m not talking tail dragging/upright at this point…we are well past that I hope.

If he had asked about Apatosaurus, I could have written him a novel. But it is a point of pride with me not to contribute to the over-application of human attention to T. rex; not only would it be vulgar, it would also be a waste of resources, considering how many people already have that covered. So, you theropod workers and avocational “rexperts”, we’re finally inviting you to the high table. Please, tell us–and Doug–what separates the good T. rex mounts from the crappy ones. Big piles of SV-POW! bucks will be showered on whoever brings the most enlightenment, especially if you adhere to the requested List of 5 Things format.

The comment lines are open–go!

[This is part 4 in an ongoing series on our recent PLOS ONE paper on sauropod neck cartilage. See also part 1, part 2, and part 3.]

Big Bend Vanessa 182 small

Weird stuff on the ground, Big Bend, 2007.

Here’s a frequently-reproduced quote from Darwin:

About thirty years ago there was much talk that geologists ought only to observe and not theorise; and I well remember some one saying that at this rate a man might as well go into a gravel-pit and count the pebbles and describe the colours. How odd it is that anyone should not see that all observation must be for or against some view if it is to be of any service!

It’s from a letter to Henry Fawcett, dated September 18, 1861, and you can read the whole thing here.

I’ve known this quote for ages, having been introduced to it at Berkeley–a copy used to be taped to the door of the Padian Lab, and may still be. It’s come back to haunt me recently, though. An even stronger version would run something like, “If you don’t know what you’re looking for, you won’t make the observation in the first place!”

OLYMPUS DIGITAL CAMERA

Kent Sanders looking at scans of BYU 12613, a posterior cervical of either Kaatedocus or an anomalously small Diplodocus, at the University of Utah in May, 2008.

For example: I started CT scanning sauropod vertebrae with Rich Cifelli and Kent Sanders back in January, 1998. Back then, I was interested in pneumaticity, so that’s what I looked for, and that’s what I found–work which culminated in Wedel et al. (2000) and Wedel (2003). It wasn’t until earlier this year that I wondered if it would be possible to determine the spacing of articulated vertebrae from CT scans. So everything I’m going to show you, I technically saw 15 years ago, but only in the sense of “it crossed my visual field.” None of it registered at the time, because I wasn’t looking for it.

A corollary I can’t help noting in passing: one of the under-appreciated benefits of expanding your knowledge base is that it allows you to actually make more observations. Many aspects of nature only appear noteworthy once you have a framework in which to see them.

OLYMPUS DIGITAL CAMERA

BYI 12613 going through a CT scanner at the University of Utah medical center. We were filming for the “Megasaurus” episode of Jurassic CSI. That shoot was crazy fun.

So anyway, the very first specimen we scanned way back when was the most anterior of the three plaster jackets that contain the four cervical vertebrae that make up OMNH 53062, which was destined to become the holotype of Sauroposeidon. I’ve written about the taphonomy of that specimen here, and you can read more about how it was excavated in Wedel and Cifelli (2005). We scanned that jacket first because, although the partial vertebrae it contains are by far the most incomplete of the four, the jacket is a lot smaller and lighter than the other two (which weigh hundreds of pounds apiece). Right away we saw internal chambers in the vertebrae, and that led to all of the pneumaticity work mentioned above.

Sauroposeidon C5 cross section Wedel 2007b fig 14

Internal structure of a cervical vertebra of Sauroposeidon, OMNH 53062. A, parts of two vertebrae from the middle of the neck. The field crew that dug up the bones cut though one of them to divide the specimen into manageable pieces. B, cross section of C6 in posterior view at the level of the break, traced from a CT image and photographs of the broken end. The left side of the specimen was facing up in the field and the bone on that side is badly weathered. Over most of the broken surface the internal structure is covered by plaster or too damaged to trace, but it is cleanly exposed on the upper right side (outlined). C, the internal structure of that part of the vertebra, traced from a photograph. The arrows indicate the thickness of the bone at several points, as measured with a pair of digital calipers. The camellae are filled with sandstone. Wedel (2007: fig. 14).

