Somehow this seems to have slipped under the radar: National Science Foundation announces plan for comprehensive public access to research results. They put it up on 18 March, two whole months ago, so our apologies for not having said anything until now!

This is the NSF’s rather belated response to the OSTP memo on Open Access, back in January 2013. This memo required all Federal agencies that spend $100 million in research and development each year to develop OA policies, broadly in line with the existing one of the NIH which gave us PubMed Central. Various agencies have been turning up with policies, but for those of us in palaeo, the NSF’s the big one — I imagine it funds more palaeo research than all the others put together.

So far, so awesome. But what exactly is the new policy? The press release says papers must “be deposited in a public access compliant repository and be available for download, reading and analysis within one year of publication”, but says nothing about what repository should be used. It’s lamentable that a full year’s embargo has been allowed, but at least the publishers’ CHORUS land-grab hasn’t been allowed to hobble the whole thing.

There’s a bit more detail here, but again it’s oddly coy about where the open-access works will be placed: it just says they must be “deposited in a public access compliant repository designated by NSF”. The executive summary of the actual plan also refers only to “a designated repository”

Only in the full 31-page plan itself does the detail emerge. From page 5:

In the initial implementation, NSF has identified the Department of Energy’s PAGES (Public Access Gateway for Energy and Science) system as its designated repository and will require NSF-funded authors to upload a copy of their journal articles or juried conference paper to the DOE PAGES repository in the PDF/A format, an open, non-proprietary standard (ISO 19005-1:2005). Either the final accepted version or the version of record may be submitted. NSF’s award terms already require authors to make available copies of publications to the Cognizant Program Officers as part of the current reporting requirements. As described more fully in Sections 7.8 and 8.2, NSF will extend the current reporting system to enable automated compliance.

Future expansions, described in Section 7.3.1, may provide additional repository services. The capabilities offered by the PAGES system may also be augmented by services offered by third parties.

So what is good and bad about this?

Good. It makes sense to me that they’re re-using an existing system rather than wasting resources and increasing fragmentation by building one of their own.

Bad. It’s a real shame that they mandate the use of PDF, “the hamburger that we want to turn back into a cow”. It’s a terrible format for automated analysis, greatly inferior to the JATS XML format used by PubMed Central. I don’t understand this decision at all.

Then on page 9:

In the initial implementation, NSF has identified the DOE PAGES system to support managing journal articles and juried conference papers. In the future, NSF may add additional partners and repository services in a federated system.

I’m not sure where this points. In an ideal world, it would mean some kind of unifying structure between PAGES and PubMed Central and whatever other repositories the various agencies decide to use.

Anyone else have thoughts?

Update from Peter Suber, later that day

Over on Google+, Peter Suber comments on this post. With his permission, I reproduce his observations here:

My short take on the policy’s weaknesses:

  • will use Dept of Energy PAGES, which at least for DOE is a dark archive pointing to live versions at publisher web sites
  • plans to use CHORUS (p. 13) in addition to DOE PAGES
  • requires PDF
  • silent on open licensing
  • only mentions reuse for data (pp. v, 18), not articles, and only says it will explore reuse
  • silent on reuse for articles even tho it has a license (p. 10) authorizing reuse
  • silent on the timing of deposits

I agree with you that a 12 month embargo is too long. But that’s the White House recommended default. So I blame the White House for this, not NSF.

To be more precise, PAGES favors publisher-controlled OA in one way, and CHORUS does it in another way. Both decisions show the effect of publisher lobbying on the NSF, and its preference for OA editions hosted by publishers, not OA editions hosted by sites independent of publishers.

So all in all, the NSF policy is much less impressive than I’d initially thought and hoped.

In response to my post Copyright from the lens of reality and other rebuttals of his original post, Elseviers General Counsel Mark Seeley has provided a lengthy comment. Here’s my response (also posted as a comment on the original article, but I’m waiting for it to be moderated.)


Hi, Mark, thanks for engaging. You write:

With respect to the societal bargain, I would simply note that, in my view, the framers believed that by providing rights they would encourage creative works, and that this benefits society as a whole.

Here, at least, we are in complete agreement. Where we part company is that in my view the Eldred v. Ashcroft decision (essentially that copyright terms can be increased indefinitely) was a travesty of the original intent of copyright, and clearly intended for the benefit of copyright holders rather than that of society on general. (I further note in passing that those copyright holders are only rarely the creative people, but rights-holding corporations whose creative contribution is negligible.)

