## How light could a giant azhdarchid be?

### October 19, 2015

I imagine that by now, everyone who reads this blog is familiar with Mark Witton’s painting of a giant azhdarchid pterosaur alongside a big giraffe. Here it is, for those who haven’t seen it:

(This is the fifth and most recent version that Mark has created, taken from 9 things you may not know about giant azhdarchid pterosaurs.)

It’s one of those images that really kicks you in the brain the first time you see it. The idea that an animal the size of a giraffe could fly under its own power seems ludicrous — yet that’s what the evidence tells us.

But wait — what do we mean by “an animal the size of a giraffe”? Yes, the pterosaur in this image is the same height as the giraffe, but how does its weight compare?

Mark says “The giraffe is a big bull Masai individual, standing a healthy 5.6 m tall, close to the maximum known Masai giraffe height.” He doesn’t give a mass, but Wikipedia, citing Owen-Smith (1988), says “Fully grown giraffes stand 5–6 m (16–20 ft) tall, with males taller than females. The average weight is 1,192 kg (2,628 lb) for an adult male and 828 kg (1,825 lb) for an adult female with maximum weights of 1,930 kg (4,250 lb) and 1,180 kg (2,600 lb) having been recorded for males and females, respectively.” So it seems reasonable to use a mass intermediate between those of an average and maximum-sized male, (1192+1930)/2 = 1561 kg.

So much for the giraffe. What does the azhdarchid weigh? The literature is studded with figures that vary wildly, from the 544 kg that Henderson (2010) found for Quetzalcoatlus, right down to the widely cited 70 kg that Chatterjee and Templin (2004) found for the same individual — and even the astonishing 50 kg that seems to be favoured by Unwin (2005:192). In the middle is the 259 kg of Witton (2008).

It occurred to me that I could visualise these mass estimates by shrinking the giraffe in Mark’s image down to the various proposed masses, and seeing how credible it looks to imagine these reduced-sized giraffes weighting the same as the azhdarchid. The maths is simple. For each proposed azhdarchid mass, we figure out what it is as a proportion of the giraffe’s 1561 kg; then the cube root of that mass proportion gives us the linear proportion.

• 544 kg = 0.389 giraffe masses = 0.704 giraffe lengths
• 259 kg = 0.166 giraffe masses = 0.549 giraffe lengths
• 70 kg =0.0448 giraffe masses = 0.355 giraffe lengths

Let’s see how that looks.

On the left, we have Mark’s artwork, with the giraffe massing 1561 kg. On the right, we have three smaller (isometrically scaled) giraffes of masses corresponding to giant azhdarchid mass estimates in the literature. If Don Henderson (2010) is right, then the pterosaur weighs the same as the 544 kg giraffe, which to me looks pretty feasible if it’s very pneumatic. If Witton (2008) is right, then it weighs the same as the 259 kg giraffe, which I find hard to swallow. And if Chatterjee and Templin (2004) are right, then the giant pterosaur weighs the same as the teeny tiny 70 kg giraffe, which I find frankly ludicrous. (For that matter, 70 kg is in the same size-class as Georgia, the human scale-bar: the idea that she and the pterosaur weigh the same is just silly.)

What is the value of such eyeball comparisons? I’m not sure, beyond a basic reality check. Running this exercise has certainly made me sceptical about even the 250 kg mass range which now seems to be fairly widely accepted among pterosaur workers. Remember, if that mass is correct then the pterosaur and the 259 kg giraffe in the picture above weight the same. Can you buy that?

Or can we find extant analogues? Are there birds and mammals with the same mass that are in the same size relation as these images show?

# References

• Chatterjee, Sankar, and R. J. Templin. 2004. Posture, locomotion, and paleoecology of pterosaurs. Geological Society of America, Special Paper 376. 68 pages.
• Henderson, Donald M. 2010. Pterosaur body mass estimates from three-dimensional mathematical slicing. Journal of Vertebrate Paleontology 30(3):768-785.
• Witton, Mark P. 2008. A new approach to determining pterosaur body mass and its implications for pterosaur flight. Zitteliana 28:143-159.

## So what were apatosaurs doing with their crazy necks?

### September 14, 2015

We’ve noted that the Taylor et al. SVPCA abstract and talk slides are up now up as part of the SVPCA 2015 PeerJ Collection, so anyone who’s interested has probably taken a look already to see what it was about. (As an aside, I am delighted to see that two more abstracts have been added to the collection since I wrote about it.)