Happily for me, that first jacket contains not only the posterior two-thirds of the first vertebra (possibly C5), but also the front end of the second vertebra. Whoever decided to plow through the second vertebra to divide the specimen into manageable chunks in the field made a savvy choice. Way back in 2004 I realized that the cut edge of the second vertebra was not obscured by plaster, and therefore the internal structure could be seen and measured directly, which is a lot cleaner than relying on the artifact-heavy CT scans. (The CT scans are noisy because the hospital machines we had access to start to pant a bit when asked to punch x-rays through specimens this large and dense.) A figure derived from that work made it into a couple of papers and this post, and appears again above.

But that’s pneumaticity, which this post is allegedly not about. The cut through the second vertebra was also smart because it left the intervertebral joint intact.

Figure 11. Fifth and partial sixth cervical vertebrae of Sauroposeidon. Photograph and x-ray scout image of C5 and the anterior portion of C6 of Sauroposeidon OMNH 53062 in right lateral view. The anterior third of C5 eroded away before the vertebra was collected. C6 was deliberately cut through in the field to break the multi-meter specimen into manageable pieces for jacketing (see [37] for details). Note that the silhouettes of the cotyle of C5 and the condyle of C6 are visible in the x-ray.

Fifth and partial sixth cervical vertebrae of Sauroposeidon.
Photograph and x-ray scout image of C5 and the anterior portion of C6 of Sauroposeidon OMNH 53062 in right lateral view. The anterior third of C5 eroded away before the vertebra was collected. C6 was deliberately cut through in the field to break the multi-meter specimen into manageable pieces for jacketing (see Wedel and Cifelli 2005 for details). Note that the silhouettes of the cotyle of C5 and the condyle of C6 are visible in the x-ray. Taylor and Wedel (2013: figure 11).

Here are a photo of the jacket and a lateral scout x-ray. The weird rectangles toward the left and right ends of the x-ray are boards built into the bottom of the jacket to strengthen it.

Figure 12. CT slices from fifth cervical vertebrae of Sauroposeidon. X-ray scout image and three posterior-view CT slices through the C5/C6 intervertebral joint in Sauroposeidon OMNH 53062. In the bottom half of figure, structures from C6 are traced in red and those from C5 are traced in blue. Note that the condyle of C6 is centered in the cotyle of C5 and that the right zygapophyses are in articulation.

CT slices from fifth cervical vertebrae of Sauroposeidon.
X-ray scout image and three posterior-view CT slices through the C5/C6 intervertebral joint in Sauroposeidon OMNH 53062. In the bottom half of figure, structures from C6 are traced in red and those from C5 are traced in blue. Note that the condyle of C6 is centered in the cotyle of C5 and that the right zygapophyses are in articulation. Taylor and Wedel (2013: figure 12).

And here’s a closeup of the C5/C6 joint, with the relevant radiographs and tracing. The exciting thing here is that the condyle is centered almost perfectly in the cotyle, and the zygapophyses are in articulation. Together with the lack of disarticulation in the cervical rib bundle (read more about that here and in Wedel et al. 2000), these things suggest to us that the vertebrae are spaced pretty much as they were in life. If so, then the spacing between the vertebrae now tells us the thickness of the soft tissue that separated the vertebrae in life.

I should point out here that we can’t prove that the spacing between the vertebrae is still the same as it was in life. But if some mysterious force moved them closer together or farther apart, it did so (1) without  decentering the condyle of C6 within the cotyle of C5, (2) without moving the one surviving zygapophyseal joint out of contact, and (3) without disarticulating the cervical ribs. The cervical ribs were each over 3 meters long in life and they formed vertically-stacked bundles on either side below the vertebrae; that’s a lot of stuff to move just through any hypothetical contraction or expansion of the intervertebral soft tissues after death. In fact, I would not be surprised if the intervertebral soft tissues did contract or expand after death–but I don’t think they moved the vertebrae, which are comparatively immense. The cartilage probably pulled away from the bone as it rotted, allowing sediment in. Certainly every nook and cranny of the specimen is packed with fine-grained sandstone now.