You continue:

[Journal] services and competencies need to be supported through a business model, however, and in the mixed economy that we have at the moment, this means that many journals will continue to need subscription and purchase models.

This is a circular argument. It comes down to “we use restrictive copyright on scholarly works at present, so we therefore need to continue to do so”. In fact, this this is not an argument at all, merely an assertion. If you want it to stick, you need to demonstrate that the present “mixed economy” is a good thing — something that is very far from evident.

The alternatives to a sound business model rooted in copyright are in my view unsustainable. I worry about government funding, patronage from foundations, or funding by selling t-shirts—I am not sure that these are viable, consistent or durable. Governments and foundations can change their priorities, for example.

If governments and foundations decide to stop funding research, we’re all screwed, and retention of copyright on the papers we’re no longer able to research and write will be the least of our problems. The reality is that virtually everyone in research is already dependent on governments and foundations for the 99% of their funding that covers all the work before the final step of publication. Taking the additional step of relying on those same sources for the last 1% of funding is eminently sensible.

On Creative Commons licences, I don’t think we have any material disagreement.

Now we come to the crucial question of copyright terms (already alluded to via Eldred v. Ashcroft above). You content:

Copyright law was most likely an important spur for the author or publisher to produce and distribute the work [that is now in the public domain] in the first place.

In principle, I agree — as of course did the framers of the US Constitution and other lawmakers that have passed copyright laws. But as you will well know, the US’s original copyright act of 1790, which stated its purpose as “encouragement of learning”, offered a term of 14 years, with an optional renewal of a further 14 years if the author was still alive at the end of the initial term. This 14-year was considered quite sufficient to incentivise the creation of new works. The intent of the present law seems to be that authors who have been dead for 70 years still need to receive royalties for their works, and in the absence of such royalties would not have created in the first place. This is self-evident nonsense. No author in the history of the world every said “I would have written a novel if I’d continued to receive royalties until 70 years after my death, but since royalties will only last 28 years I’m not going to bother”.

But — and this can’t be stated strongly enough — even if there were some justification for the present ridiculous copyright terms in the area of creative works, it would still say nothing whatsoever about the need to copyright scientific writing. No scientific researcher ever wrote a paper who would not have written it in the absence of copyright. That’s what we’re talking about here. One of the tragedies of copyright is that it’s been extruded from a domain where it has some legitimate purpose into a domain where it has none.

The Budapest Open Access Initiative said it best and most clearly: “the only role for copyright in this domain [scholarly research] should be to give authors control over the integrity of their work and the right to be properly acknowledged and cited“. (And several of the BOAI signatories have expressed regret over even the controlling-integrity-of-the-work part of this.)


See also David Roberts’ response to Seeley’s posting.

This post is a response to Copyright from the lens of a lawyer (and poet), posted a couple of days ago by Elsevier’s General Counsel, Mark Seeley. Yes, I am a slave to SIWOTI syndrome. No, I shouldn’t be wasting my time responding to this. Yes, I ought to be working on that exciting new manuscript that we SV-POW!er Rangers have up and running. But but but … I can’t just let this go.


Copyright from the lens of a lawyer (and poet) is a defence of Elsevier’s practice of having copyright encumber scientific publishing. I tried to read it in the name of fairness. It didn’t go well. The very first sentence is wrong:

It is often said that copyright law is about a balance of interests and communities, creators and users, and ultimately society as a whole.

No. Copyright is not a balance between competing interests; it’s a bargain that society makes. We, the people, give up some rights in exchange for incentivising creative people to make new work, because that new work is of value to society. To quote the US constitution’s helpful clause, copyrights exist “To promote the Progress of Science and useful Arts” — not for authors, but for wider society. And certainly not of publishers who coerce authors to donate copyright!

(To be fair to Seeley, he did hedge by writing “It is often said that copyright law is about a balance”. That is technically true. It is often said; it’s just wrong.)

Well, that’s three paragraphs on the first sentence of Elsevier’s defence of copyright. I suppose I’d better move on.

The STM journal publishing sector is constantly adjusting to find the right balance between researcher needs and the journal business model, as refracted through copyright.