It was my privilege to present a talk on our hypothesis that the distinctive and bizarre toblerone-shaped necks of apatosaurs were an adaptation for intraspecific combat. This talk was based on an in-progress manuscript that Matt is lead-authoring. Also on board is the third SV-POW!sketeer, the silent partner, Darren Naish; and artist/ethologist Brian Engh.

Here is our case, briefly summarised from five key slides. First, let’s take a look at what is distinctive in the morphology of apatosaur cervicals:

Here I’m using Brontosaurus, which is among the more extreme apatosaurs, but the same features are seen developed to nearly the same extent in Apatosaurus louisae, the best-known apatosaur, and to some extent in all apatosaurs.

Now we’ll look at the four key features separately.

First, the cervicals ribs of sauropods (and other saurischians, including birds) anchored the longus colli ventralis and flexor colli lateralis muscles — ventral muscles whose job is to pull the neck downwards. By shifting the attachments points of these muscles downwards, apatosaurs enabled them to work with improved mechanical advantage — that is, to bring more force to bear.

Second, by redirecting the diapophyses and parapophyses ventrally, and making them much more robust than in other sauropods, apatosaurs structured their neck skeletons to better resist ventral impacts.

Third, because the low-hanging cervical ribs created an inverted “V” shape below the centrum, they formed a protective cradle for the vulnerable soft-tissue that is otherwise exposed on the ventral aspect of the neck: trachea, oesophagus, major blood vessels. In apatosaurus, all of these would have been safely wrapped in layers of connective tissue and bubble-wrap-like pneumatic diverticula. The presence of diverticula ventral to the vertebral centrum is not speculative – most neosauropods have fossae on the ventral surfaces of their cervical centra, and apatosaurines tend to have foramina that connect to internal chambers as well (see Lovelace et al. 2007: fig. 4, which is reproduced in this post).

Fourth, most if not all apatosaurs have distinctive ventrally directed club-like processes on the front of their cervical ribs. (It’s hard to tell with Apatosaurus ajax, because the best cervical vertebra of that species is so very reconstructed.) How did these appear in life? It’s difficult to be sure. They might have appeared as a low boss; or, as with rhinoceros horns, they might even have carried keratinous spikes.

Putting it all together, we have an animal whose neck can be brought downwards with great force; whose neck was mechanically capable of resisting impacts on its ventral aspect; whose vulnerable ventral-side soft-tissue was well protected; and which probably had prominent clubs or spikes all along the ventral aspect of the neck. And all of this was accomplished at the cost of making the neck a lot heavier than it would have been otherwise. Off the cuff, it seems likely that the cervical series alone would have massed twice as much in apatosaurines as in diplodocines of the same neck length.

Doubling the mass of the neck is a very peculiar thing for a sauropod lineage to do – by the Late Jurassic, sauropods were the leading edge of an evolutionary trend to lengthen and lighten the neck that had been running for almost 100 million years, through basal ornithodirans, basal dinosauromorphs, basal saurischians, basal sauropodomorphs, and basal sauropods. Whatever the selective pressures that led apatosaurines to evolve such robust and heavy necks, they must have been compelling.

The possibility that apatosaurs were pushing or crashing their necks ventrally in some form of combat accounts for all of the weird morphology documented above, and we know that sexual selection is powerful force that underlies a lot of bizarre structures in extant animals, and probably in extinct ornithodirans as well (see Hone et al. 2012, Hone and Naish 2013).

What form of combat, exactly? There are various possibilities, which we’ll discuss another time. But I’ll leave you with Brian Engh’s beautiful illustration of one possible form of combat: a powerful impact of one neck brought down onto the dorsal aspect of another.

We’re aware that this proposal is necessarily somewhat speculative. But we’re just not able to see any other explanation for the distinctive apatosaur neck. Even if we’re wrong about the ventrolateral processes on the cervical ribs supporting bosses or spikes, the first three points remain true, and given how they fly in the face of sauropods’ long history of making their necks lighter, they fairly cry out for explanation. If anyone has other proposals, we’ll be happy to hear them.

# References

• Hone, D. W., Naish, D., & Cuthill, I. C. (2012). Does mutual sexual selection explain the evolution of head crests in pterosaurs and dinosaurs?. Lethaia 45(2):139-156.
• Hone, D. W. E., & Naish, D. (2013). The ‘species recognition hypothesis’ does not explain the presence and evolution of exaggerated structures in non‐avialan dinosaurs. Journal of Zoology 290(3):172-180.
• Lovelace, D. M., Hartman, S. A., & Wahl, W. R. (2007). Morphology of a specimen of Supersaurus (Dinosauria, Sauropoda) from the Morrison Formation of Wyoming, and a re-evaluation of diplodocid phylogeny. Arquivos do Museu Nacional, Rio de Janeiro 65(4):527-544.

