Someone on Facebook asked whether sauropods had subcutaneous fat, and by the time my answer hit five paragraphs I thought, “The merciful thing to do here is blog this and link to it.” So here are some things to keep in mind regarding the integumentary systems of sauropods.

Emu dissection at UC Santa Cruz back in 2004. Note the fat pad on the chest and how it abruptly comes to an end.

Sauropods may have had some subcutaneous fat – we can’t rule it out – but it probably wasn’t broadly distributed as it is in mammals. In the interaction of their air sac systems with connective tissue, sauropods were probably a lot like birds. Most birds don’t have subcutaneous fat all over their bodies. Instead, they have subcutaneous air sacs (or pneumatic diverticula) over parts or all of their bodies – in pelicans these are like bubble wrap under the skin, presumably for impact padding and insulation (Richardson 1939, 1943). The diverticula go everywhere and most places they go, they replace adipose tissue, even the harmless bits of fat between muscles that are basically the body’s packing peanuts (broiler chickens don’t count here, they’ve been artificially selected to be radically unhealthy). We suspect that sauropods had subcutaneous diverticula because so many other aspects of their pneumatic systems correspond to those of living birds (see the discussion in Wedel and Taylor 2013b for more on that).

Contrast the narrow line of adipose tissue down the ventral midline with the almost-completely-lean hindlimb.

That’s not to say that birds don’t have subcutaneous fat, just that it tends to be highly localized. Back in grad school I got to help dissect an emu (link) and a rhea (one, two), and in both cases the fat was concentrated in two places: huge paired fat pads over the pelvis, like big lozenges, and a concentration over the sternum with extensions along the ventral midline from the base of the neck to the cloaca. It was weird, the fat would be present and then it would just stop, like somebody flipped a switch. We pulled 18 lbs of fat off a 102-lb emu, so it wasn’t a trivial part of the body composition. IME, even relatively fatty birds like ducks tend to have the fat start and stop abruptly, and again, the fat deposits tend to be concentrated on the breast and tummy and over the hips.

Fat-tailed gecko, borrowed from here.

A lot of lizards and crocs and even some turtles carry fat deposits in their tails, and that is one aspect of sauropod anatomy that is definitely un-bird-like. So some sauropods might have had fat tails.

We can be pretty sure that at least some sauropods had thick skin. Osteoderms (armor plates) from Madagascar show that the bits that were embedded in the skin could be up to 7cm thick, so the surrounding skin was at least that thick and possibly even thicker (Dodson et al. 1998). And that was most likely on Rapetosaurus, which was not a huge sauropod. So giant sauropods might have had even thicker skin. Maybe. Remember that big-ass-ness (here arbitrarily defined as 40+ metric tons) evolved independently in:

They probably didn’t all get there looking the same way, beyond sharing the basic sauropod bauplan.

I’m too lazy to write about the fossil evidence for scaly skin and keratinous spines in sauropods – see this post and the references therein.

One final thing to think about is scar tissue. The scar tissue on the chest of a male elephant seal can be up to 5cm thick. Some sauropods might have had calluses or patches of scar tissue in predictable places, from combat, or habitually pushing down trees with their chests or tails, or doing whatever weird things real animals do when we’re not looking.

So in the toolbox of things to play with in reconstructing the integument of sauropods, we have thick skin, keratinous spines, osteoderms, fat pads (possibly concentrated over the hips and shoulders or on the tail), subcutaneous diverticula, calluses, and scar tissue. And that’s just the stuff we have found or reasonably inferred so far, barely 150 years into our exploration of animals we know mostly from bits and bobs, whose size means they mostly got buried in big sediment-dumping events that would not preserve delicate integumentary structures. Give us a millennium of Yixian Formations and Mahajanga Basins and Howe Quarries and the picture will probably change, and the arrow of history dictates that it will change for the weirder.

Likely? Probably not. But roll the evolutionary dice for 160 million years and you’ll get stranger things than this. Recycled from this post.

