Plateosaurus trossingensis AMNH 6810 sacrum and pelvis in left dorsolateral view

We’ve shown a lot of sauropod sacra around here lately (for example here, here, and here), so here’s a little look back down the tree.

You haven’t heard from me much lately because I’ve been busy teaching anatomy. Still, I get to help people dissect for a living, so I can’t complain.

Further bulletins as events warrant.

Plateosaurus engelhardti (originally P. trossingensis) SMNS 13200 cervical vertebrae 3-8 in left lateral view. C8 is roughly 15 cm long.

In the recent post on serial variation in sauropod cervicals, I wrote:

Even in ‘adult’ sauropods like the big mounted Apatosaurus and Diplodocus skeletons, the anterior cervicals are less complex than the posterior ones. Compared to posterior cervicals, anterior cervicals tend to have simpler pneumatic fossae and foramina, fewer laminae, and unsplit rather than bifid spines. In all of these things the anterior cervicals are similar to those of juveniles of the same taxa, and to those of adults of more basal taxa. This is also true in prosauropods–in Plateosaurus, the full complement of vertebral laminae is not present until about halfway down the neck.

I was working from memory there and actually understated things a bit. Plateosaurus presacral vertebrae don’t have well-developed spinal laminae, but they do eventually get the four major diapophyseal laminae–the anterior centrodiapophyseal lamina (ACDL), posterior centrodiapophyseal lamina (PCDL), prezygodiapophyseal lamina (PRDL), and postzygodiapophyseal lamina (PODL–please see the lamina tutorial if you need a refresher on these and the other 15 commonly identified laminae). But they aren’t all present halfway down the neck–the ACDL doesn’t really show up until the cervicodorsal transition. The other three kick in sequentially down the neck, as shown in the above image. I think that’s pretty cool, that you get different character states expressed at different points along the neck, in one individual organism, at one time. And possibly also at different times–in sauropods, the anterior cervicals tend to look more ‘juvenile’ or ‘primitive’, even in adult animals, so all of the cervicals go through a juvenile stage, but not all of them grow out of it. I don’t know if there’s a word for that–some kind of serial heterochronotopomorphy or the like–but hopefully someone will enlighten me.

I took the original photo in the collections at the Staatliches Museum für Naturkunde Stuttgart in the spring of 2004. Markus Moser and Rainer Schoch were wonderful hosts during my visit. Mike did all the work of turning the raw photo into a figure, so thanks to him for getting this off my hard drive and out into the world.

Item 1: With his new piece at the Guardian,  “Persistent myths about open access scientific publishing”, Mike continues to be a thorn in the side of exploitative commercial publishers, who just can’t seem to keep their facts straight. This time Mike unravels some choice bits of nonsense that keep getting circulated about open access publishing: that OA publishing must necessarily cost as much as barrier-based publishing, that the peer review process is expensive for publishers, and that authors who can’t pay OA publication fees will be left out in the cold. It’s cleanly and compellingly argued–go read for yourself.

Item 2: The Yates et al. prosauropod pneumaticity paper is officially published in the latest issue of Acta Palaeontologica Polonica, and I have updated the citation and links accordingly. This may not seem like big news, in that the accepted manuscript has been available online for 13 months, and the final published version does not differ materially from that version other than being pretty. But it’s an opportunity to talk about something that we haven’t really addressed here before, which is the potential for prompt publication to accelerate research.

A bit of background: standard practice at APP is to post accepted manuscripts as soon as they’re, well, accepted, unless the authors ask otherwise (for example, because the paper contains taxonomic acts and the first public version needs to be the version of record). Not everyone likes this policy–I know Darren objects, and I’m sure there are others. The chief complaint is that it muddies the waters around when the paper is published. Is a paper published when a manuscript is posted to a preprint server like arXiv, or when the accepted manuscript is made freely available by a journal, or when the official, formatted version is published online, or when it arrives in printed hardcopy?

Now, this is an interesting question to ponder, but I think it’s only interesting from the standpoint of rules (e.g., codes governing nomenclature) and how we’re going to decide what counts. From the standpoint of moving science forward, the paper is published as soon as it is available for other researchers to use openly–i.e., not just to use in private in their own research, but also to cite. And since that’s the axis I care most about, I prefer to see accepted manuscripts made widely available as soon as possible, and I support APP’s policy. In the case of Yates et al. (2012), having the accepted manuscript online for the past year meant that it was available for Butler et al. (2012) to use, and cite, in their broad reassessment of pneumaticity in Triassic archosaurs. If our manuscript has not been published, that might not have been the case; Adam gave a talk on our project at the 2009 SVP in Bristol, but Butler et al. might have been loathe to cite an abstract, and some journals explicitly forbid it.

So I say bring it on. Let’s really accelerate research, by letting people see the content as early as possible. Making other researchers wait just so they can see a prettier version of the same information seems to me to be a triumph of style over science.

References

UPDATE April 16, 2012: The paper is officially published now. I’ve updated the citation and link below accordingly.

More new goodies:

Yates, A.M., Wedel, M.J., and Bonnan, M.F. 2012. The early evolution of postcranial skeletal pneumaticity in sauropodomorph dinosaurs. Acta Palaeontologica Polonica 57(1):85-100. doi: http://dx.doi.org/10.4202/app.2010.0075

This is only kinda sorta published. The accepted manuscript is now posted on the APP website, and it has a DOI, but it’s not formatted or available in print yet. But after discussing it amongst ourselves, we authors agreed that (1) the paper is globally available and it’s silly to pretend otherwise, (2) there are no nomenclatural ramifications of that fact, and (3) we’re tired of not being able to talk about this stuff. So we’re gonna, starting…now.

A brief tale of Serendipity in Science (TM):

Back in 2004 I was in my third year of grad school at Berkeley. My fellow grad student, Brian Kraatz, gave me a heads up about the 19th International Congress of Zoology coming up in Beijing. Attendees could submit 500-word abstracts or 2000-word short papers. I didn’t plan on doing either one, until the night before they were due, when I changed my mind and wrote almost all of what would become this paper in a single six-hour session (don’t be too impressed; I’ve been trying to replicate that feat for seven years with no success).

That summer, I met up with Brian in Beijing a week before the congress, and we spent the extra time working in the collections of the Institute of Vertebrate Paleontology and Paleoanthropology (IVPP). Paul Barrett was there, working on prosauropods, and he and I had some long and fascinating conversations. We also gave our talks in the same session at the congress. Paul must have decided I was not a complete moron because he invited me to give a talk in the basal sauropodomorph symposium at SVP in 2005.