## Help us assemble all of the museum abbreviations

### March 6, 2015

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]

## The “finger-like” parapostzygapophyseal processes of Qijianglong

### January 29, 2015

There’s a new mamenchisaurid in town! It’s called Qijianglong (“dragon of Qijiang”), and it’s the work of Xing et al. (2015).

Life restoration of Qijianglong, by Cheung Chungtat.

As far as I can make out, the life restoration is also due to Xing Lida: at least, every instance of the picture I’ve seen says “Credit: Xing Lida”. If that’s right, it’s an amazing display of dual expertise to produce both the science and the art! We could quibble with details, but it’s a hundred times better than I could ever do. [Update: no, it’s by Cheung Chungtat, but being uniformly mis-attributed in the media. Thanks to Kevin for the correction in the comment below.]

There’s a mounted skeleton of this new beast in the museum local to where it was found, though I don’t know how much of the material is real, or cast from the real material. Here it is:

A reconstructed skeleton of Qijianglong now on display in Qijiang Museum

A new sauropod is always great news, of course, and it’s a source of shame to us that we cover so few of them here on SV-POW!. (Just think of some of the ones we’ve missed recently … Leikupal, for example.)

But as is so often the case, the most interesting thing about this new member of the club is its vertebrae — specifically the cervicals. Here they are:

Xing et al. (2015), FIGURE 11. Anterior cervical series of Qijianglong guokr (QJGPM 1001) in left lateral views unless otherwise noted. A, axis; B, cervical vertebra 3; C, cervical vertebra 4; D, cervical vertebrae 5 and 6; E, cervical vertebra 7 and anterior half of cervical vertebra 8 (horizontally inverted; showing right side); F, posterior half of cervical vertebra 8 and cervical vertebra 9; G, cervical vertebra 10; H, cervical vertebra 11; I, close-up of the prezygapophy- sis-postzygapophysis contact between cervical vertebrae 3 and 4 in dorsolateral view, showing finger-like process lateral to postzygapophysis; J, close- up of the postzygapophysis of cervical vertebra 5 in dorsal view, showing finger-like process lateral to postzygapophysis. Arrow with number indicates a character diagnostic to this taxon (number refers to the list of characters in the Diagnosis). All scale bars equal 5 cm. Abbreviations: acdl, anterior centrodiapophyseal lamina; cdf, centrodiapophyseal fossa; plc, pleurocoel; pocdl, postcentrodiapophyseal lamina; poz, postzygapophysis; pozcdf, post- zygapophyseal centrodiapophyseal fossa; pozdl, postzygodiapophyseal lamina; ppoz, finger-like process lateral to postzygapophysis; ppozc, groove for contact with finger-like process; przdl, prezygodiapophyseal lamina; sdf, spinodiapophyseal fossa.

(At first, I couldn’t figure out what this pocdl abbreviation meant. Then I realised it was a vanilla posterior centrodiapophyseal lamina. Come on, folks. That element has had a standard abbreviation since 1999. Let’s use our standards!)

The hot news in these cervicals is the presence of what the authors call “a distinct finger-like process extending from the postzygapophyseal process beside a zygapophyseal contact”. They don’t give a name to these things, but I’m going to call them parapostzygapophyses since they’re next to the postzygapophyses. [Update: see the comment from Matt below.]

You can get some sense of this morphology from the figure above — although it doesn’t help that we’re looking at tiny greyscale images which really don’t convey 3d structure at all. The best illustration is part J of the figure:

What are these things? The paper itself says disappointingly little about them. I quote from page 9:

From the axis to at least the 14th cervical vertebra, a finger- like process extends posteriorly above the postzygapophysis and overlaps onto the dorsolateral surface of the prezygapophysis of the next vertebra (Fig. 11I, J). These processes are unique to Qijianglong, unlike all previously known mamenchisaurids that are preserved with cervical vertebrae (e.g., Chuanjiesaurus, Mamenchisaurus spp., Omeisaurus spp., Tonganosaurus). Therefore, the neck of Qijianglong presumably had a range of motion restricted in sideways.

That’s it.