Wrong wrong wrong. We don’t look for a balance between researchers needs (i.e. science) and the journal business model. Journals are there to serve science. That’s what they’re for.

Then we have the quote from Mark Fischer:

I submit that society benefits when the best creative spirits can be full-time creators and not part-timers doing whatever else (other than writing, composing, painting, etc.) they have to do to pay the rent.

This may be true. But it is totally irrelevant to scholarly copyright. That should hardly need pointing out, but here it is for those hard of thinking. Scholars make no money from the copyright in the work they do, because (under the Elsevier model) they hand that copyright over to the publisher. Their living comes in the form of grants and salaries, not royalties.

Ready for the next one?

The alternatives to a copyright-based market for published works and other creative works are based on near-medieval concepts of patronage, government subsidy […]

Woah! Governments subsidising research and publication is “near-medieval”? And there we were thinking it was by far the most widespread model. Silly us. We were all near-medieval all this time.

Someone please tell me this is a joke.

Moving swiftly on …

Loud advocates for “copyright reform” suggest that the copyright industries have too much power […] My comparatively contrarian view is that this ignores the enormous creative efforts and societal benefits that arise from authoring and producing the original creative work in the first place: works that identify and enable key scientific discoveries, medical treatments, profound insights, and emotionally powerful narratives and musical experiences.

Wait, wait. Are we now saying that … uh, the only reason we get scientific discoveries and medical treatment because … er … because of copyright? Is that it? That can’t be it. Can it?

Copyright has no role in enabling this. None.

In fact, it’s worse than that. The only role of copyright in modern scholarly publishing is to prevent societal benefits arising from scientific and medical research.

The article then wanders off into an (admittedly interesting) history of Seeley’s background as a poet, and as a publisher of literary magazines. The conclusion of this section is:

Of course creators and scientists want visibility […] At the very least, they’d like to see some benefit and support from their work. Copyright law is a way of helping make that happen.

This article continues to baffle. The argument, if you want to dignify it with that name, seems to be:

  • poets like copyright
  • => we copyright other people’s science
  • => … profit!

Well, that was incoherent. But never mind: finally we come to part of the article that makes sense:

  • There is the “idea-expression” dichotomy — that copyright protects expression but not the fundamental ideas expressed in a copyright work.

This is correct, of course. That shouldn’t be cause for comment, coming from a copyright lawyer, but the point needs to be made because the last time an Elsevier lawyer blogged, she confused plagiarism with copyright violation. So in that respect, this new blog is a step forward.

But then the article takes a sudden left turn:

The question of the appropriateness of copyright, or “authors’ rights,” in the academic field, particularly with respect to research journal articles, is sometimes controversial. In a way quite similar to poets, avant-garde literary writers and, for that matter, legal scholars, research academics do not rely directly on income from their journal article publishing.

Er, wait, what? So you admit that scholarly authors do not benefit from copyright in their articles? We all agree, then, do we? Then … what was the first half of the article supposed to be about?

And in light of this, what on earth are we to make of this:

There is sometimes a simplistic “repugnance” about the core publishing concept that journal publishers request rights from authors and in return sell or license those rights to journal subscribers or article purchasers.

Seeley got that much right! (Apart from the mystifyingly snide use of “simplistic” and the inexplicable scare-quotes.) The question is why he considers this remotely surprising. Why would anyone not find such a system repugnant? (That was a rhetorical question, but here’s the answer anyway: because they make a massive profit from it. That is the only reason.)

Well, we’re into the final stretch. The last paragraph

Some of the criticism of the involvement of commercial publishing and academic research is simply prejudice, in my view;

Yes. Some of us are irrationally prejudiced against a system where, having laboriously created new knowledge, it’s then locked up behind a paywall. It’s like the irrational prejudice some coal-miners have against the idea of the coal they dig up being immediately buried again.

And finally, this:

Some members of the academic community […] base their criticism on idealism.

Isn’t that odd? I have never understood why some people consider “idealism” to be a criticism. I accept it as high praise. People who are not idealists have nothing to base their pragmatism on. They are pragmatic, sure, but to what end?

So what are we left with? What is Seeley’s article actually about? It’s very hard to pick out a coherent thread. If there is one, it seems to be this: copyright is helpful for some artists, so it follows that scholarly authors should donate their copyright to for-profit publishers. That is a consequence that, to my mind, does not follow particularly naturally from the hypothesis.