A brief aside: many of the animals I grew up calling prosauropods ended up outside of the monophyletic Prosauropoda that is anchored on Plateosaurus. Some are now basal sauropods, some are closer to sauropods than to Plateosaurus but outside of Sauropoda, and some are outside of Prosauropoda + Sauropoda. The phylogenetically correct term encompassing all of the nonsauropods is  ‘basal sauropodomorphs’, and it means roughly what ‘prosauropods’ did until a decade or so ago. I often slip into informally using ‘prosauropods’, but I try to remember to put the term in quotes so as not to mislead anyone.

I had been to England in 2004 and 2005 and seen the putatively pneumatic vertebrae of Erythrosuchus and what was then known as Thecodontosaurus caducus (and is currently trading under the name Pantydraco caducus for reasons that it would be otiose, for the moment, to rehearse)–and, not incidentally, had finally met Mike in person, although we’d been corresponding since 2000. I’d also been to Stuttgart primarily to see the appendicular material of Janenschia and ended up spending some quality time with Plateosaurus. (Since the theme here is serendipity, note that the Janenschia work–my raison d’etre for going to Germany–died on the table, whereas I’ve now been an author on three ‘prosauropod’ papers and have more in the works. Weird!)

Anyway, with all of that accidental experience with ‘prosauropods’ and other interesting critters like Erythrosuchus, I found that I actually had something to say in 2005 SVP symposium. I titled my talk, ‘What pneumaticity tells us about “prosauropods”, and vice versa’, and it turned into the 2007 paper of the same title.

None of this would have happened if Brian hadn’t hounded me about going to Beijing, and if I hadn’t ended up talking so much with Paul on that trip, and if I hadn’t finished up with Janenschia on my first day in Stuttgart and spent the rest of the week playing with Plateosaurus. And so on. Science is unpredictable, especially for scientists.

When I sent around the PDF of the paper to friends and colleagues, I included this quip: “Were prosauropods pneumatic? The fossils don’t say. Somehow I stretched that out to 16 pages.” Mike claims that because of this quip he’s never been able to take that paper seriously. But it is my favorite among my solo efforts. It includes loads of stuff on the origins of air sacs and pneumaticity that I wasn’t able to get into my earlier papers, either because it wasn’t directly relevant or because some reviewer forced me to excise it.

BUT.

Almost immediately after the paper came out, Adam Yates and Matt Bonnan went and found roughly a zillion pneumatic ‘prosauropods’, which was a bit embarrassing since I’d just concluded that the evidence for ‘prosauropod’ pneumaticity was thin to nonexistent. So it is a damn good thing for me that I was already on friendly terms with both of them, because instead of taking the opportunity to smack me down, they invited me on board. Which led to Adam’s talk at SVP in Bristol in 2009, and to the new paper.

And actually, the depth of my incorrectness was even greater than I had thought. I reckon that literally millions of people have seen the mounted Plateosaurus skeleton in the AMNH, and any of them who have looked closely have seen this:

(Click for full size, unlabeled version.)

You see the problem here, I’m sure: the semi-big, semi-obvious fossa divided by an accessory lamina, not consistent with a muscle attachment point or fat pad or cartilage or infection, but very consistent in both form and location with the pneumatic fossae of other, more derived sauropodomorphs. On the lateral face of the vertebra, probably seen by millions, obvious to anyone who cares to look. A pneumatic prosauropod, in other words, right out in public for decades and decades (this time I don’t have to use the scare quotes because Plateosaurus actually IS a prosauropod sensu stricto). I didn’t even notice the first time I visited the AMNH back in 2006. I took the above photos, which are the basis for Figure 4 in the paper, in 2009.

So: ‘prosauropods’ were pneumatic. Some of them. A little bit. If you’d like to know more, please read the paper–it’s free.

Finally, a big thank-you to Adam and Matt for inviting me to be part of this. I think it’s pretty cool stuff, and I’m sure I’ll have more to say about it in the future. They might too–you should be reading their blogs, Dracovenator and Jurassic Journeys, anyway.

We’re still not done with Brontomerus, by the way. If nothing else, there’s the long-overdue post on how sauropod ilia change (or rather fail to change) through ontogeny. But that’s something we’ll have to get back to next week. Stay tuned.

Why we do mass estimates

Mass estimates are a big deal in paleobiology. If you want to know how much an animal needed in terms of food, water, and oxygen, or how fast it could move, or how many offspring it could produce in a season, or something about its heat balance, or its population density, or the size of its brain relative to its body, then at some point you are going to need a mass estimate.

All that is true, but it’s also a bit bogus. The fact is, people like to know how big things are, and paleontologists are not immune to this desire. We have loads of ways to rationalize our basic curiosity about the bigness of extinct critters. And the figuring out part is both very cool and strangely satisfying. So let’s get on with it.

Two roads diverged

There are two basic modes for determining the mass of an extinct animal: allometric, and volumetric. Allometric methods rely on predictable mathematical relationships between body measurements and body mass. You measure a bunch of living critters, plot the results, find your regression line, and use that to estimate the masses of extinct things based on their measurements. Allometric methods have a couple of problems. One is that they are absolutely horrible for extrapolating to animals outside the size range of the modern sample, which ain’t so great for us sauropod workers. The other is that they’re pretty imprecise even within the size range of the modern sample, because real data are messy and there is often substantial scatter around the regression line, which if faithfully carried through the calculations produces large uncertainties in the output. The obvious conclusion is that anyone calculating extinct-animal masses by extrapolating an allometric regression ought to calculate the 95% confidence intervals (e.g. “Argentinosaurus massed 70000 kg, with a 95% confidence interval of 25000-140000 kg), but, oddly, no-one seems to do this.

Volumetric methods rely on creating a physical, digital, or mathematical model of an extinct animal, determining the volume of the model, multiplying by a scale factor to get the volume of the animal in life, and multiplying that by the presumed density of the living animal to get its mass. Volumetric methods have three problems: (1) many extinct vertebrates are known from insufficient material to make a good 3D model of the skeleton; (2) even if you have a complete skeleton, the method is very sensitive to how you articulate the bones–especially the ribcage–and the amount of flesh you decide to pack on, and there are few good guidelines for doing this correctly; and (3) relatively small changes in the scale factor of the model can produce big changes in the output, because mass goes with the cube of the linear measurement. If your scale factor is off by 10%, you mass will be off by 33% (1.1^3=1.33).

On the plus side, volumetric mass estimates are cheap and easy. You don’t need hundreds or thousands of measurements and body masses taken from living animals; you can do the whole thing in your kitchen or on your laptop in the space of an afternoon, or even less. In the old days you’d build a physical model, or buy a toy dinosaur, and use a sandbox or a dunk tank to measure the volume of sand or water that the model displaced, and go from there. Then in the 90s people started building digital 3D models of extinct animals and measuring the volumes of those.

But you don’t need a physical model or a dunk tank or even a laptop to do volumetric modeling. Thanks to a method called graphic double integration or GDI, which is explained in detail in the next section, you can go through the whole process with nothing more than pen and paper, although a computer helps.