Anyway, barring actual preserved cartilage, this is a best-case scenario for trying to infer intervertebral spacing in a fossil. If articulation of the centra, zygs, and cervical ribs doesn’t indicate legitimate geometry, nothing ever will. So if we’re going to use the fossils to help settle this at all, we’re never going to have a better place to start.

Figure 14. Geometry of opisthocoelous intervertebral joints. Hypothetical models of the geometry of an opisthocoelous intervertebral joint compared with the actual morphology of the C5/C6 joint in Sauroposeidon OMNH 53062. A. Model in which the condyle and cotyle are concentric and the radial thickness of the intervertebral cartilage is constant. B. Model in which the condyle and cotyle have the same geometry, but the condyle is displaced posteriorly so the anteroposterior thickness of the intervertebral cartilage is constant. C. the C5/C6 joint in Sauroposeidon in right lateral view, traced from the x-ray scout image (see Figure 12); dorsal is to the left. Except for one area in the ventral half of the cotyle, the anteroposterior separation between the C5 cotyle and C6 condyle is remarkably uniform. All of the arrows in part C are 52 mm long.

Geometry of opisthocoelous intervertebral joints.
Hypothetical models of the geometry of an opisthocoelous intervertebral joint compared with the actual morphology of the C5/C6 joint in Sauroposeidon OMNH 53062. A. Model in which the condyle and cotyle are concentric and the radial thickness of the intervertebral cartilage is constant. B. Model in which the condyle and cotyle have the same geometry, but the condyle is displaced posteriorly so the anteroposterior thickness of the intervertebral cartilage is constant. C. the C5/C6 joint in Sauroposeidon in right lateral view, traced from the x-ray scout image (see Figure 12); dorsal is to the left. Except for one area in the ventral half of the cotyle, the anteroposterior separation between the C5 cotyle and C6 condyle is remarkably uniform. All of the arrows in part C are 52 mm long. Taylor and Wedel (2013: figure 14).

So, by now, you know I’m a doofus. I have been thinking about this problem literally for years and the data I needed to address it was sitting on my hard drive the entire time. One of the things I pondered during those lost years is what the best shape for a concave-to-convex intervertebral joint might be. Would the best spacing be radially constant (A in the figure above), or antero-posteriorly constant (B), or some other, more complicated arrangement? The answer in this case surprised me–although the condyle is a lot smaller in diameter than the cotyle, the anteroposterior separation between them in almost constant, as you can see in part C of the above figure.

Figure 13. Joint between sixth and seventh cervicals vertebrae of Sauroposeidon. X-ray scout image of the C6/C7 intervertebral joint in Sauroposeidon OMNH 53062, in right lateral view. The silhouette of the condyle is traced in blue and the cotyle in red. The scale on the right is marked off in centimeters, although the numbers next to each mark are in millimeters.

Joint between sixth and seventh cervicals vertebrae of Sauroposeidon.
X-ray scout image of the C6/C7 intervertebral joint in Sauroposeidon OMNH 53062, in right lateral view. The silhouette of the condyle is traced in blue and the cotyle in red. The scale on the right is marked off in centimeters, although the numbers next to each mark are in millimeters. Taylor and Wedel (2013: figure 13).

Don’t get too worked up about that, though, because the next joint is very different! Here’s the C6/C7 joint, again in a lateral scout x-ray, with the ends of the bones highlighted. Here the condyle is almost as big in diameter as the cotyle, but it is weirdly flat. This isn’t a result of overzealous prep–most of the condyle is still covered in matrix, and I only found its actual extent by looking at the x-ray. This is flatter than most anterior dorsal vertebrae of Apatosaurus–I’ve never seen a sauropod cervical with such a flat condyle. Has anyone else?

