Norwescon 41 Guests of Honor: Ken Liu, Galen Dara, and, er, me. Mike would like to remind you that you can get your own ‘Kylo Stabbed First’ t-shirt here.

The week before last I was fortunate to be the Science Guest of Honor at Norwescon 41 in Seattle (as threatened back when). I had a fantastic time. I got to give talks on binocular stargazing and the sizes of the largest sauropods and whales (ahem), participate on panels on alien biology and creature drawing, and meet a ton of cool people, including my fellow Guests of Honor, multiple-award-winning author Ken Liu and multiple-award-winning artist Galen Dara, both of whom turned out to be humble, easygoing, regular folks (if frighteningly talented).

I also had a lot of great conversations with folks who were attending the con, which is exactly what I wanted. One of the most interesting was a hallway conversation with a fellow DM named Shawn Connor. He had a great question for me, which I liked so much I wanted to answer it here on the blog. Here’s his question, copied with permission from a follow-up email:

I run tabletop RPGs, and in my current game one of the characters is a caveman type who naturally grew up hunting dinosaurs. As one does. His weapon is a dinosaur bone, customized and used as a club. I have attached the picture that he came up with [below]. Now understanding the picture is obviously not of a real dinosaur bone – it’s probably a chicken bone or a cow bone or something – let’s assume for the sake of this exercise that it is and that it is four feet long stem to stern. Given that, two questions: discounting the extra bling attached how heavy would such a bone be, and what kind of dinosaur could it have come from?

I’m going to answer those questions out of order. Advance warning: this will be a loooong post that will go down several rabbit holes that are likely of more intense interest to me, personally, than to anyone else on the planet. Read on at your own risk.

Whose femur is in the image?

First, Shawn is correct in noting that the femur in the image provided by his player is not a dinosaur femur. The prominent trochanters and spherical head offset on a narrow neck clearly make it a mammal femur, and if it’s four feet long, it could only have come from an elephant or an indricothere. Or a giant humanoid, I suppose, which is what the anatomy of the bone in the image most closely resembles. (It also appears to be foreshortened to make the distal end look bigger, or deliberately distorted to enhance the clubby-ness.)

Mounted elephant at the Museum of Osteology in Oklahoma City, with Tyler Hunt for scale.

But let’s play along and assume it’s from a non-human mammal. How big? Back in 2016 I was fortunate to get to measure most of the mounted large mammal skeletons at the Museum of Osteology in Oklahoma City, along with Tyler Hunt, then a University of Oklahoma undergrad and now finishing up his MS thesis under my mentor, Rich Cifelli.* The mounted elephant at the Museum of Osteology has a shoulder height of 254 cm (8 ft, 4 in) and a femur length of 102 cm (3 ft, 4 in). Assuming isometric scaling, a world record elephant with a shoulder height of 366 cm (12 ft) would have a femur length of 147 cm (4 ft, 10 in). So a four-foot (122 cm) femur would belong to an elephant roughly in the middle of that range, about ten feet (3 m) tall at the shoulder. That’s the size of the big bull elephant mounted at the Field Museum in Chicago.

The big mounted bull elephant at the Field Museum is 10 feet tall at the shoulder and weighed 6 tons in life. Note Mike for scale on the lower right. He and the elephant are about equidistant from the camera, so he should make a roughly accurate scale bar. Photo from our visit in 2005!

* Two further notes: first, I have roughly a zillion awesome photos from that 2016 visit to the Museum of Osteology, both of the specimens and of Tyler and me measuring them – not having posted them yet is one of the things I was whingeing about in the post that kicked off our return-to-weekly-posting thing this year. And second, I owe a belated and public thanks to the folks at the Museum of Osteology for accommodating Tyler and me. They helped us with ladders and so on and basically gave us free rein to play with collect data from their mounted skeletons, which was incredibly generous and helpful, and fortunately reflects the pro-research and pro-researcher attitude of most museums.

Which dinos had four-foot femora?

As for what kind of dinosaur a four-foot femur could have come from, we can rapidly narrow it down to a handful of clades: sauropods, ornithopods, theropods, and stegosaurs.

  • Sauropods. The longest complete femora of Patagotitan are 238 cm (7 ft, 10 in; Carballido et al. 2017), and an incomplete femur of Argentinosaurus has an estimated complete length of 250 cm (8 ft, 2 in; Mazzetta et al. 2004). So a four-foot femur would not be from a particularly large sauropod – something about elephant-sized, as you might expect from the elephant comparison above. Our old friend Haplocanthosaurus will fit the bill, as we’ll see in a bit.
  • Ornithopods. Femora of 172 cm (5 ft, 8 in) are known for the hadrosaurs Shantungosaurus (Hone et al. 2014) and Huaxiaosaurus (Zhao and Li 2009), and Zhao et al. (2007) reported a 170 cm (5 ft, 7 in) femur for Zhuchengosaurus (Huaxiaosaurus and Zhuchengosaurus may be junior synonyms of Shantungosaurus). But those are all monsters, well over 10 metric tons in estimated mass. So a four-foot femur would be from a large but not insanely large hadrosaur.

Mmmmmm…suffering. OM NOM NOM NOM!!