So what are these things? The authors — who after all have seen the actual fossils, not just the rather inadequate pictures — seem to assume that they are a stiffening adaptation, but don’t discuss their reasoning. My guess — and it’s only a guess — it that they assumed that this is what was going on with these processes because it’s what people have assumed about extra processes on xenarthrous vertebrae. But as best as I can determine, that’s not been demonstrated either, only assumed. Funny how these things seem to get a pass.

Armadillo lumbar vertebrae in posterior, anterior and right lateral views.

So what are these processes? It’s hard to say for sure without having seen the fossils, or at least some better multi-view photos, but the obvious guess is that they are our old friends epipophyses, in extreme form. That is, they are probably enlarged attachment points for posteriorly directed dorsal muscles, just as the cervical ribs are attachment points for posteriorly directly ventral muscles.

It’s a shame that Xing et al. didn’t discuss this (and not only because it would probably have meant citing our paper!) Their new beast seems to have some genuinely new and interesting morphology which is worthy of a bit more attention than they gave it, and whose mechanical implications could have been discussed in more detail. Until more is written about these fossils (or better photographs published) I think I am going to have to suspend judgement on the as-yet unjustified assumption that the parapostzygs were there to make the neck rigid against transverse bending.

A final thought: doesn’t JVP seem terribly old-fashioned now? It’s not just the paywall — apologies to those many of you who won’t be able to read the paper. The greyscaling of the figures is part of it — something that makes no sense at all in 2015. The small size and number of the illustrations is also a consequence of the limited page-count of a printed journal — it compares poorly with, for example, the glorious high-resolution colour multiview illustrations in Farke et al.’s (2013) hadrosaur description in PeerJ. Seems to me that, these days, all the action is over at the OA journals with infinite space — at least when it comes to descriptive papers.

# References

• Farke, Andrew A., Derek J. Chok, Annisa Herrero, Brandon Scolieri and Sarah Werning. (2013) Ontogeny in the tube-crested dinosaur Parasaurolophus (Hadrosauridae) and heterochrony in hadrosaurids. PeerJ 1:e182. doi:10.7717/peerj.182
• Xing Lida, Tetsuto Miyashita, Jianping Zhang, Daqing Li, Yong Ye, Toru Sekiya, Fengping Wang & Philip J. Currie. 2015. A new sauropod dinosaur from the Late Jurassic of China and the diversity, distribution, and relationships of mamenchisaurids. Journal of Vertebrate Paleontology. doi:10.1080/02724634.2014.889701

### December 10, 2014

Life restoration of Aquilops by Brian Engh. Farke et al. (2014: fig. 6C). CC-BY.

Today sees the description of Aquilops americanus (“American eagle face”), a new basal neoceratopsian from the Cloverly Formation of Montana, by Andy Farke, Rich Cifelli, Des Maxwell, and myself, with life restorations by Brian Engh. The paper, which has just been published in PLOS ONE, is open access, so you can download it, read it, share it, repost it, remix it, and in general do any of the vast scope of activities allowed under a CC-BY license, as long as we’re credited. Here’s the link – have fun.

Obviously ceratopsians are much more Andy’s bailiwick than mine, and you should go read his intro post here. In fact, you may well be wondering what the heck a guy who normally works on huge sauropod vertebrae is doing on a paper about a tiny ceratopsian skull. The short, short version is that I’m here because I know people.

OMNH 34557, the holotype of Aquilops

The slightly longer version is that OMNH 34557, the holotype partial skull of Aquilops, was discovered by Scott Madsen back in 1999, on one of the joint Cloverly expeditions that Rich and Des had going on at the time (update: read Scott’s account of the discovery here). That the OMNH had gotten a good ceratopsian skull out of Cloverly has been one of the worst-kept secrets in paleo. But for various complicated reasons, it was still unpublished when I got to Claremont in 2008. Meanwhile, Andy Farke was starting to really rock out on ceratopsians at around that time.

For the record, the light bulb did not immediately go off over my head. In fact, it took a little over a year for me to realize, “Hey, I know two people with a ceratopsian that needs describing, and I also know someone who would really like to head that up. I should put these folks together.” So I proposed it to Rich, Des, and Andy in the spring of 2010, and here we are. My role on the paper was basically social glue and go-fer. And I drew the skull reconstruction – more on that in the next post.

One of the world’s smallest ceratopsians meets one of the largest: the reconstructed skull of Aquilops with Rich Cifelli and Pentaceratops for scale. Copyright Leah Vanderburg, courtesy of the Sam Noble Oklahoma Museum of Natural History.