The condyle of C6 is a bit flatter than expected, too–certainly a lot flatter than the cervical condyles in Giraffatitan and the BYU Brachiosaurus vertebrae. As we said in the paper,

It is tempting to speculate that the flattened condyles and nearly constant thickness of the intervertebral cartilage are adaptations to bearing weight, which must have been an important consideration in a cervical series more than 11 meters long, no matter how lightly built.

Anyway, obviously here the anteroposterior distance between condyle and cotyle could not have been uniform because they are such different shapes. Wacky. The zygs are missing, so they’re no help, and clearly the condyle is not centered in the cotyle. Whether this posture was attainable in life is debatable; I’ve seen some pretty weird stuff. In any case, we didn’t use this joint for estimating cartilage thickness because we had no reason to trust the results.

Figure 15. First and second dorsal vertebrae of Apatosaurus CM 3390. Articulated first and second dorsal vertebrae of Apatosaurus CM 3390. A. Digital model showing the two vertebrae in articulation, in left lateral (top) and ventral (bottom) views. B-G. Representative slices illustrating the cross-sectional anatomy of the specimen, all in posterior view. B. Slice 25. C. Slice 31. D. Slice 33. E. Slice 37. F. Slice 46. G. Slice 61. Orthogonal gaps are highlighted where the margins of the condyle and cotyle are parallel to each other and at right angles to the plane of the CT slice. 'Zygs' is short for 'zygapophyses', and NCS denotes the neurocentral synchondroses.

First and second dorsal vertebrae of Apatosaurus CM 3390.
Articulated first and second dorsal vertebrae of Apatosaurus CM 3390. A. Digital model showing the two vertebrae in articulation, in left lateral (top) and ventral (bottom) views. B-G. Representative slices illustrating the cross-sectional anatomy of the specimen, all in posterior view. B. Slice 25. C. Slice 31. D. Slice 33. E. Slice 37. F. Slice 46. G. Slice 61. Orthogonal gaps are highlighted where the margins of the condyle and cotyle are parallel to each other and at right angles to the plane of the CT slice. ‘Zygs’ is short for ‘zygapophyses’, and NCS denotes the neurocentral synchondroses. Taylor and Wedel (2013: figure 15).

Kent Sanders and I had also scanned several of the smaller sauropod vertebrae from the Carnegie collection (basically, the ones that would fit in the trunk of my car for the drive back to Oklahoma). Crucially, we’d scanned a couple of sets of articulated vertebrae, CM 3390 and CM 11339, both from juvenile individuals of Apatosaurus. In both cases, the condyles and cotyles are concentric (that’s what the ‘orthogonal gaps’ are all about in the above figure) and the zygs are in articulation, just as in Sauroposeidon. These are dorsals, so we don’t have any cervical ribs here to provide a third line of evidence that the articulation is legit, but all of the evidence that we do have is at least consistent with that interpretation.

So, here’s an interesting thing: in CM 3390, above, the first dorsal is cranked up pretty sharply compared to the next one, but the condyle is still centered in the cotyle and the zygs are in articulation. Now, the vertebrae have obviously been sheared by taphonomic deformation, but that seems to have affected both vertebrae to the same extent, and it’s hard to imagine some kind of taphonomic pressure moving one vertebra around relative to the next. So I think it’s at least plausible that this range of motion was achievable in life. Using various views and landmarks, we estimate the degree of extension here somewhere between 31 and 36 degrees. That’s a lot more than the ~6 degrees estimated by Stevens and Parrish (1999, 2005). And, as we mentioned in the paper, it nicely reinforces the point made by Upchurch (2000), that flexibility in the anterior dorsals should be taken into account in estimating neck posture and ROM.

Figure 16. Dorsal vertebrae of Apatosaurus CM 11339. Articulated middle or posterior dorsal vertebrae of Apatosaurus CM 11339. A. X-ray scout image showing the two vertebrae in articulation, in left lateral view. B–D. Slices 39, 43 and and 70 in posterior view, showing the most anterior appearance of the condyles and cotyles.