Finally, and related to my observation about big-ass-ness: sauropods were a globally-distributed radiation of animals from horse-sized to whale-sized that existed from the Late Triassic to the end of the Cretaceous. The chances that all of them had the same integumentary specializations, for display or combat or insulation or camouflage or whatever, are pretty darned low. Support weird sauropods – and vanilla ones, too.

Almost immediate update: I’ve just been reminded about Mark Witton’s excellent post on dinosaur fat from a couple of years ago. Go read that, and the rest of his blog. I’m sure I missed other relevant posts at other excellent blogs – feel free to remind me in the comments.



One of the field trips for last year’s SVPCA meeting was a jaunt to Nottingham to see the Dinosaurs of China exhibit at Wollaton Hall. We got to see a lot of stuff, including original fossils of some pretty famous feathered dinos – but of course what really captured our attention was the mounted Mamenchisaurus. This is a cast of the good old M. hochuanensis holotype specimen that has been put up all over the world, including in a car-park in Copenhagenon stilts in Chicago and even in a flooded basement in Slovenia.

Wollaton Hall houses the Nottingham Museum of Natural History, which is a fantastic trove of weird and wonderful things from around the world. We should really post about those things – I had them in mind when I was recently lamenting my lousy conversion rate of museum visit photos into blog posts. That will have to wait for another time. I’ll just note in closing that grand buildings and mounted sauropods go together like peanut butter and chocolate, and that this field trip was outstanding.

Mike Taylor, Matt Wedel, Darren Naish, and Bob Nicholls (kneeling) at Wollaton Hall, with Mamenchisaurus hochuanensis for scale.

Back in 2009, I posted on a big cervical series discovered in Big Bend National Park. Then in 2013 I posted again about how I was going to the Perot Museum in Dallas to see that cervical series, which by then was fully prepped and on display but awaiting a full description. Ron Tykoski and Tony Fiorillo (2016) published that description a couple of years ago, and after almost five years it’s probably time I posted an update.

I did visit the Perot Museum in 2013 and Ron and Tony kindly let me hop the fence and get up close and personal with their baby. I got a lot of nice photos and measurements of the big specimen. It’s an impressive thing. Compared to the other big sauropod cervicals I’ve gotten to play with, these vertebrae aren’t all that long – the two longest centra are about 80cm, compared to ~120cm for Sauroposeidon, Puertasaurus, and Patagotitan, and 137cm for Supersaurus (more details here) – but they are massive. According to the table of measurements (yay!) in Tykoski and Fiorillo (2016), which accord well with the measurements I took when I was there, the last vert is 117.5cm tall from the bottom of the cervical rib to the top of the neural spine, 98.4cm wide across the diapophyses, and has a cotyle measuring 29cm tall by 42cm wide. Here it is with me for scale:

I guarantee you, standing next to that thing and imagining it being inside the neck of a living animal is a breathtaking experience.

I failed in my mission in one way. In a comment on my 2013 post, I said, “I’ll try to get some good lateral views of the mount with as little perspective as possible.” But it can’t be done – the geometry of the room and the size of the skeleton don’t allow it, as Ron noted in the very next comment. There is one place in the exhibit hall where you can get the whole skeleton into the frame, and that’s a sort of right anterolateral oblique view. Here’s my best attempt:

So, this is an awesome specimen and you should go see it. As you can see from the photos, the vertebrae are right on the other side of the signage, with no glass between you and them, so you can see a lot. The rest of the exhibits are top notch as well. Definitely worth a visit if you find yourself within striking distance of Dallas.


Tykoski, R.S. and Fiorillo, A.R. 2016. An articulated cervical series of Alamosaurus sanjuanensis Gilmore, 1922 (Dinosauria, Sauropoda) from Texas: new perspective on the relationships of North America’s last giant sauropod. Journal of Systematic Palaeontology 15(5):339-364.

This past weekend I was camping up the coast at Hearst San Simeon State Park, with my son, London, and Brian Engh.