Volumetric methods in general, and GDI in particular, have one more huge advantage over allometric methods: they’re more precise and more accurate. In the only published study that compares the accuracy of various methods on extant animals of known mass, Hurlburt (1999) found that GDI estimates were sometimes off by as much as 20%, but that allometric estimates were much worse, with several off by 90-100% and one off by more than 800%. GDI estimates were not only closer to the right answers, they also varied much less than allometric methods. On one hand, this is good news for GDI afficionados, since it is the cheapest and easiest of all the mass estimation methods out there. On the other hand, it should give us pause that on samples of known mass, the best available method can still be off by as much as a fifth even when working with complete bodies, including the flesh. We should account for every source of error that we can, and still treat our results with appropriate skepticism.

Graphic Double Integration

GDI was invented by Jerison (1973) to estimate the volumes of cranial endocasts. Hurlburt (1999) was the first to apply it to whole animals, and since then it has been used by Murray and Vickers-Rich (2004) for mihirungs and other extinct flightless birds, yours truly for small basal saurischians (Wedel 2007), Mike for Brachiosaurus and Giraffatitan (Taylor 2009), and probably many others that I’ve missed.

GDI is conceptually simple, and easy to do. Using orthogonal views of a life restoration of an extinct animal, you divide the body into slices, treat each slice as an ellipse whose dimensions are determined from two perspectives, compute the average cross-sectional area of each body part, multiply that by the length of the body part in question, and add up the results. Here’s a figure from Murray and Vickers-Rich (2004) that should clarify things:

One of the cool things about GDI is that it is not just easy to separate out the relative contributions of each body region (i.e., head, neck, torso, limbs) to the total body volume, it’s usually unavoidable. This not only lets you compare body volume distributions among animals, it also lets you tinker with assigning different densities to different body parts.

An Example: Plateosaurus

Naturally I’m not going to introduce GDI without taking it for a test drive, and given my proclivities, that test drive is naturally going to be on a sauropodomorph. All we need is an accurate reconstruction of the test subject from at least two directions, and preferably three. You could get these images in several ways. You could take photographs of physical models (or toy dinosaurs) from the front, side, and top–that could be a cool science fair project for the dino-obsessed youngster in your life. You could use the white-bones-on-black-silhouette skeletal reconstructions that have become the unofficial industry standard. You could also use orthogonal photographs of mounted skeletons, although you’d have to make sure that they were taken from far enough away to avoid introducing perspective effects.

For this example, I’m going to use the digital skeletal reconstruction of the GPIT1 individual of Plateosaurus published by virtual dino-wrangler and frequent SV-POW! commenter Heinrich Mallison (Mallison et al 2009, fig. 14). I’m using this skeleton for several reasons: it’s almost complete, very little distorted, and I trust that Heinrich has all the bits in the right places. I don’t know if the ribcage articulation is perfect but it looks reasonable, and as we saw last time that is a major consideration. Since Heinrich built the digital skeleton in digital space, he knows precisely how big each piece actually is, so for once we have scale bars we can trust. Finally, this skeleton is well known and has been used in other mass estimate studies, so when I’m done we’ll have some other values to compare with and some grist for discussion. (To avoid accidental bias, I’m not looking at those other estimates until I’ve done mine.)

Of course, this is just a skeleton, and for GDI I need the body outline with the flesh on. So I opened the image in GIMP (still free, still awesome) and drew on some flesh. Here we necessarily enter the realm of speculation and opinion. I stuck pretty close to the skeletal outline, with the only major departures being for the soft tissues ventral to the vertebrae in the neck and for the bulk of the hip muscles. As movie Boromir said, there are other paths we might take, and we’ll get to a couple of alternatives at the end of the post.

This third image is the one I used for actually taking measurements. You need to lop off the arms and legs and tote them up separately from the body axis. I also filled in the body outlines and got rid of the background so I wouldn’t have any distracting visual clutter when I was taking measurements. I took the measurements using the measuring tool in GIMP (compass icon in the toolbar), in orthogonal directions (i.e., straight up/down and left/right), at regular intervals–every 20 pixels in this case.

One thing you’ll have to decide is how many slices to make. Ideally you’d do one slice per pixel, and then your mathematical model would be fairly smooth. There are programs out there that will do this for you; if you have a 3D digital model you can just measure the voxels (= pixels cubed) directly, and even if all you have is 2D images there are programs that will crank the GDI math for you and measure every pixel-width slice (Motani 2001). But if you’re just rolling with GIMP and OpenOffice Calc (or Photoshop and Excel, or calipers and a calculator), you need to have enough slices to capture most of the information in the model without becoming unwieldy to measure and calculate. I usually go with 40-50 slices through the body axis and 9 or 10 per limb.

The area of a circle is pi*r^2, and the area of an ellipse is pi*r*R, where r and R are the radii of the minor and major axes. So enter the widths and heights of the body segments in pixels in two columns (we’ll call them A and B) in your spreadsheet, and create a third column with the function 3.14*A1*B1/4. Divide by four because the pixel counts you measured on the image are diameters and the formula requires radii. If you forget to do that, you are going to get some wacky numbers.

One obvious departure from reality is that the method assumes that all of the body segments of an animal have elliptical cross-sections, when that is often not exactly true. But it’s usually close enough for the coarse level of detail that any mass estimation method is going to provide, and if it’s really eating you, there are ways to deal with it without assuming elliptical cross-sections (Motani 2001).

For each body region, average the resulting areas of the individual slices and multiply the resulting average areas by the lengths of the body regions to get volumes. Remember to measure the lengths at right angles to your diameter measurements, even when the body part in question is curved, as is the tail of Heinrich’s Plateosaurus.

For sauropods you can usually treat the limbs as cylinders and just enter the lateral view diameter twice, unless you are fortunate enough to have fore and aft views. It’s not a perfect solution but it’s probably better than agonizing over the exact cross sectional shape of each limb segment, since that will be highly dependent on how much flesh you (or some other artist) put on the model, and the limbs contribute so little to the final result. For Plateosaurus I made the arm circular, the forearm and hand half as wide as tall, the thigh twice as long as wide, and the leg and foot round. Don’t forget to double the volumes of the limbs since they’re paired!

We’re not done, because so far all our measurements are in pixels (and pixels cubed). But already we know something cool, which is what proportion each part of the body contributes to the total volume. In my model based on Heinrich’s digital skeleton, segmented as shown above, the relative contributions are as follows:

  • Head: 1%
  • Neck: 3%
  • Trunk: 70%
  • Tail: 11%
  • Forelimbs (pair): 3%
  • Hindlimbs (pair): 12%

Already one of the great truths of volumetric mass estimates is revealed: we tend to notice the extremities first, but really it is the dimensions of the trunk that drive everything. You could double the size of any given extremity and the impact on the result would be noticeable, but small. Consequently, modeling the torso accurately is crucial, which is why we get worried about the preservation of ribs and the slop inherent in complex joints.