While Mike’s been off having fun at the Royal Society, this has been happening:

Lots of feathers flying right now over the situation at the Medical Journal of Australia (MJA). The short, short version is that AMPCo, the company that publishes MJA, made plans to outsource production of the journal, and apparently some sub-editing and administrative functions as well, to Elsevier. MJA’s editor-in-chief, Professor Stephen Leeder, raised concerns about the journal getting involved with one of the most ethically problematic publishing companies in existence. And also about this having been done without consultation.

He was sacked for his trouble.

After Leeder was pushed out, his job was offered to MJA’s deputy editor, Tania Janusic. She declined, and resigned from the journal, as did 19 of the 20 members of the journal’s editorial advisory committee. (Some accounts say 18. Anyway, 90%+ of the committee is gone.)

When we first discussed the situation via email, Mike wrote, “My take is that at the present stage of the OA transition, editorial board resignations from journals controlled by predatory legacy publishers are about the most important visible steps that can be taken. Very good news for the world, even though it must be a mighty pain for the people involved.”

Yes. I feel pretty bad for the people involved, but I’m hugely supportive of what they’re doing.

I don’t know what we can do to materially contribute here, beyond amplifying the signal and lending our public support to Leeder, Janusic, and the 19 editors who resigned. That’s a courageous thing to do, but no-one should have to do it. The sooner we move to a world where scientific results and other forms of scholarly publication are freely available to all, instead of under the monopolistic control of a handful of exploitative, hugely profitable corporations, the better.

A short list of links, nowhere near exhaustive, if you’d like to read more:

UPDATE: In the first comment below, Alex Holcombe pointed us to this post written by Leeder himself, explaining the reasoning and consequences of his decision.

Also, dunno how I forgot this – if you haven’t already, you might be interested in signing the Cost of Knowledge boycott against Elsevier. Here’s the link.

Peggy Sue's Diner-saurs - London with sauropod

A couple of weekends ago, London and I went camping and stargazing at Afton Canyon, a nice dark spot about 40 miles east of Barstow. On the way home, we took the exit off I-15 at Ghost Town Road, initially because we wanted to visit the old Calico Ghost Town. But then we saw big metal dinosaurs south of the highway, and that’s how we came to Peggy Sue’s Diner and in particular the Diner-saur Park.

Peggy Sue's Diner-saurs - spinosaur

The Diner-saur Park is out behind the diner and admission is free. There are pools with red-eared sliders, paved walkways, grass, trees, a small gift shop, and dinosaurs. Here’s a Spinosauruscuriously popular in the Mojave Desert, those spinosaurs.

Peggy Sue's Diner-saurs - stegosaur

Ornithischians are represented by two stegosaurs, this big metal one and a smaller concrete one under a tree.

Peggy Sue's Diner-saurs - turtles

The turtles are entertaining. They paddle around placidly and crawl out to bask on the banks of the pools, and on little islands in the centers.

Peggy Sue's Diner-saurs - sign

The gift shop is tiny and the selection of paleo paraphernalia is not going to blow away any hard-core dinophiles. But it is not without its charm. And, hey, when you find a dinosaur gift shop in the middle of nowhere, you don’t quibble about size. London got some little plastic turtles and I got some cheap and horribly inaccurate plastic dinosaur skeletons to make a NecroDinoMechaLaser Squad for our Dinosaur Island D&D campaign.

Now, about that sauropod. The identification sign on the side of the gift shop notwithstanding, this is not a Brachiosaurus. With the short forelimbs and big back end, this is clearly a diplodocid. The neck is too skinny for Apatosaurus or the newly-resurrected Brontosaurus, and too long for Diplodocus. I lean toward Barosaurus, although I noticed in going back through these photos that with the mostly-straight, roughly-45-degree-angle neck, it is doing a good impression of the Supersaurus from my 2012 dinosaur nerve paper. Compare this:

Peggy Sue's Diner-saurs - sauropod 1

to this:

Wedel RLN fig1 - revised

If I had noticed it sooner, I would have maneuvered for a better, more comparable shot.

Guess I’ll just have to go back.