We went to see the elephant seal colony at Piedras Blancas. It was my first time seeing elephant seals in the wild. Not having done any research in advance, I was expecting something like this:

In other words, a small number of elephant seals, not doing much, basically at binocular distance from the viewing area. Obviously we did get some of that, since I have a picture of it. But that was up the coast a bit, at the start of Boucher Trail near the Piedras Blancas lighthouse.

We spent most of our time at the main Piedras Blancas rookery, where just the southeastern half of the viewing area looked – and sounded – like this:

We also saw a lot of this (semi-groady iPhone-through-binocular shot by me):

and even some of this (much nicer photo courtesy of Brian Engh):

I’ll have a lot more to say about this real soon, including more video, but it’s late and I need sleep. Stay tuned!

When I was nine, a copy of Don Glut’s The New Dinosaur Dictionary turned up in my local Waldenbooks. It wasn’t my first dinosaur book, by far – I’d been a dinosaurophile since the age of three. But The New Dinosaur Dictionary was different.

Up to that point, I had subsisted on a heavy diet of kids’ dino books and the occasional article in National Geographic and Ranger Rick. The kids’ books were aimed at kids and the magazine articles were pitched at an engagingly popular level. I didn’t understand every word, but they were clearly written for curious layfolk, not specialists.

A typical spread from The New Dinosaur Dictionary (Glut, 1982). The armored sauropod blew my young mind.

The New Dinosaur Dictionary was something else entirely. It had photos of actual dinosaur bones and illustrations of skeletons with cryptic captions like, “Skeleton of Daspletosaurus torosus. (After Russell)”. Okay, clearly this Russell cove was out there drawing dinosaur skeletons and this book had reproduced some of them. But nobody I knew talked like that, and the books I had access to up to that point held no comparable language.

The New Dinosaur Dictionary (Glut, 1982: p. 271)

Then there was stuff like this: “The so-called Von Hughenden sauropod restored as a brachiosaurid by Mark Hallett”. A chain of fascinating and pleasurable ideas detonated in my brain. “The so-called” – say what now? Nobody even knew what to call this thing? Somehow I had inadvertently sailed right to the edge of human knowledge of dinosaurs, and was peering out into taxa incognita. “Restored as a brachiosaurid” – so this was just one of several possible ways that the animal might have looked. Even the scientists weren’t sure. This was a far cry from the bland assurances and blithely patronizing tones of all my previous dinosaur books.

“By Mark Hallett.” I didn’t know who this Hallett guy was, but his art was all over the book, along with William Stout and some guy named Robert T. Bakker and a host of others who were exploding my conception of what paleo art could even be. Anyway, this Mark Hallett was someone to watch, not only because he got mentioned by name a lot, but because his art had a crisp quality that teetered on some hypercanny ridge between photorealism and scribbling. His sketches looked like they might just walk off the page.

In case that line about scribbling sounds dismissive: I have always preferred sketches by my favorite artists to their finished products. The polished works are frequently inhumanly good. They seem to have descended in a state of completed perfection from some divine realm, unattainable by mere mortals. Whereas sketches give us a look under the hood, and show how a good artist can conjure light, shadow, form, weight, and texture from a few pencil strokes. Put it this way: I am anatomist by temperament first, and by training and occupation second. Of course I want to see how things are put together.

The New Dinosaur Dictionary (Glut, 1982: p. 75)

Anyway, The New Dinosaur Dictionary was something completely new in my experience. It wasn’t aimed at kids and written as if by kids, like lots of kids’ books. It wasn’t even written by adults talking down (deliberately or inadvertently) to kids, or trying to reach a wide audience that might include kids. It was written by an adult, aiming at other adults. And it was admitting in plain language that we didn’t know everything yet, that there were lots of animals trembling on the outer threshold of scientific knowledge. I didn’t understand half of it – I was down in an ontogenetic trench, looking up as these packets of information exploded like fireworks over my head.