Scale factor

The 170 cm scale bar in Heinrich’s figure measures 292 pixels, or 0.582 cm per pixel. The volume of each body segment must be multiplied by 0.582 cubed to convert to cubic cm, and then divided by 1000 to convert to liters, which are the lingua franca of volumetric measurement. If you’re a math n00b, your function should look like this: volume in liters = volume in pixels*SF*SF*SF/1000, where SF is the scale factor in units of cm/pixel. Don’t screw up and use pixels/cm, or if you do, remember to divide by the scale factor instead of multiplying. Just keep track of your units and everything will come out right.

If you’re not working from an example as perfect as Heinrich’s digital (and digitally measured) skeleton, you’ll have to find something else to use for a scale bar. Something big and reasonably impervious to error is good. I like the femur, if nothing else is available. Any sort of multi-segment dimension like shoulder height or trunk length is going to be very sensitive to how much gloop someone thought should go between the bones. Total length is especially bad because it depends not only on the intervertebral spacing but also on the number of vertebrae, and even most well-known dinos do not have complete vertebral series.

Density

Finally, multiply the volume in liters by the assumed density to get the mass of each body segment. Lots of people just go with the density of water, 1.0 kg/L, which is the same as saying a specific gravity (SG) of 1. Depending on what kind of animal you’re talking about, that may be a little bit off or it may be fairly calamitous. Colbert (1962) found SGs of 0.81 and 0.89 for an extant lizard and croc, which means an SG of 1.0 is off by between 11% and 19%. Nineteen percent–almost a fifth! For birds, it’s even worse; Hazlehurst and Rayner (1992) found an SG of 0.73.

Now, scroll back up to the diagram of the giant moa, which had a mass of 257.5 kg “assuming a specific gravity of 1”. If the moa was as light as an extant bird–and its skeleton is highly pneumatic–then it might have had a mass of only 188 kg (257.5*0.73). Or perhaps its density was higher, like that of a lizard or a croc. Without a living moa to play with, we may never know. Two points here: first, the common assumption of whole-body densities of 1.0 is demonstrably incorrect* for many animals, and second, since it’s hard to be certain about the densities of extinct animals, maybe the best thing is to try the calculation with several densities and see what results we get. (My thoughts on the plausible densities of sauropods are here.)

* Does anyone know of actual published data indicating a density of 1.0 for a terrestrial vertebrate? Or is the oft-quoted “bodies have the same density as water” basically bunk? (Note: I’m not disputing that flesh has a density close to that of water, but bones are denser and lungs and air spaces are lighter, and I want to know the mean density of the whole organism.)

Back to Plateosaurus. Using the measurements and calculations presented above, the total volume of the restored animal is 636 liters. Here are the whole body masses (in kg) we get using several different densities:

  • SG=1.0 (water), 636 kg
  • SG=0.89 (reptile high), 566 kg
  • SG=0.81 (reptile low), 515 kg
  • SG=0.73 (bird), 464 kg

I got numbers. Now what?

I’m going to describe three possible things you could do with the results once you have them. In my opinion, two of them are the wrong the thing to do and one is the right thing to do.

DON’T mistake the result of your calculation for The Right Answer. You haven’t stumbled on any universal truth. Assuming you measured enough slices and didn’t screw up the math, you know the volume of a mathematical model of an organism. If you crank all the way through the method you will always get a result, but that result is only an estimate of the volume of the real animal the model was based on. There are numerous sources of error that could plague your results, including: incomplete skeletal material, poorly articulated bones, wrong scale factor, wrong density, wrong amount of soft tissue on the skeleton. I saved density and gloop for last because you can’t do much about them; here the strength of your estimate relies on educated guesses that could themselves be wrong. In short, you don’t even know how wrong your estimate might be.

Pretty dismal, eh?

DON’T assume that the results are meaningless because you don’t know the actual fatness or the density of the animal, or because your results don’t match what you expected or what someone else got. I see this a LOT in people that have just run their first phylogenetic analysis. “Why, I could get any result I wanted just by tinkering with the input!” Well, duh! Like I said, the method will always give you an answer, and it won’t tell you whether the answer is right or not. The greatest advantage of explicit methods like cladistics and GDI is that you know what the input is, and so does everyone else if you are honest about reporting it. So if someone disagrees with your character coding or with how much the belly sags on your model sauropod, you can have a constructive discussion and hopefully science as a whole gets closer to the right answer (even if we have no way of knowing if or when we arrive, and even if your pet hypothesis gets trampled along the way).

DO be appropriately skeptical of your own results without either accepting them as gospel or throwing them out as worthless. The fact that the answer changes as you vary the parameters is a feature, not a bug. Investigate a range of possibilities, report all of those results, and feel free to argue why you think some of the results are better than others. Give people enough information to replicate your results, and compare your results to those of other workers. Figure out where yours differ and why.

Try to think of more interesting things you could do with your results. Don Henderson went from digitally slicing critters (Henderson 1999) to investigating floating sauropods (Henderson 2004) to literally putting sauropods through their paces (Henderson 2006)–not to mention working on pterosaur flight and swimming giraffes and other cool stuff. I’m not saying you should run out and do those exact things, but rather that you’re more likely to come up with something interesting if you think about what you could do with your GDI results instead of treating them as an end in themselves.

How massive was GPIT1, really?

Beats me. I’m not the only one who has done a mass estimate based on that skeleton. Gunga et al. (2007) did not one but two volumetric mass estimates based on GPIT1, and Mallison (2010) did a whole series, and they published their models so we can see how they got there. (In fact, many of you have probably been reading this post in slack-jawed horror, wondering why I was ignoring those papers and redoing the mass estimate the hard way. Now you know!) I’m going to discuss the results of Gunga et al. (2007) first, and come back to Mallison (2010) at the end.

Here’s the “slender” model of Gunga et al. 2007 (their fig. 3):

and here’s their “robust” model (Gunga et al. 2007:fig. 4):

(These look a bit…inelegant, let’s say…because they are based on the way the physical skeleton is currently mounted; Heinrich’s model looks much nicer because of his virtual remount.)

For both mass estimates they used a density of 0.8, which I think is probably on the low end of the range for prosauropods but not beyond the bounds of possibility. They got a mass of 630 kg for the slender model and 912 kg for the robust one.

