Rabbit skulls strike back: Kraatz and Sherratt (2016)
September 22, 2016

Facial tilt in extant leporids is strongly correlated with locomotor mode – fast movers have more strongly tilted faces. There’s a lot of homoplasy, which is to be expected with characters that are strongly driven by current function. Kraatz and Sherratt (2016: fig. 5).
Judgmental readers will recall that I have dabbled in mammal skulls, thanks to the corrupting influence of my friend and colleague, Brian Kraatz. At the end of my last post on this sordid topic, I mentioned that Brian and Emma Sherratt were working on a version 2.0 based in 3D morphometrics. The first volley from that project was published today in PeerJ.
Happily for all of us, Brian and Em confirmed the relationship between facial tilt and locomotor mode that we first documented last year, using more taxa, more landmarks, and two more dimensions (Kraatz and Sherratt 2016: 12):
…in accordance with previous findings by Kraatz et al. (2015), facial tilt angle is correlated with locomotor mode (D-PGLS, F(2,17) = 11.13, P = 0.003), where lower facial tilt angle, meaning more pronounced cranial flexion, is found in cursorial species, and high angles are found in generalist species.
That’s just the most personally relevant tip of a very large, multifaceted iceberg, including a monster supplementary info package on FigShare with, among other things, 3D models of bunny skulls. It’s all free and awesome, so go have fun.

That homoplastic pattern shown in figure 5, above? It’s been going on for a while. I’m gonna go out on a limb and guess that Hypolagus was a rocket. Kraatz and Sherratt (2016: fig. 7).
References
Should we still give taxonomic authorities?
September 18, 2016
I have before me the reviews for a submission of mine, and the handling editor has provided an additional stipulation:
Authority and date should be provided for each species-level taxon at first mention. Please ensure that the nominal authority is also included in the reference list.
In other words, the first time I mention Diplodocus, I should say “Diplodocus Marsh 1878″; and I should add the corresponding reference to my bibliography.

Marsh (1878: plate VIII in part). The only illustration of Diplodocus material in the paper that named the genus.
What do we think about this?
I used to do this religiously in my early papers, just because it was the done thing. But then I started to think about it. To my mind, it used to make a certain amount of sense 30 years ago. But surely in 2016, if anyone wants to know about the taxonomic history of Diplodocus, they’re going to go straight to Wikipedia?
I’m also not sure what the value is in providing the minimal taxonomic-authority information rather then, say, morphological information. Anyone who wants to know what Diplodocus is would be much better to go to Hatcher 1901, so wouldn’t we serve readers better if we referred to “Diplodocus (Hatcher 1901)”
Now that I come to think of it, I included “Giving the taxonomic authority after first use of each formal name” in my list of
Idiot things that we we do in our papers out of sheer habit three and a half years ago.
Should I just shrug and do this pointless busywork to satisfy the handling editor? Or should I simply refuse to waste my time adding information that will be of no use to anyone?
References
- Hatcher, Jonathan B. 1901. Diplodocus (Marsh): its osteology, taxonomy and probable habits, with a restoration of the skeleton. Memoirs of the Carnegie Museum 1:1-63 and plates I-XIII.
- Marsh, O. C. 1878. Principal characters of American Jurassic dinosaurs, Part I. American Journal of Science, series 3 16:411-416.
What is the nature and purpose of a type specimen?
September 13, 2016
It’s been interesting seeing the response to my comment on the ICZN petition to establish Diplodocus carnegii as the replacement type species of the genus Diplodocus. In particular, Mickey Mortimer’s opposition to the petition seems to be based primarily on this argument:
The dinosaur community has recently lost sight of the fact that the type concept was never meant to indicate the most well preserved or described specimen/species.
I find this unconvincing, on the basis that the ICZN was never designed with dinosaurs in mind in the first place. For the great majority of the species that have been named under its rules, the selection of the obvious holotype has been perfectly adequate, because extant animals — by far the majority — are nearly all represented by complete and well-preserved specimens.