In Seeing In the Dark, the best book about why you should go out stargazing for yourself, Timothy Ferris writes about growing up on Florida’s Space Coast in the early 1960s, and watching the first generation of artificial satellites pass overhead:

I felt like an ancient lungfish contemplating the land from the sea. We could get up there.

That’s precisely the effect that The New Dinosaur Dictionary had on me: I could get up there. Maybe not immediately. But there were steps, bodies of knowledge that could be mastered piecemeal, and most of all, mysteries to be resolved. The book itself was like a sketch, showing how from isolated and broken bones and incomplete skeletons, scientists and artists reconstructed the world of the past, one hypothesis at a time. Now I take it for granted, because I’ve been behind the curtain for a couple of decades. But to my 9-year-old self, it was revolutionary.

This has all come roaring back because of something that came in the mail this week. Or rather, something that had been waiting in the mailroom for a while, that I finally picked up this week: a package from Mark Hallett, enclosing a copy of his 2018 dinosaur calendar. And also this:


An original sketch, which he gave to me as a Christmas present. The published version appears on one of the final pages of our book, where we discuss the boundaries between the known – the emerging synthesis of sauropod biology that we hoped to bring to a broader audience by writing the book in the first place – and the unknown – the enduring mysteries that Mark and I think will drive research in sauropod paleobiology for the next few decades. Presented without a caption or commentary, the sketch embodies sauropods as we see them: emerging from uncertainty and ignorance one hard-won line at a time, with ever-increasing solidity.

Thank you, Mark, sincerely. That sketch, what it evokes, both for me now and for my inner 9-year-old – you couldn’t have chosen a better gift. And I couldn’t be happier. Except perhaps to someday learn that our book exploded in the mind of a curious kid the way that The New Dinosaur Dictionary did for me 34 years ago, a time that now seems as distant and romantic as the primeval forests of the Mesozoic.

Computer programmer, essayist and venture capitalist Paul Graham writes:

In most fields, prototypes have traditionally been made out of different materials. Typefaces to be cut in metal were initially designed with a brush on paper. Statues to be cast in bronze were modelled in wax. Patterns to be embroidered on tapestries were drawn on paper with ink wash. Buildings to be constructed from stone were tested on a smaller scale in wood.

What made oil paint so exciting, when it first became popular in the fifteenth century, was that you could actually make the finished work from the prototype. You could make a preliminary drawing if you wanted to, but you weren’t held to it; you could work out all the details, and even make major changes, as you finished the painting.

You can do this in software too. A prototype doesn’t have to be just a model; you can refine it into the finished product. I think you should always do this when you can. It lets you take advantage of new insights you have along the way. But perhaps even more important, it’s good for morale.

– Paul Graham, “Design and Research

Mike and I have long been drawn by the idea that blog posts, like conference talks and posters, could be first drafts of research papers. In practice, we haven’t generated many successful examples. We basically wrote our 2013 neural spine bifurcation paper as a series of blog posts in 2012. And Mike’s 2014 neck cartilage paper grew out of this 2013 blog post, although since he accidentally ended up writing 11 pages I suppose the blog post was more of a seed than a draft.

I should also note that we are far from the first people to do the blog-posts-into-papers routine. The first example I know of in paleo was Darren’s Tet Zoo v1 post on azhdarchid paleobiology, which formed part of the skeleton of Witton and Naish (2008).

Nevertheless, the prospect of blogging as a way to generate research papers remains compelling.

And as long as I’m on about blogging and papers: sometimes people ask if blogging doesn’t get in the way of writing papers. I can’t speak for anyone else, but for me it goes in the opposite direction: I blog most when I am most engaged and most productive, and drops in blogging generally coincide with drops in research productivity. I think that’s because when I’m rolling on a research project, I am constantly finding or noticing little bits that are cool and new, but which aren’t germane to what I’m working on at the moment. I can’t let those findings interfere with my momentum, but I don’t want to throw them away, either. So I blog them. Also the blog gives me a place to burn off energy at the end of the day, when I can still produce words but don’t have the discipline to write technical prose.