Their 630-kg estimate for the slender model is deceptively close to the upper end of my range; deceptive because their 630-kg estimate assumes a density of 0.8 and my 636-kg one assumes a density of 1.0. The volumes are more directly comparable: 636 L for mine, 790 L for their slender one, and 1140 L for their robust one. I think that’s pretty good correspondence, and the differences are easily explained. My version is even more skinnier than their slender version; I made it about as svelte as it could possibly have been. I did that deliberately, because it’s always possible to pack on more soft tissue but at some point the dimensions of the skeleton establish a lower bound for how voluminous a healthy (i.e., non-starving) animal could have been. The slender model of Gunga et al. (2007) looks healthier than mine, whereas their robust version looks, to my eye, downright corpulent. But not unrealistically so; fat animals are less common than skinny ones but they are out there to be found, at least in some times and places. It pays to remember that the mass of a single individual can fluctuate wildly depending on seasonal food availability and exercise level.

For GPIT1, I think something like 500 kg is probably a realistic lower bound and 900 kg is a realistic upper bound, and the actual mass of an average individual Plateosaurus of that size was somewhere in the middle. That’s a big range–900 kg is almost twice 500 kg. It’s hard to narrow down because I really don’t know how fleshy Plateosaurus was or what it’s density might have been, and I feel less comfortable making guesses because I’ve spent much less time working on prosauropods than on sauropods. If someone put a gun to my head, I’d say that in my opinion, a bulk somewhere between that of my model and the slender model of Gunga et al. is most believable, and a density of perhaps 0.85, for a result in the neighborhood of 600 kg. But those are opinions, not hypotheses, certainly not facts.

I’m happy to see that my results are pretty close to those of Mallison (2010), who got 740 L, which is also not far off from the slender model of Gunga et al. (2007). So we’ve had at least three independent attempts at this and gotten comparable results, which hopefully means we’re at least in the right ballpark (and pessimistically means we’re all making mistakes of equal magnitude!). Heinrich’s paper is a goldmine, with loads of interesting stuff on how the skeleton articulates, what poses the animal might have been capable of, and how varying the density of different body segments affects the estimated mass and center of mass. It’s a model study and I’d happily tell you all about it but you should really read it for yourself. Since it’s freely available (yay open access!), there’s no barrier to you doing so.

Conclusion

So: use GDI with caution, but do use it. It’s easy, it’s cool, it’s explicit, it will give you lots to think about and give us lots to talk about. Stay tuned for related posts in the not-too-distant future.

References

I Cannot Brain Today, I Have the Dumb

Man, I hate making mistakes. The only thing worse than making mistakes is making them in public, and the only thing worse than that is finding them in published papers when it’s too late to do anything about them. About the only consolation left–if you’re lucky–is getting to be the one to rat yourself out (we have to do this a lot). So here goes.

fig4-head-and-neck-angles 480

Neck angle FAIL

In our figure 4 (from Taylor et al. 2009) we showed the skulls of three sauropodomorphs, Massospondylus, Camarasaurus, and Diplodocus, posed with horizontal semicircular canals (HSCCs) level, angled 30 degrees above horizontal, and angled 20 degrees below horizontal, as it is written (by Duijm 1951). We also showed the angle of the occipital condyle when the HSCCs are level; if the craniocervical joint was in osteologically neutral pose (ONP), that line would indicate the angle of the anterior cervicals.

Trouble is, we put the neck lines for Diplodocus and Camarasaurus in the wrong places.

As any idiot can see from Sereno et al. (2008: fig 1), the brain, brainstem, and occipital condyle form a line that runs from roughly the upper part of the orbit (in lateral see-through view) out the back of the head. Now if you look at our fig. 4 you’ll see that the ONP lines for Camarasaurus and Diplodocus are much too inclined, so that if the brain was in line with the anterior neck–which it should be, in ONP–it would be sticking out the back of the head.

If that doesn’t make sense, just look at the above illustration, imagine the brain and spinal cord in a straight line parallel to the black neck line but also dorsal to it, and you’ll see that the brain would be outside the skull. Those incorrect neck lines don’t represent impossible postures, but they don’t represent ONP, either.

Sauropodomorph head figure redone 480

Taxonomic variation WIN!

Here’s a corrected up version of the figure to show what I mean. The black lines are still the ONP neck lines, and now I’ve put in shadowy necks at +30 and -20 to go with the shadowy heads. The 50 degree spans marked out by the shadowy necks are the ranges within which the neck could articulate in ONP with skulls stuck in the 50-degree “Duijm window”.

Caution: it is very easy to misread the shadowy necks as showing a range of movement within an individual; in fact, the neck lines are ‘anchored’ to the skulls in ONP as the skulls rotate through the 50 degrees allowed by the HSCCs. They are not individual movement but the possible range of taxonomic variation in HSCC orientation according to Duijm (1951).

Worth noting here is the likelihood that Massospondylus had a more elevated neck than any of the neosauropods studied so far–certainly a finding at odds with the traditional depictions of basal sauropodomorphs. (It is just a likelihood, though, since the top, neck-wise, of Massospondylus‘s Duijm window overlaps with the windows of the other taxa a bit.)

Nigersaurus, buddy, why so down?

Nigersaurus, buddy, why so down?

In this version I’ve gone one step farther and included Nigersaurus (modified from Sereno et al. (2008: fig 1). Nigersaurus differs from Diplodocus in the angle of the face from the HSCCs and occipital condyle, not in the angle between the HSCCs and the occipital condyle, which is remarkably similar in Camarasaurus, Diplodocus, and Nigersaurus. This suggests that Nigersaurus held its head differently than other sauropods, but not necessarily its neck.

Keep in mind, though, that the difference in facial angle between Diplodocus and Nigersaurus is less than 50 degrees, and that some of the head postures in the respective Duijm windows of the two taxa are identical. So we can’t say for certain that Nigersaurus held its head differently than Diplodocus; it is possible that they held their heads at the same angle and that Nigersaurus just carried its HSCCs at a different angle. If that were the case, the neck of Nigersaurus would have been more inclined than that of Diplodocus. I’m not arguing that that’s likely–it seems perfectly plausible that the two taxa might have held their necks similarly and their heads differently, as suggested above–I’m just pointing out the very wide range of possibilities allowed by the data. To reiterate one of the points of the paper, HSCCs aren’t useless for determining habitual head posture, they just can’t narrow things down very far on their own.

Also note that some of the neck postures allowed by the Duijm window have the anterior cervicals running down, below horizontal, not up. And many of the allowed neck postures for the neosauropods are close to horizontal. So, we were wrong and HSCCs + occipital condyles show that most sauropods held their necks close to level and not strongly elevated after all, right?

Onward and Upward, or Down in Flames?

Not so fast. Remember that all of the neck lines in the above figures show the angle of the anterior neck if the neck was in ONP with the skull. But Vidal et al. (1986) found that the skull is habitually flexed on the neck, even in lizards, and we have since verified this for salamanders, turtles, and more. And sometimes the flexion is dramatic.