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

This abomination — a proposal for a “UK National Licence” for open-access papers, making the available only in the UK, is not an April Fool joke. It’s a serious proposal, put forward by HEPI, the Higher Education Policy Institute, which styles itself “the UK’s only independent think tank devoted to higher education” (though I note without comment that they routinely partner with Elsevier).

It’s desperately disappointing that British academics should propose something as small-minded and xenophobic as this, which I can only refer to as the UKIP Licence.  Let’s start counting some of the ways this is a terrible, terrible idea.

1. It’s not open access by any existing definition of the term. For example, the Budapest Open Access Initiative, which first coined the term, describes OA as “free availability on the public internet” (i.e. not a subnet), “permitting any users” (i.e. not just British users) “without financial, legal, or technical barriers” (i.e. no filtering on IP addresses).

2. It positions the UK as the one country in the world willing to poison the open-access well, prepared to destroy value for 199 countries in the hope of increasing it for one. This makes it a classic prisoner’s-dilemma “defect” strategy — an approach which has been shown by multi-algorithm tournaments to reliably downgrade the defector’s outcome.

3. British people gain more when 200 countries are working on advances in health, education, etc., than when only one is. This tiny-minded licence, if adopted, would hobble British innovation, health and education, as well as that of the rest of the world.

4. Most important, it’s mean. We have to be better than this. Publishing research about diseases that kill millions in third-world countries, then preventing scientists in those countries from reading that research is not just stupid, it’s despicable. It’s hard to imagine behaviour more unrepresentative of the values that we like to imagine the UK embodies.

Oh, and 5. it won’t work, of course. Barring access by IP address is a notoriously flawed approach, which hides content from Brits abroad while allowing access to anyone anywhere who knows how to use a Web proxy.

Putting it all together, this is about the most misguided proposal imaginable. I would like to see its authors, both of them senior at UCL, withdraw it with all possible haste, and with an appropriate apology.

[I would have left this comment on HEPI’s blog-post announcing their proposal,  but comments are turned off — perhaps not surprisingly. I did leave a version of it on the Times Higher Education article about this.]


Update the next day: see also David Kernohan’s post A local licence for Henbury.

Update 9th April: this post, lightly modified, is published as a letter in today’s Times Higher Education. More importantly, you should all go and read Stephen Curry’s much more dispassionate, but equally critical, analysis of the National Licence proposal.

Kraatz et al 2015 Figure 1 - rabbit skull freak gallery

Meet some of my new friends: (A) Brachylagus idahoensis, (B) Lepus capensis, (C) Poelagus marjorita, (D) Pronolagus crassicaudatus, (E) Lepus americanus, (F) Oryctolagus cuniculus, (G) Nesolagus timminsi, (H) Bunolagus monticularis, and (I) Romerolagus diazi. Kraatz et al. (2015: fig. 1).

I have a new paper out today in PeerJ: “Ecological correlates to cranial morphology in leporids (Mammalia, Lagomorpha)”, with coauthors Brian Kraatz, Emma Sherratt, and Nick Bumacod. Get it free here.

I know, I know, I have fallen from grace. First Aquilops, now rabbits. And, and…skulls! I know what you’re thinking: that maybe I’m not just experimenting with the non-vertebrae of non-sauropods anymore – maybe I have an actual problem. But I don’t. I can quit anytime! You’ll see.

Actually rabbits are the freakiest of all mammals and their skulls are wicked cool. They have double incisors, with the second set right behind the first, hence the name Duplicidentata for rabbits and their close relatives. They have weird fenestrations in their maxillae (pretty much all taxa) and parietal and occipital bones (some more than others) – I’ll come back to that in a bit. And, as we discuss in our new paper, you can tell something about how a rabbit runs by looking at its skull. I thought it would be fun to relate how we figured that out, and why.

A long time ago in a graduate seminar far, far away…

1950: DuBrul, Laskin, and Moss

I met Brian Kraatz at Berkeley, where he and I were part of the cohort of students that came into the Integrative Biology Department in the fall of 2001 (faithful readers may remember Brian from his work tracking oliphaunts from, gosh, three years ago already). We took a lot of classes together, including a seminar by Marvalee Wake on evolutionary morphology. I’m pretty sure that seminar was the first time I’d actually read DuBrul and Laskin (1961), “Preadaptive potentialities of the mammalian skull: an experiment in growth and form”, or as I think of it, “How to turn a rat skull into a pika skull for fun and profit.”

