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!)

References

  • 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.  https://dx.doi.org/10.7717/peerj.844
  • 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.

In 2012, Matt and I spent a week in New York, mostly working at the AMNH on Apatosaurusminimus and a few other specimens that caught our eye. But we were able to spend a day at the Yale Peabody Museum up in New Haven, Connecticut, to check out the caudal pneumaticity in the mounted Apatosaurus (= “Brontosaurus“) excelsus, YPM 1980, and the bizarrely broad cervicals of the Barosaurus lentus holotype YPM 429.

While we there, it would have been churlish not to pay some attention to the glorious and justly famous Age of Reptiles mural, painted by Rudolph F. Zallinger from 1944-1947.

So here it is, with the Brontosaurus neck for scale:

IMG_0501-zallinger-mural

Click through for high resolution (3552 × 2664).

And here is a close-up of the most important, charismatic, part of the mural:

IMG_0500-zallinger-mural

Again, click through for high resolution (3552 × 2664).

That’s your lot for now. We’ve long promised a proper photo post of the Brontosaurus mount itself, and I’ll try to get that done soon. For now, it’s just scenery.

Just launched: a new open-access journal of vertebrate paleontology, brought to you by the University of Alberta, Canada! It’s called VAMP (Vertebrate Anatomy Morphology Palaeontology), and it charges no APC. Here’s a illustration from one of the two articles in its first issue.

Holmes (2104:fig 12A). Synsacrum and pelvis of Chasmosaurus belli (ROM 843) in dorsal view.

Holmes (2014:fig 12A). Synsacrum and pelvis of Chasmosaurus belli (ROM 843) in dorsal view.

VAMP uses the canonical open-access licence, Creative Commons Attribution 4.0 International (CC By), which means it fulfils both the letter and the spirit of the Budapest Open Access Initiative’s definition of OA.

It’s great that we in vertebrate palaeontology can add this journal to the roster of OA journals in our field, already including Palaeontologia Electronica, Acta Palaeontologica Polonica, Palarch’s Journal of Vertebrate Paleontology, The Fossil Record and others. (Plus of course there is lots of vertebrate palaeontology in PLOS ONE and PeerJ.) I think that as a field, we are ahead of the curve in making the transition towards an all-OA literature.

 

Kaatedocus by Brian Engh

Kaatedocus is heading to the sidebar to help the cause.

We have a new page on the sidebar – here – where we’re collecting as many museum abbreviations as possible, the idea being that you can copy and paste them into your papers to rapidly populate the ‘Museum Abbreviations’ section. I grabbed about 100 from my own previous papers and a handful of others, so currently the list is highly skewed toward museums with (1) sauropods (2) that I’ve had reason to yap about. I’ve probably missed tons of museums that are important for people working on hadrosaurs or stegosaurs or (shudder) mammals. From here on out the list will grow as people suggest additions and edits in the comments on that page. So please get on over there and contribute!

Completely unrelated eyeball-bait art courtesy of Brian Engh, who writes,

I don’t even remember drawing this, I just found it lying around and spruced it up a bit today. It’s supposed to be some kinda diplodocid, maybe Kaatedocus, but I think the main goal of the drawing was to draw one with a sense of weight that felt right given that their center of mass is supposed to be so far back. I like the idea of them getting startled and popping up every now and again… [see also–MJW]

I’ve been taking a long-overdue look at some of the recently-described giant sauropods from China, trying to sort out just how big they were. Not a new pursuit for me, just one I hadn’t been back to in a while. Also, I’m not trying to debunk anything about this animal – as far as I know, there was no bunk to begin with – I’m just trying to get a handle on how big it might have been, for my own obscure purposes.

‘Huanghetitan’ ruyangensis was named by Lu et al. (2007) on the basis of a sacrum, the first 10 caudal vertebrae, some dorsal ribs and haemal arches, and a partial ischium. The holotype is 41HIII-0001 in the Henan Geological Museum. Lu et al. (2007) referred the new animal to the genus Huanghetitan, which was already known from the type species H. liujiaxiaensis (You et al., 2006). However, Mannion et al. (2013) found that the two species are not sister taxa and therefore ‘H.’ ruyangensis probably belongs to another genus, which has yet to be erected. Hence my use of scare quotes around the genus name.