Our figure 1 (from Taylor et al. 2009) shows the cranium, cervicals, and first few dorsals from a hare in ONP and in the posture shown by Vidal et al. (1986: fig. 4b). The difference between the anteriorly-directed ONP pose and the backward-leaning Vidal-compliant pose is striking. I measured the angle between the cervical column and the maxillary toothrow to be ~110 degrees in the ONP pose and ~70 degrees in the Vidal-compliant pose (try it yourself with Paint or Photoshop, or download some free image manipulation software). That means the head is flexed on the neck by 40 degrees! That is a big angle. If sauropods did the same, you could take the neck lines shown above and crank them down by 40 degrees (remember that the heads are “fixed” into the 50-degree Duijm windows allowed by the HSCCs), which would make Mike’s elevated Diplodocus look not just achievable, but perhaps even conservative.

Where does all that leave us? In sauropods for which HSCC orientation is known, putting the HSCCs level the anterior neck is still inclined, and even with the HSCCs angled 20 degrees down the ONP neck would only be slightly below horizontal, and if the head was Vidal-compliant (strongly flexed on the neck), the neck would have to be above horizontal. So heads still tell us about necks, and in particular they tell us that the necks angled up. Our neck lines for Camarasaurus and Diplodocus are not correct for ONP, but probably represent attainable postures. My first head ‘n necks post has the angles too exaggeraged for ONP, too, but again all of those poses are not just possible but likely if the head was flexed on the neck.

Miscellanea

We owe mad props to Brian Engh, a.k.a. The Historian, who burst on the paleo-rap scene with a rap video about crocodilian predation and almost certainly the first ever kung-fu rap video to name-check titanosaurs. Brian stumbled across Mike’s extra goodies page for the new paper about week before the paper was due out, and kindly suppressed the information until after D-Day. You can and should download his entire album, Earth Beasts Awaken (open access, yo), and kick it old school.

Congratulations to Francisco “Paco” Gasco, who just got funding for a PhD to do a complete morphological and paleobiological workup on the giant Spanish sauropod Turiasaurus. You’ll be hearing more about Paco in the not-too-distant future, we promise.

Finally, here’s that video of an elephant grabbing an ostrich by the neck that you ordered.

ostrichvselephant

The End of the Beginning?

This brings us to the end of ten solid days of new posts, which is a new record for us and one not likely to be broken for a long time, if ever. We never planned to do all this; in the beginning we each were going to contribute one post and that would have been that. But we kept finding things that we felt needed to be discussed.

As all of us have been saying in every available medium, this is not the end of anything. The sauropod neck posture debate is not over; in a few years we may look back and see that in 2009 we were still stumbling to the real starting line. We don’t think this stuff is unimportant or unknowable, and we’re going to keep working on it, and we hope lots of others do as well.

We’ll see you out there.

Ridem dino 480

Up, boy, up! Heyaaah!!

References

In case you haven’t heard, Taylor et al. (2009) recently argued that sauropods naturally held their cervico-dorsal junctions in extension, and their cranio-cervical joints in flexion… at least, when they weren’t foraging, feeding or engaged in other such activities [if you need help with those terms please see the Tet Zoo article here].

neck-posture_montage

Given that we here at SV-POW! are predominantly interested in sauropods, and given that the amazing necks of these animals have long been such a source of debate, it stands to reason that sauropods might get used as the ‘poster children’ or exemplars for any particular argument about neck pose in fossil tetrapods. However, as we’ve said here and there – I certainly mentioned it in my Tet Zoo article on the subject – the contention (that cervico-dorsal junctions are maintained in extension, and that cranio-cervical joints are maintained in flexion) holds true for all terrestrial amniotes and, to a degree, all crown-group tetrapods. In this article we’re going to do something a little odd for SV-POW! – we’re going to look at other fossil amniotes to see if and how this affects them. Have any of them also been reconstructed in poses that are not compliant with the data from living animals?

The short answer is yes, yes they have.

First off, fossil mammals mostly get by ok, which is what you’d expect given that they are generally very similar to their extant relatives. Likewise, there aren’t any fossil birds that have been reconstructed incorrectly, and again you’d hope not given that they’re generally highly similar to extant forms. The extinct moa from New Zealand (that’s moa in the plural sense) are sometimes shown standing at rest with non-extended cervico-dorsal junctions, but with extremely strong extension in the anterior part of the neck that makes up for this (Worthy & Holdaway 2002). While extension at the cervico-dorsal junction may be subtle or absent in living ratites when they are feeding or foraging, in relaxed individuals extension at the neck base is indeed present.

Stegosaurs really need a makeover

What about other dinosaurs? Here’s where we do find quite a few reconstructions that contradict our contention. For a start, basal sauropodomorphs – the animals conventionally lumped together as prosauropods – have often been shown with non-extended cervico-dorsal junctions and fully extended cranio-cervical junctions: that is, with necks that emerge in a straight line from the body, and heads that have their long axis parallel to that of the neck. Classic examples include Kermack’s reconstruction of the animal formerly known as Thecodontosaurus and Weishampel & Westphal’s Plateosaurus. There are many others.

Historically, non-avian theropods have been depicted with elevated necks, flexed cranio-cervical junctions and all that. So far so good. One specific exception does come to mind however: Tarsitano (1983) produced a truly awful theropod reconstruction in which the neck was shown as straight and with a non-extended cervico-dorsal junction. The latter is a no-no, and so is straightening the neck this much, as the shapes of the centra and neural arches show that the cervical vertebrae of theropods were held elevated and in a gentle S-curve (see Molnar & Farlow 1990). A few artistic reconstructions of non-avian theropods have given them non-elevated necks (Neave Parker’s megalosaur picture from the 1970s comes to mind), and if you look at the allosaurs that featured in Walking With Dinosaurs you’ll note that their cranio-cervical junctions are extended, not flexed as they should be.

Tarsitano_1983_theropod_posture

Theropod posture as reconstructed by Tarsitano (1983). Tarsitano mostly argued that non-avian theropods were more like crocodilians than birds in musculature and some aspects of posture.

On to ornithischians. It was difficult to keep a tight lip back in February 2009 when the long-necked stegosaur Miragaia longicollum was published. Like the WWD diplodocoids, Miragaia was given a non-extended cervico-dorsal junction and extended cranio-cervical junction: in other words, its neck and head were illustrated projecting forwards in a straight line, as a continuation of the animal’s dorsal column (Mateus et al. 2009). Based on what we know about living animals, it’s more likely that the cervico-dorsal junction was extended, and that the cranio-cervical junction was flexed: in other words, that the neck was strongly elevated relative to the dorsal vertebrae, and that the head was held at an angle to the neck. Given the remarkable length of its neck, this at least makes it possible that Miragaia was a high-browser. I look forward to seeing artistic reconstructions that show this animal with its head held up above its back, rather than extending forwards and parallel to it (actually, I’ve already seen two, but you know what I mean).