Pikas (Ochotonidae) are the sister group to rabbits (Leporidae) and together these groups make up crown Lagomorpha. If you’re not familiar with pikas, Brian describes them as starting with bunny rabbits and then making them even cuter and cuddlier. Seriously, go do an image search for ‘pika’ and try not to die of cute overload.

Pikas are interesting because in many ways their skulls are intermediate between those of rodents, especially rats, and rabbits. This is maybe not surprising since rodents are the sister group to lagomorphs and are united with them in the clade Glires. E. Lloyd DuBrul was all over this rat-pika-rabbit thing back in the mid-twentieth century. Here’s an illustration from DuBrul (1950: plate 2; labels added by me):

Rattus Ochotona and Lepus skulls compared - DuBrul 1960 plate 2

So DuBrul knew from pikas and in particular he had the idea that you could maybe just tweak a rat skull – say, by knocking out the basicranial sutures in a baby rat to limit the growth of the skull base – and produce a gently domed skull like that of a pika. That’s what DuBrul and Laskin (1961) is all about. They did that experiment and here are their results (DuBrul and Laskin (1961: plate 3). Normal rat skull on the right, and dotted in the bottom diagram; experimental “pika-morph” rat skull on the left, and solidly outlined below.

Experimental skull doming in rats - DuBrul and Laskin 1961 plate 3

What’s going on here morphogenetically is that the facial skeleton is getting tilted down and away from the back end of the skull. DuBrul was hip to that, too – here’s a relevant image from his 1950 paper (plate 4; labels added by me):

Skull tilting in Rattus Ochotona and Lepus - DuBrul 1960 plate 4

The common reference point against which these skulls are registered is the cranial base (the floor of the braincase just forward of the foramen magnum). Again, the pika is a pretty good intermediate between the rat and a ‘normal’ rabbit, and the dang-near-dog-sized Flemish Giant rabbit takes the lagomorph face-tilting thing to its extreme. (‘Flemish Giant rabbit’ is another entertaining image search that I will leave you as homework.)

Turns out there’s another way you can get rat skulls with different geometries: you can cut off their legs and make them walk on two feet. In an experiment that you might have trouble getting past an Institutional Animal Care and Use Committee today, Moss (1961) lopped off the forefeet or hindfeet in two experimental batches of rats, to see what effect this would have on their skulls. I’ll let Moss speak for himself on this one (Moss, 1961: pp. 301-303, emphasis in the original):

Circumnatal amputation of the forelimbs has successfully produced what are in essence “bipedal rats,” i.e., rats whose habitual mode of kinetic and static posture is permanently altered. […] The animals never became bipedal in the exact sense; that is, they never walked erect on two limbs at all times. […] Nevertheless, bipedal posture and motion were more frequently observed than in controls. […]

Animals whose hind limbs were removed represented another picture. They most certainly did not walk about on their intact forelimbs. Neither did they seem able to use their hind limb stumps as satisfactory substitutes. Their gait was not uniform and seemed to consist in a series of short pushes or hops. The most noticeable thing about them was, among other things, apparent accentuation of their cervical vertebral curvature. The sum of these changes was an upward rotation of the skull.

He wasn’t kidding: when the two groups of bipedal rats grew up, their facial skeletons were tilted relative to the control group, but in different directions (Moss, 1961: fig 3; ‘fore’ and ‘hind’ refer to which limbs the animals had left to locomote with):

Skull deformation in bipedal rats - Moss 1961 fig 3

Brian and I read Moss back at Berkeley, too. In fact, we were minor Moss junkies. If you’re interested in how living forms come into being, you owe it to yourself to read Moss (1968), “A theoretical analysis of the functional matrix”.

The upshot of all of this is that although neither Brian nor I had done anything with our deep (and, okay, deeply weird) knowledge of how to experimentally jack up rat skulls by the time we graduated from Berkeley, we were also primed to be thinking about how skulls attain their shapes – especially the skulls of rodents and rabbits.