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The photo at the top of the post is of Giraffatitan dorsal vertebrae in a case at the MfN Berlin, from Mike’s and my visit with the DfG 533 group back in late 2008. I picked that photo so I could make the following dumb off-topic observation: with its upturned transverse processes, the dorsal on the right looks like it’s being all faux melodramatic, a la:

This post started out as a comment on this thread, kicked off by Dale McInnes, in which Mike Habib got into a discussion with Mike Taylor about the max size of sauropods. Stand by for some arm-waving. All the photos of outdoor models were taken at Dino-Park Münchehagen back in late 2008.

I think it’s all too easy to confuse how big things do get from how big they could get, assuming different selection pressures and ecological opportunities. I’m sure someone could write a very compelling paper about how elephants are as big as they could possibly be, or Komodo dragons, if we didn’t have indricotheres and Megalania to show that the upper limit is elsewhere. This is basically what Economos (1981) did for indricotheres, either forgetting about sauropods or assuming they were all aquatic.

Truly, a mammal of excellence and distinction. With Mike and some dumb rhino for scale.

In fact, I’ll go further: a lot of pop discussions of sauropod size assume that sauropods got big because of external factors (oxygen levels, etc.) but were ultimately limited by internal factors, like bone and cartilage strength or cardiovascular issues. I think the opposite is more likely: sauropods got big because of a happy, never-repeated confluence of internal factors (the Sander/et al. [2008, 2011, 2013] hypothesis, which I think is extremely robust), and their size was limited by external, ecological factors.

Take a full-size Argentinosaurus or Bruhathkayosaurus – even modest estimates put them at around 10x the mass of the largest contemporary predators. Full-grown adults were probably truly predator-immune, barring disease or senescence. So any resources devoted to pushing the size disparity higher, instead of invested in making more eggs, would basically be wasted.

If there was reproductive competition among the super-giants, could the 100-tonners have been out-reproduced by the 70-tonners, which put those extra 30 tonnes into making babies? Or would the 100-tonners make so many more eggs than the 70-tonners (over some span of years) that they’d still come out on top? I admit, I don’t know enough reproductive biology to answer that. (If you do, speak up in the comments!) But if – if – 70-tonners could out-reproduce 100-tonners, that by itself might have been enough to put a cap on the size of the largest sauropods.

Another possibility is that max-size adult sauropods were neither common nor the target of selection. In most populations most of the time, the largest individuals might have been reproductively active but skeletally-immature and still-growing subadults (keep in mind that category would encompass most mounted sauropod skeletons, including the mounted brachiosaurs in Chicago and Berlin). If such individuals were the primary targets of selection, and they were selected for a balance of reproductive output and growth, then the few max-size adults might represent the relatively rare instances in which the developmental program “overshot” the selection target.

Dave Hone and Andy Farke and I mentioned this briefly in our 2016 paper, and it’s come up here on the blog several times before, but I still have a hard time wrapping my head around what that would mean. Maybe the max-size adults don’t represent the selective optimum, but rather beneficial traits carried to extreme ends by runaway development. It seems at least conceivable that the bodies of such animals might have been heavily loaded with morphological excrescences – like 15- to 17-meter necks – that were well past the selective optimum. As long as those features weren’t inherently fatal, they could possibly have been pretty darned inefficient, riding around on big predator-immune platforms that could walk for hundreds of kilometers and survive on garbage.

What does that swerve into weird-but-by-now-well-trod ground have to do with the limits on sauropod size? This: if max-size adults were not heavy selection targets, either because the focus of selection was on younger, reproductively-active subadults, or because they’d gotten so big that the only selection pressure that could really affect them was a continent-wide famine – or both – then they might not have gotten as big as they could have (i.e., never hit any internally-imposed, anatomical or biomechanical limits) because nothing external was pushing them to get any bigger than they already were.

Or maybe that’s just a big pile of arm-wavy BS. Let’s try tearing it down, and find out. The comment thread is open.