In fact, like sauropods, stegosaurs have been flat out abused by palaeontologists, with non-extended cervico-dorsal junctions and fully extended cranio-cervical junctions being the norm across more than 100 years of description and reconstruction. And don’t use the excuse that these reconstructions are all meant to show the animals engaged in feeding or foraging: they’re not. Many of them clearly depict the animals standing, in relaxed poses, and doing nothing. Marsh started it in 1891: he showed the skull of Stegosaurus armatus (then S. stenops) fully extended, rather than flexed, on the neck, and showed the neck continuing in (approximately) a straight line from the dorsals. This reconstruction was hugely influential, of course, and even today the popular conception of the stegosaur – with its horrible over-arched back and down-sloping tail – is based on Marsh’s drawing. Later stegosaur reconstructions by Lull and Gilmore perpetuated the idea of non-extended cervico-dorsal junctions and fully extended cranio-cervical junctions in Stegosaurus (see Czerkas 1987 for a review of stegosaur life reconstructions), and the same posture was later reconstructed for Huayangosaurus, Kentrosaurus, Tuojiangosaurus and others (see Galton & Upchurch 2004).

Tuojiangosaurus_skeleton_NHM

Tuojiangosaurus, as displayed in the Natural History Museum, London. (c) NHM (image from wikipedia). Note the lack of extension at the cervico-dorsal junction and the slight hyper-extension at the cranio-cervical junction.

Those stegosaur reconstructions you can see in some museums – some of which show the cranio-cervical junction in slight hyper-extension (look at the Tuojiangosaurus shown here) – are flat-out horrible and totally contradict the data we have from neck and head posture in extant amniotes (Taylor et al. 2009). In recent decades, reconstructions by artists like Stephan Czerkas and Greg Paul have given stegosaurs raised necks where the cervico-dorsal junction is extended in proper fashion (as per the data from living amniotes). I get the impression, however, that such reconstructions have not been taken seriously by ‘mainstream’ palaeontologists, at least some of whom still seem to think that stegosaurs walked around with their heads two inches off the ground.

Similar mistakes have been made with ankylosaurs: most classic reconstructions show non-extended cervico-dorsal junctions where the neck emerges in a straight line or even slopes downwards, and cranio-cervical junctions that are in full extension. This goes for Ken Carpenter’s Euoplocephalus [shown in composite above] and Sauropelta, Richard Lull’s Nodosaurus, and others (Lull 1921, Carpenter 1982, 1984). Again, the reconstructions that show these neck and head postures do not definitely show the animals in feeding, foraging or searching postures: they are meant to depict the ‘normal’ (viz, relaxed) pose for the animal. A gently elevated neck with an extended cervico-dorsal junction and a flexed cranio-cervical junction is, again, what we should expect given what living animals do, and this has been correctly portrayed by some.

Other ornithischians have generally been reconstructed accurately (at least as goes neck and head posture), but there are, however, a few ceratopsian reconstructions showing non-extended cervico-dorsal junctions and fully extended cranio-cervical junctions. Granger & Gregory (1923) reconstructed Protoceratops in this manner, for example, and ceratopsids have sometimes been shown this way too (Lull 1933). Again, the reconstructions I’m referring to are meant to show normal, relaxed poses, rather than feeding or foraging poses, so criticism is justified. Putting extension into the ceratopsian cervico-dorsal junction raises the head somewhat, such that the top of the frill is now higher than the top of the back rather than lower than it. Notably, some articulated skeletons are displayed this way: the Centrosaurus panel-mount AMNH 5351, shown here, is one of the best examples. Lull thought that the neck had been elevated too much and that the neck posture ‘is that of death rather than that of life’! Peter Galton’s Hypsilophodon (which has mostly been superseded by Greg Paul’s reconstruction these days anyway) should also be considered suspect in view of the strongly extended cranio-cervical junction (Galton 1971, 1974), but the animal was clearly meant to be running at speed, so you could argue that it was shown holding its head and neck in a decidedly un-relaxed pose.

AMNH_Centrosaurus_oblique_June-2009

The excellent Centrosaurus specimen AMNH 5351. Photo borrowed from Traumador the Tyrannosaur. Thanks, Traumador :)

I should point out again at this point that our contention (that cervico-dorsal junctions should be shown in extension in a relaxed animal, and cranio-cervical junctions should be shown in flexion in a relaxed animal) is a hypothesis. It’s possible (unlikely perhaps, but possible) that some stegosaurs, or ankylosaurs, or therapsids, or whatever, did some funky stuff with their occipital condyles or vertebrae and evolved a relaxed head and neck posture different from that of living amniotes, and indeed (as we’ll see in a moment) there surely are at least some exceptions within Amniota. However, if you think a given animal represents a special case, you’re gonna have to demonstrate it.

Shock horror, marine reptiles on SV-POW!

Elsewhere among Reptilia, the data from living lizards and crocodilians indicates that, generally speaking, we should expect fossil forms to hold their necks elevated at moderate angles of between 20-40° relative to the dorsal column when in normal, relaxed pose. Many fossil, non-dinosaurian archosaurs (like rauisuchians and aetosaurs) have been reconstructed this way (mostly because people have looked at living crocodilians when reconstructing these animals), as have fossil squamates and many others.

large_Nicholson_plesiosaur_June-2009

A very old reconstruction of the Jurassic plesiosaur Plesiosaurus.

We do, however, have a contradiction of sorts when we come to sauropterygians (the plesiosaurs and their relatives). Reconstructions of plesiosaurs have evolved in similar fashion to those depicting sauropods: some old reconstructions (some, not all) depict them with extended cervico-dorsal junctions and flexed cranio-cervical junctions (such reconstructions typically show the animals sticking their necks well up out of the water and peering around) (e.g., Williston 1914), but many others show them with non-extended cervico-dorsal junctions and fully extended cranio-cervical junctions. In other words, with the neck and head continuing in a straight line from the dorsal column.

Denver_elasmosaur_June-2009

The elasmosaurid Thalassomedon haningtoni, as displayed at the Denver Museum of Nature and Science. Check out the big neural spines.

Contradicting the idea that plesiosaurs looked down on their prey from above is the fact that their orbits often face slightly or strongly upwards, and there are also indications from their narial and ear anatomy that they were specialised for detecting sensory cues in water, not in air (Cruickshank et al. 1991, Storrs & Taylor 1996). Furthermore, their high-density, often pachyostotic skeletons indicate that they were negatively buoyant animals that were trying their hardest to stay submerged and beneath the surface. All of this indicates that plesiosaurs were subaqueous predators that mostly kept their necks and heads beneath the surface of the water (except when breathing). This makes it very unlikely that their necks were elevated, and indeed – in strong contrast to sauropods and other dinosaurs – there are indications from their vertebral anatomy that neck elevation was not possible in the group (in elasmosaurs, for example, the neural spines on both the cervicals and dorsals are tall and sub-rectangular and it’s difficult to imagine how this would have allowed anything more than extremely subtle extension at the cervico-dorsal junction). So I am going to go out on a limb here (or, more accurately, I’m going to agree with everyone who works on plesiosaurs) and say that plesiosaurs did not hold their necks in the same manner as the extant amniotes that we looked at (Taylor et al. 2009). Is this because they were aquatic, and hence not under the same gravitational constraints as terrestrial amniotes? That looks likely, but we really need to thrash this out once and for all: further work on this is obviously needed, and perhaps it will appear soon. I know from many discussions that plesiosaur researchers talk as much about long necks as sauropod researchers do.