2009: American Museum of Natural History

I went to the AMNH in February, 2009, to visit Brian, who was on a postdoc there at the time, and to spend one day looking at sauropods with Mike, who was over from England for a conference. What Brian and I planned to work on was the fenestration of rabbit skulls, because I’m always interested in the strategic loss of bone from skeletal structures. We spent probably half a day talking about that, and I filled a whole page in my notebook with related noodlings:

AMNH rabbit skull sketch 1

But as the sketch on the right shows, it didn’t take us long to figure out that there was something even more interesting to do with rabbit skulls. Brian had a whole shedload of rabbit skulls from different taxa sitting on his desk, and we noticed pretty quickly that one of the primary ways they varied was in the tilt of the facial skeleton relative to the back of the skull. Here’s the very next page of my notes from that trip:


The skull up top belongs to Caprolagus, the Hispid hare, which I tend to think of as the “bulldozer hare”. Seriously, it looks like a tank. It doesn’t bound or even hop, it scrambles. Here, stare into the abyss:

Caprolagus from ARKive

That rabbit will cut you, man. And just look at how flat its skull is. Even in life Caprolagus looks more rodent-y than rabbit-y. Or, more precisely, more Ochotona-y.

At the the other extreme are taxa like Bunolagus and Pronolagus, which really push the “I’m going to cute you to death by dint of my incredible bunnosity” thing:

Bunolagus from ARKive

As Brian and I started going through skulls of as many extant rabbits as we could, we noticed that the flatter-skulled taxa, with less pronounced facial tilt, tended to be the stolid, foursquare scramblers like Caprolagus, whereas the speed demons tended to have more strongly tilted skulls. It also seemed like the latter group were achieving that pronounced facial tilt by changing the geometry of the occipital region of the skull. Look back up at the red quadrilaterals I drew on the Caprolagus and Bunolagus skulls in my notebook – those mark the basioccipital ventrally and the dorsal exposure of the supraoccipital. Perhaps unsurprisingly, supraoccipital length is not the whole story; it turns out that some face-tilters get that way by having longer or more strongly arched parietals, BUT it remains true that if you find a rabbit skull with a long dorsal exposure of the supraoccipital, it will also have pronounced facial tilt.

ANYWAY, by my last night in New York, Brian and I decided that the best way to attack this would be to go down to the basement and stay up most of the night drinking beer and measuring rabbit skulls. We then tried to correlate the various measurements and angles with information on the locomotor and burrowing habits of each species. That was a big job, and after a couple of years with little forward progress (to be fair, Brian was moving across the country and taking his first tenure-track job in this interval, and I was helping birth a sauropod) we brought in Brian’s graduate student, Nick Bumacod, to do most of it. Later on the three of us were forced to acknowledge that we knew enough statistics to get ourselves into trouble but not enough to get back out. Brian had taken a geometric morphometrics course for which Emma Sherratt was a TA, and he started bugging her for help with the stats. Emma has been involved in writing new software packages for R, and we realized that the paper would be a lot stronger if we just brought her on as an author and gave her free rein with the data. Along the way Brian and Nick were giving presentations on the project everywhere from the local Western Area Vert Paleo meeting to the World Lagomorph Conference in Vienna. I got my name on four abstracts along the way, which I think is record abstract-to-paper ratio for me (especially considering that 90% of my effort on the paper was invested in a single evening in 2009 over a couple of six-packs).

But enough navel-gazing, what did we find?

2015: Rabbit skulls reveal their mode of locomotion

Our results, which you can read for free, support the hunch that Brian and I had back in 2009: slow-moving rabbits that locomote by scrambling or scampering instead of hopping tend to have less facial tilt, and faster-moving saltatorial (hopping) and cursorial (leaping and bounding) rabbits have more facial tilt. Interestingly, facial tilt does not distinguish the saltators from the cursors. So the break here is between scrambling and any kind of hopping or leaping, but not between hoppers and leapers.

Kraatz et al 2015 fig 5a

Kraatz et al. (2015: fig. 5a)

Why would that be so? We don’t know for sure yet, but our top hypothesis is that if you’re moving fast, it pays to see the ground in front of you more clearly, and getting your nose down out of the way probably helps with that. This is pretty similar to the hypothesis that tyrannosaurs have pinched nasals for better binocular vision (Stevens, 2006). Rabbits are prey animals and probably can’t afford to point their eyes forward, and they may need wide nasal airways as air intakes while they’re sprinting. Tilting the nose down may be the next best thing.