Alphina nigrosignata (type specimen; photo courtesy Geert Goemans, Department of Ecology and Evolutionary Biology, The University of Connecticut; specimen loaned from Herbert Zettel from The Museum of Natural History Vienna (NHMV). From University of Delaware, College of Agriculture & Natural Resources.
Dinosaurs — which in many cases are represented by eroded and distorted fossils of a tiny part of the animal — are already an aberration from the perspective of the ICZN, and that is why they sometimes need special treatment.
What are type specimens for, after all? The Code itself says “The fixation of the name-bearing type of a nominal taxon provides the objective standard of reference for the application of the name it bears” (Article 61.1); and comments that type specimens “are the international standards of reference that provide objectivity in zoological nomenclature” (Article 72.10). That is a role that YPM 1920 is simply not capable of fulfilling — and, more to the point, a role that it is not filling. The Diplodocus carnegii holotype CM 84 is the international standard of reference that provides objectivity in Diplodocus nomenclature. Slavishly following the usual provisions of the Code to retain the fiction that YPM 1920 fulfils this role simply does not reflect reality.
Some people occasionally object to the nomination of neotype specimens or replacement type species on the grounds that the Code does not require this. Of course it doesn’t: if it did, there would be no need for petitions. The fact that the Code allows for petitions constitutes explicit recognition that its usual provisions do not always suffice to produce the “sense and stability for animal names” that the Commission’s web-site used to have as its banner before the last redesign. Petitions exist precisely to allow the setting aside of the usual rules when sense and stability is served by doing do.
My comment in support of the Diplodocus carnegii ICZN petition
September 8, 2016
If you keep an eye on the wacky world of zoological nomenclature, you’ll know that earlier this year Emanuel Tschopp and Octávio Mateus published a petition to the International Commission on Zoological Nomemclature, asking them to establish Diplodocus carnegii, represented by the ubiquitous and nearly complete skeleton CM 84, as the type species of Diplodocus.
That is because Marsh’s (1878) type species, YPM 1920, is a pair of non-diagnostic mid-caudals which no-one has paid any attention to since 1901:

Tschopp and Mateus (2016: fig. 1). More anterior of the only two reasonably complete caudal vertebrae of the type specimen of Diplodocus longus (YPM 1920) in dorsal (A), anterior (B), left (C), posterior (D), right (E), and ventral (F) views. The neural spine is lost. The estimated position within the caudal column is caudal vertebra 17â24. Note the transverse ridge between the prezygapophyses shared with AMNH 223 (1).
I have now submitted a formal comment to the ICZN in support of the petition, in which I argue:
In its use as the definitive exemplar of the genus Diplodocus, as the foundation for numerous palaeobiological studies of the genus, and as the specifier for numerous important clades, the species D. carnegii is already effectively functioning as the type species of Diplodocus. Therefore the petition of Tschopp and Mateus (2016) requests only that the commission recognises de jure what is already the case de facto.
Anyone else who has strong feelings either in favour of or against the establishment of D. carnegii as a replacement type species for Diplodocus is welcome to submit their own comment to the ICZN. (I know of at least one person who has submitted a comment opposing the petition.)
The procedure is straightforward: just write your comment and email it to the Commision at iczn@nhm.ac.uk. (But it’s best also to copy your email to iczn@nus.edu.sg, as that seems to be where the ICZN is operating out of now: it took the NHM address four days to reply to my initial inquiry, but the Singaporean address responds quickly.)
References
- Marsh, O.C. 1878. Principal characters of American Jurassic dinosaurs, Part I. American Journal of Science (series 3) 16:411–416.
- Tschopp, Emanuel, and Octávio Mateus. 2016. Case 3700: Diplodocus Marsh, 1878 (Dinosauria, Sauropoda): proposed designation of D. carnegii Hatcher, 1901 as the type species. Bulletin of Zoological Nomenclature 73(1):17-24.
The Hallett and Wedel sauropod book is out now
September 2, 2016
I came home from SVPCA to find a heavy box waiting for me. It had my author’s copies of the book. I figured maybe they had gone out in advance of wide release, but nope, the book is shipping right now. This is welcome but unexpected. We only got the final files in a little over three months ago. I have no idea what alchemy the folks at Johns Hopkins University Press worked, to get the book out so fast, but I’m grateful.
We’ll have a book signing at SVP, at one in the LA area with both of us, and probably some local ones in Oregon and SoCal with just one author. I’ll announce those when we get them set up.
I have about a zillion people I need to thank, and I’ll probably do a FAQ about the book soon, but for now: it’s out. JHUP page. Amazon page.