Finally – dicynodonts and other synapsids

Moving now well away from dinosaurs and archosaurs and even reptiles, I’ve had non-mammalian synapsids on my mind an awful lot during all of this. While many of them have relatively short necks, members of some groups have still been shown in downright unlikely postures. Dinocephalians have consistently been shown (correctly) with extended cervico-dorsal junctions and flexed cranio-cervical junctions, so those reconstructions of such things as Titanophoneus and Moschops with their necks held high and their heads at an angle are correct based on the data from living amniotes. However, some reconstructions of some caseids (Stovall’s Cotylorhynchus), dicynodonts (I’m looking at you, Watson’s rendition of Lystrosaurus [shown at top of composite image used above] and Cluver’s Cistecephalus) and gorgonopsids (Colbert’s Lycaenops, for example; shown below) have the cranio-cervical junction in extended or even hyper-extended pose, which again is a total no-no unless there is evidence to the contrary. While some of these animals have been reconstructed in walking or running poses (and hence might be holding their necks and heads in special searching or foraging poses), plenty of others are shown standing on all fours, in relaxed, ‘normal’ poses, so their unusual neck and head poses are, we can assume, meant to be the relaxed, ‘normal’ poses (further examples include King’s Dicynodon and Dinodontosaurus).

Lycaenops_ornatus_AMNM_June-2009

The relatively short necks of these animals mean that, even with the cervico-dorsal junction in full extension, the neck is only elevated by a slight and thoroughly believable 20-40° relative to the dorsal column. Similarly, showing the cranio-cervical junction in flexion is no big deal, as all it does is rotate the skull such that its long axis is at an angle to the neck, rather than acting as a straight-line continuation of it. It seems that more extreme cervico-dorsal extension and cranio-cervical flexion evolved within Mammalia, and hence that non-mammalian synapsids were more like other ‘average’ amniotes in head and neck posture. Nevertheless – again – reconstructions that show the neck and head as straight-line extensions of the back should be considered inconsistent with what we know of neck and neck posture in living amniotes.

Final thoughts

We really hope that our paper will inspire some much-needed debate, and instigate some new work. As you’ll know if you’ve been following the comments on blogs and such, and the media coverage we’ve been getting, there’s every indication that this is exactly what will happen. But what makes this work of particular interest to people in general – and not just to specialists who spend their time worrying about cervical rib morphology and its correlation with functional morphology, or whether the bifurcate neural spines of some sauropods are homologous with the single neural spines of others, and so on – is that it has a real and obvious effect on the life appearance of a fossil animal. And, as I’ve tried to show here, our hypothesis extends beyond the limits of Sauropoda. Stegosaurs and dicynodonts need never look the same way again.

References

  • Carpenter, K. 1982. Skeletal and dermal armor reconstruction of Euoplocephalus tutus (Ornithischia: Ankylosauridae) from the Late Cretaceous Oldman Formation of Alberta. Canadian Journal of Earth Sciences 19, 689-697.
  • Carpenter, K. 1984. Skeletal reconstruction and life restoration of Sauropelta (Ankylosauria: Nodosauridae) from the Cretaceous of North America. Canadian Journal of Earth Sciences 21, 1491-1498.
  • Cruickshank, A. R. I., Small, P. G. & Taylor, M. A. 1991. Dorsal nostrils and hydrodynamically driven underwater olfaction in plesiosaurs. Nature 352, 62-64.
  • Czerkas, S. A. 1987. A reevaluation of the plate arrangement on Stegosaurus stenops. In Czerkas, S. J. & Olson, E. C. (eds) Dinosaurs Past and Present, Volume II. Natural History Museum of Los Angeles County/University of Washington Press (Seattle and Washington), pp. 82-99.
  • Galton, P. M. 1971. Hypsilophodon, the cursorial non-arboreal dinosaur. Nature 231, 159-161.
  • Galton, P. M. 1974. The ornithischian dinosaur Hypsilophodon from the Wealden of the Isle of Wight. Bulletin of the British Museum (Natural History) 25, 1-152.
  • Galton, P. M. & Upchurch, P. 2004. Stegosauria. In Weishampel, D. B., Dodson, P. & Osmólska, H. (eds) The Dinosauria, Second Edition. University of California Press (Berkeley), pp. 343-362.
  • Granger, W. & Gregory, W. K. 1923. Protoceratops andrewsi, a pre-ceratopsian dinosaur from Mongolia. American Museum Novitates 72, 1-9.
  • Lull, R. S. 1921. The Cretaceous armored dinosaur, Nodosaurus textilis Marsh. American Journal of Science 1, 97-126.
  • Lull, R. S. 1933. A revision of the Ceratopsia or horned dinosaurs. Memoirs of the Peabody Museum of Natural History 3, 1-175.
  • Mateus, O., Maidment, S. C. R. & Christiansen, N. A. 2009. A new long-necked ‘sauropod mimic’ stegosaur and the evolution of the plated dinosaurs. Proceedings of the Royal Society of London, Series B (doi:10.1098/rspb.2008.1909)
  • Molnar, R. E & Farlow, J. O. 1992. Carnosaur paleobiology. In Weishampel, D. B., Dodson, P. & Osmólska, H. (eds) The Dinosauria. University of California Press (Berkeley), pp. 210-224.
  • Storrs, G. W. & Taylor, M. A. 1996. Cranial anatomy of a new plesiosaur genus from the lowermost Lias (Rhaetian/Hettangian) of Street, Somerset, England. Journal of Vertebrate Paleontology 16, 403-420.
  • Tarsitano, S. F. 1983. Stance and gait in theropod dinosaurs. Acta Palaeontologica Polonica 28, 251-264.
  • Taylor, M. P., Wedel, M. J. & Naish, D. 2009. Head and neck posture in sauropod dinosaurs inferred from extant animals. Acta Palaeontologica Polonica 54, 213-220.
  • Williston, S. W. 1914. Water Reptiles of the Past and Present. University of Chicago Press, Chicago.
  • Worthy, T. H., Holdaway, R. N. 2002. The Lost World of the Moa. Indiana University Press, Bloomington, Indiana.