Guinea pig and mara skulls - DuBrul 1960 plate 6

Some circumstantial support for this comes from the Caviidae, the family of South American rodents that includes guinea pigs, cavies, maras, and capybaras. Here’s another plate from DuBrul (1950: plate 6) contrasting the flatter skull of the guinea pig (Cavia porcellus, top) with the decidedly arched skull of the mara or Patagonian hare (Dolichotis magellanica, bottom). Compare the mara skull to the sectioned rabbit skull in the other DuBrul plate, above – there aren’t a lot of obvious characters to separate the two (beyond the lack of double incisors in the mara).

Mara photo from Wikipedia

Mara photo from Wikipedia

Despite being commonly referred to as ‘hares’ and looking a lot like short-eared rabbits, maras are rodents that evolved their rabbit-like form independently. The acquisition of pronounced facial tilt in two separate lineages of small fast-moving herbivorous mammals is further evidence for the influence of locomotor mode on skull form. Irritatingly, I think we neglected to mention the guinea pig : mara :: pika : rabbit correspondence in the paper. Oh well, it wasn’t our novel observation, and we did cite DuBrul (1950).

Kraatz et al 2015 Figure 4 - skull measurements

Relevant to the next paragraph: DILU is ‘diastema length upper’ and BLD is ‘bulla diameter’. Kraatz et al. (2015: fig. 4).

We found lots of other interesting things, too. The PCA plots we produced from our data separate the living rabbits in unexpected ways. The length of the diastema (the toothless portion of the upper jaw) and the diameter of the auditory bulla seem to be particularly important. Diastema length isn’t too hard to figure out – most of the face-tilters have long diastemas, and the flat-heads tend to have short ones. We have no idea what bulla diameter means yet. I mean, obviously something to do with hearing, but we don’t have any ecological variables in our analysis to address that because we didn’t see it coming. So there’s a chunk of new science waiting to be done there.

Speaking of new science, or at least a relatively new thing in science, we published the full peer-review history alongside the paper, just as Mike and I did back in 2013 and as Mike did with his stand-alone paper last December. More than 80% of PeerJ authors elect to publish the peer review histories for their papers. I can’t wait until it’s 100%. PeerJ reviews are citeable – each one gets a DOI and instructions on how to cite it – and I’m tired of having my effort as a peer reviewer used once and then thrown away forever.

If you’ve been reading this whole post with gritted teeth, wondering why we were using linear measurements instead of geometric morphometrics, chillax. Brian and Emma are on that. They’ve been CT scanning the skulls of as many extant rabbits as possible and plotting landmarks for 3D morphometrics – if you were at SVP last fall, you may have seen their talk (Kraatz and Sherratt, 2014). So stay tuned for what will soon be a new ongoing series, Rabbit Skulls: The Next Generation.

I probably won’t be on that voyage. I’ve had fun getting acquainted with a completely different part of the tree of life, but there are an awful lot of shards of excellence – busted-up sauropod vertebrae, that is – crying out for my attention, and I need to stop neglecting them. I’m done with rabbit skulls, I promise. I’m going clean. (Wish me luck!)


  • DuBrul, E. L. (1950). Posture, locomotion and the skull in Lagomorpha. American Journal of Anatomy, 87(2), 277-313.
  • DuBrul, E. L., & Laskin, D. M. (1961). Preadaptive potentialities of the mammalian skull: an experiment in growth and form. American Journal of Anatomy, 109(2), 117-132.
  • Kraatz, B., and Sherratt, E. (2014). Evolution, ecology, and modularity of the lagomorph skull. Journal of Vertebrate Paleontology, 35(3, Supplement), 162A.
  • Kraatz, B.P., Sherratt, E., Bumacod, N., and Wedel, M.J. 2015. Ecological correlates to cranial morphology in leporids (Mammalia, Lagomorpha). PeerJ3:e844.
  • Moss, M. L. (1961). Rotation of the otic capsule in bipedal rats. American Journal of Physical Anthropology, 19(3), 301-307.
  • Moss, M. L. (1968). A theoretical analysis of the functional matrix. Acta Biotheoretica, 18(1), 195-202.
  • Stevens, K. A. (2006). Binocular vision in theropod dinosaurs. Journal of Vertebrate Paleontology, 26(2), 321-330.

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