Dorsal vertebrae of Apatosaurus CM 11339.
Articulated middle or posterior dorsal vertebrae of Apatosaurus CM 11339. A. X-ray scout image showing the two vertebrae in articulation, in left lateral view. B–D. Slices 39, 43 and and 70 in posterior view, showing the most anterior appearance of the condyles and cotyles. Taylor and Wedel (2013: figure 16).

Here’s our last specimen, CM 11339. No big surprises here, although if you ever had a hard time visualizing how hyposphenes and hypantra fit together, you can see them in articulation in parts C and D (near the top of the specimen). Once again, by paging through slices we were able to estimate the separation between the vertebrae. Incidentally, the condyle IS centered in the cotyle here, it just doesn’t look that way because the CT slice is at an angle to the joint–see the lateral scout in part A of the figure to see what I mean.

So, what did we find? In Sauroposeidon the spacing between C5 and C6 is 52mm. That’s pretty darn thick in absolute terms–a shade over two inches–but really thin in relative terms–only a little over 4% of the length of each vertebra. In both of the juvenile Apatosaurus specimens, the spacing between the vertebrae was about 14mm (give or take a few because of the inherent thickness of the slices; see the paper for details on these uncertainties).

Now, here’s an interesting thing: we can try to estimate the intervertebral spacing in an adult Apatosaurus in two ways–by scaling up from the juvenile apatosaurus, or by scaling sideways from Sauroposeidon (since a big Apatosaurus was in the same ballpark, size-wise)–and we get similar answers either way.

Scaling sideways from Sauroposeidon (I’m too lazy to write anymore so I’m just copying and pasting from  the paper):

Centrum shape is conventionally quantified by Elongation Index (EI), which is defined as the total centrum length divided by the dorsoventral height of the posterior articular surface. Sauroposeidon has proportionally very long vertebrae: the EI of C6 is 6.1. If instead it were 3, as in the mid-cervicals of Apatosaurus, the centrum length would be 600 mm. That 600 mm minus 67 mm for the cotyle would give a functional length of 533 mm, not 1153, and 52 mm of cartilage would account for 9.8% of the length of that segment.

Scaling up from the juveniles: juvenile sauropods have proportionally short cervicals (Wedel et al. 2000). The scanned vertebrae are anterior dorsals with an EI of about 1.5. Mid-cervical vertebrae of this specimen would have EIs about 2, so the same thickness of cartilage would give 12mm of cartilage and 80mm of bone per segment, or 15% cartilage per segment. Over ontogeny the mid-cervicals telescoped to achieve EIs of 2.3–3.3. Assuming the cartilage did not also telescope in length (i.e., didn’t get any thicker than it got taller or wider), the ratio of cartilage to bone would be 12:120 (120 from 80*1.5), so the cartilage would account for 10% of the length of the segment–almost exactly what we got from the based-on-Sauroposeidon estimate. So either we got lucky here with our tiny sample size and truckloads of assumptions, or–just maybe–we discovered a Thing. At least we can say that the intervertebral spacing in the Apatosaurus and Sauroposeidon vertebrae is about the same, once the effects of scaling and EI are removed.

Finally, we’re aware that our sample size here is tiny and heavily skewed toward juveniles. That’s because we were just collecting targets of opportunity. Finding sauropod vertebrae that will fit through a medical-grade CT scanner is not easy, and it’s just pure dumb luck that Kent Sanders and I had gotten scans of even this many articulated vertebrae way back when, since at the time we were on the hunt for pneumaticity, not intervertebral joints or their soft tissues. As Mike has said before, we don’t think of this paper as the last word on anything. It is, explicitly, exploratory. Hopefully in a few years we’ll be buried in new data on in-vivo intervertebral spacing in both extant and extinct animals. If and when that avalanche comes, we’ll just be happy to have tossed a snowball.

References