Anyway, it’s not my meager contribution that you should care about. I am fairly certain that, just as Brontomerus coasted to global fame on the strength of Paco Gasco’s dynamite life restoration, whatever attention Aquilops gets will be due in large part to Brian Engh’s detailed and thoughtful work in bringing it to life – Brian has a nice post about that here. I am very happy to report that the three pieces Brian did for us – the fleshed-out head that appears at the top of this post and as Figure 6C in the paper, the Cloverly environment scene with the marauding Gobiconodon, and the sketch of the woman holding an Aquilops – are also available to world under the CC-BY license. So have fun with those, too.

Finally, I need to thank a couple of people. Steve Henriksen, our Vice President for Research here at Western University of Health Sciences, provided funds to commission the art from Brian. And Gary Wisser in our scientific visualization center used his sweet optical scanner to generate the hi-res 3D model of the skull. That model is also freely available online, as supplementary information with the paper. So if you have access to a 3D printer, you can print your own Aquilops – for research, for teaching, or just for fun.

Cloverly environment with Aquilops and Gobiconodon, by Brian Engh (CC-BY).

Next time: Aquilöps gets röck döts.

### Reference

Farke, A.A., Maxwell, W.D., Cifelli, R.L., and Wedel, M.J. 2014. A ceratopsian dinosaur from the Lower Cretaceous of Western North America, and the biogeography of Neoceratopsia. PLoS ONE 9(12): e112055. doi:10.1371/journal.pone.0112055

## Beards of the Mesozoic: Bob Nicholls edition

### September 11, 2013

We’ve blogged a lot of Bob Nicholls‘ art (here, here, and here) and we’ll probably continue to do so for the foreseeable future. We don’t have much choice: he keeps drawing awesome things and giving us permission to post them. Like this defiantly shaggy Apatosaurus, which was probably the star of the Morrison version of Duck Dynasty. Writes Bob:

On my way home at the airport I did a sketch of your giant Apatosaurus* — see attachment.  My thought was that massive thick necks were probably pretty sexy things to apatosaurs, so maybe sexually mature individuals used simple feathers (stage 1, 2 or 3?) to accentuate the neck profile.  The biggest males would of course have the most impressive growths so in the attached sketch your giant has one of the biggest beards in Earth’s history!  What do you think of this idea?

Well, I think it’s awesome. And entirely plausible, for reasons already explained in this post.

“Now, wait,” you may be thinking, “I thought you guys said that sauropod necks weren’t sexually selected.” Actually we made a slightly different point: that the available evidence does not suggest that sexual selection was the primary driver of sauropod neck elongation. But we also acknowledged that biological structures are almost never single-purpose, and although the long necks of sauropods probably evolved to help them gather more food, there is no reason that long necks couldn’t have been co-opted as social billboards. This seems especially likely in Apatosaurus, where the neck length is unremarkable** but the neck fatness is frankly bizarre (and even inspired a Star Wars starfighter!).

I also love the “mobile ecosystem” of birds, other small dinosaurs, and insects riding on this Apatosaurus or following in its train. It’s a useful reminder that we have no real idea what effect millions of sauropods would have on the landscape. But it’s not hard to imagine that most Mesozoic terrestrial ecosystems were sauropod-driven in a thousand cascading and ramifying chains of cause and effect. I’d love to know how that worked. At heart, I’m still a wannabe chrononaut, and all my noodlings on pneumaticity and sauropod nerves and neural spines and so on are just baby steps toward trying to understand sauropod lives. Safari by way of pedantry: tally-ho!

For other speculative apatosaurs, see:

* “My” giant is the big Oklahoma Apatosaurus, which I gave a talk on at SVPCA a couple of weeks ago. See these posts for more details (123).

** Assuming we can be blasé about a neck that is more than twice as long (5 m) as a world-record giraffe neck (2.4 m), for garden variety Apatosaurus, or three times that length for the giant Oklahoma Apatosaurus (maybe 7 m).

## My attempt at a fully Engh-ed out brachiosaur

### July 26, 2013

Last Sunday I got to hang out with Brian Engh and some of his friends in LA. You may remember Brian from thisthis, this, this, and, most notoriously, this. We got to drawing dinosaurs, naturally.

Now, for me to try to draw dinosaurs next to Brian is more than a little intimidating. I really felt the need to bring my A-game. So this is what I came up with. I’m posting it not because I think it is particularly likely* but because the blog has been a little sauropod-lite this summer, and heck, it’s Friday.

* Although frigatebirds and anoles and such might have some things to say about that.