Huanghetitan ruyangensis sacrum comparison

Here’s the sacrum of ‘H.’ ruyangensis from Lu et al. (2007: fig. 2). The original small scale bar is supposed to be 10cm. You know how I feel about scale bars (or maybe you don’t, in which case read this and this), but in this case the scale seems pretty legit based on limited measurements that are also given in the paper. I comped in the sacrum of Brachiosaurus altithorax FMNH P25107 from this post (many thanks to Phil Mannion for the photos!), and scaled it according to the max width across the second pair of sacral ribs, which Riggs (1904: p. 236) gives as 105 cm. The sacrum of ‘H.’ ruyangensis is a little bigger, but not vastly bigger. ‘H.’ ruyangensis had six sacrals to Brachiosaurus‘s five, so extra length is mostly illusory, whereas the extra width is mostly legit.

According to Lu et al. (2007), the anterior face of the first caudal vertebra in ‘H.’ ruyangensis measures 26.9 cm tall by 32 cm wide, and the centrum is 18.2 cm long. The same measurements in Brachiosaurus are 28 x 33 cm for the anterior face and 16 cm for the centrum length. It’s basically a tie.

What about the big rib? Lu et al. (2007) show a complete dorsal rib of ‘H.’ ruyangensis that is 293 cm long. That’s nothing to sniff at – the longest rib of Brachiosaurus, and the cause for the specific name altithorax (‘tall-bodied’), measures 274.5 cm, so the ‘H.’ ruyangensis rib is about 7% longer. But it’s not the longest rib known for any sauropod. As far as I know, that honor goes to a Supersaurus dorsal rib measuring 305 cm (Lovelace et al., 2008). The biggest Supersaurus caudal also blows away the caudals of both ‘H.’ ruyangensis and Brachiosaurus, with a anterior face 39 cm tall by 46 cm wide. But then diplodocids were all about that bass, so there’s not much point in comparing tail size with a titanosauriform if you’re trying to get a handle on overall body size. Still, the 35-40 ton Supersaurus shows that you can have 3-meter ribs without being anywhere near Argentinosaurus territory, mass-wise.

So what’s the verdict? ‘H.’ ruyangensis was a little bigger than the holotype of Brachiosaurus altithorax, but only by a few percent. It might have been about the same size as the XV2 specimen of Giraffatitan brancai. Or, who knows, it could have had completely different proportions and massed considerably more (or less). But on the current evidence, it doesn’t seem to have been one of the biggest sauropods of all time. I hope we get some more of it one of these days.

References

 

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 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 may 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.

The Carnegie Quarry, at Dinosaur National Monument, near Jensen, Utah, is arguably the most impressive dinosaur-fossil exhibit anywhere in the world — a covered, semi-excavated quarry that’s absolutely packed with big dinosaur fossils.

It’s also notoriously difficult to photograph: too big to fit into a single photo, and with poor contrast between the bones and matrix. This is the best picture I’ve found of part of it (from here) …

PublicDomain-DouglassQuarry-DinosaurNationalMonument-NPSPhoto

… although this one (from here) conveys the scale better:

field_work_at_dinosaur_national_monument

It’s one of the great sadnesses of my life that I’ve yet to visit DNM.

The quarry is historically important: discovered by Earl Douglass in 1909, it yielded among other specimens CM 3018, the holotype of Apatosaurus louisae and the principle subject of Gilmore’s (1936) monograph.

I’ve only recently become aware (thanks, Matt!) of Ken Carpenter’s (2013) detailed treatment of the history, sedimentology and taphonomy of the quarry — an important work that deserves to be widely read. Pages 10-14 are largely taken up with parts A-E of figure 10 — a big multi-page map of the quarry, showing the location of its most important specimens. Unfortunately, the five sections of this figure are all at slightly different scales in the PDF. I’ve rescaled them and pasted them together into a single big (4387 × 1210) image which I reproduce here:

Carpenter (2013:fig 10): map of the Carnegie quarry, composited from parts A-E.

Carpenter (2013:fig 10): map of the Carnegie quarry, composited from parts A-E.

Enjoy!

Update (six hours later)

I just heard from Ken Carpenter, who created the illustration. He has kindly sent me the full-resolution version — which is four times as big as the one I extracted from the PDF — and gave me permission to post it here on SV-POW! under the CC By licence. So here it is!

DNM Quarry map

Thanks, Ken!

Second update (12 March 2015)

Over on the Extinct Monsters blog, Ben Miller has published The Carnegie Quarry Diaspora. It’s a beautiful illustrated survey of some of the most important specimens to have come out of this quarry, including no fewer than seven important sauropod individuals.

References

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