  • Theropods. Among the largest theropods, the holotype of Giganotosaurus has a femur length of 143 cm (4 ft, 8 in; Coria and Salgado 1995), and ‘Sue’ the T. rex (a.k.a. FMNH PR2081) has a right femur 132 cm long (4 ft, 4 in; Brochu 2003). So a four-foot femur from a theropod would definitely be from one of the monsters. The femur of Saurophaganax was 113.5 cm long (Chure 1995), just under four feet, which I only note as an excuse to use the above photo, which I adore.
  • Stegosaurs. I don’t know the longest femur that has been recovered from a stegosaur, but getting in the ballpark is easy. NHMUK PV R36730 has a femur 87 cm long, and the whole animal was approximately 6 m long (Maidment et al. 2015). Partial bits and bobs of the largest stegosaurs suggest animals about 9 m long, implying a femur length of about 130 cm (4 ft, 3 in), or just over the line.

I think that’s it. I don’t know of any ceratopsians or ankylosaurs with femora long enough to qualify – I assume someone will let me know in the comments if I’ve forgotten any.

How much would a four-foot femur weigh?

There are a couple of ways to get to the answer here. One is to use Graphic Double Integration, which is explained in this post.

Limb bones are not solid – in terrestrial tetrapods there is virtually always a marrow cavity of some sort, and in marine tetrapods the limb bones tend to be cancellous all the way through. Estimating the mass of a limb bone is a lot like estimating the mass of a pneumatic bone: figure out the cross-sectional areas of the cortex and marrow cavity (or air space if the bone is pneumatic), multiply by the length of the element to get volumes, and multiply those volumes by the density of the materials to get masses. I piled up all the relevant numbers and formulas in Tutorial 24, a move that has frequently made me grateful to my former self (instead of cussing his lazy ass, which is my more usual attitude toward Past Matt).

Currey and Alexander (1985: fig. 1)

Sauropod limb bones are pretty darned dense, with extremely thick cortices and smallish marrow spaces that are not actually hollow (tubular) but are instead filled with trabecular bone. My gut feeling is that even a four-foot sauropod femur would be almost too heavy to lift, let alone wield as a club, so in the coming calculations I will err in the direction of underestimating the mass, to give our hypothetical caveman the best possible chance of realizing his dream.

Some of the proportionally thinnest cortices I’ve seen in sauropod limb bones are those of the macronarian Haestasaurus becklesii NHMUK R1870, which Mike conveniently showed in cross-section in this post. I could look up the actual dimensions of the bones (in Upchurch et al 2015: table 1 – they passed the MYDD test, as expected), but for these calculations I don’t need them. All I need are relative areas, for which pixels are good enough.

First, I took Mike’s photo into GIMP and drew two diameters across each bone, one maximum diameter and a second at right angles. Then I drew tick marks about where I think the boundaries lie between the cortex and the trabecular marrow cavity. Next, I used those lines as guides to determine the outer diameters (D) and inner diameters (d) in pixels, as noted in the image.

For the radius, on the left, the mean diameters are D = 891 and d = 648. I could divide those by 2 to get radii and then plug them into the formula for the area of a circle, etc., but there’s an easier way still. For a tubular bone, the proportional area of the inner circle or ellipse is equal to k^2, where k = r/R. Or d/D. (See Wedel 2005 and Tutorial 24 for the derivation of that.) For the Haestasaurus radius (the bone, not the geometric dimension), d/D = 0.727, and that number squared is 0.529. So the marrow cavity occupies 53% of the cross-sectional area, and the cortex occupies the other 47%.

For the ulna, on the right, the mean diameters are D = 896 and d = 606, d/D = 0.676, and that number squared is 0.457. So in this element, the marrow cavity occupies 46% of the cross-sectional area, and the cortex occupies the other 54%.

(For this quick-and-dirty calculation, I am going to ignore the fact that limb bones are more complex than tubes and that their cross-sectional properties change along their lengths – what I am doing here is closer to Fermi estimation than to anything I would publish. And we’ll ground-truth it before the end anyway.)

Left: rat humerus, right: mole humerus. The mole humerus spits upon my simple geometric models, with extreme prejudice. From this post.

You can see from the photo (the Haestasaurus photo, not the mole photo) that neither bone has a completely hollow marrow cavity – both marrow cavities are filled with trabecular bone. By cutting out good-looking chunks in GIMP and thresholding them, I estimate that these trabecular areas are about 30% bone and 70% marrow (actual marrow space with no bone tissue) by cross-sectional area. According to Currey and Alexader (1985: 455), the specific gravities of fatty marrow and bone tissue are 0.93 and 2.1, respectively. The density of the trabecular area is then (0.3*2.1)+(0.7*0.93) =  1.28 kg/L, or about one quarter more dense than water.

But that’s just the trabecular area, which accounts for about one half of the cross-sectional area of each bone. The other half is cortex, which is probably close to 2.1 kg/L throughout. The estimated whole-element densities are then:

Radius: (0.53*1.28)+(0.47*2.1) = 1.67 kg/L

Ulna: (0.46*1.28)+(0.54*2.1) = 1.72 kg/L

Do those numbers pass the sniff test? Well, any skeletal elements that are composed of bone tissue (SG = 2.1) and marrow (SG = 0.93) are constrained to have densities somewhere between those extremes (some animals beat this by building parts of their skeletons out of [bone tissue + air] instead of [bone tissue + marrow]). We know that sauropod limb bones tend to have thick cortices and small marrow cavities, and that the marrow cavities are themselves a combination of trabecular bone and actual marrow space, so we’d expect the overall density to be closer to the 2.1 kg/L end of the scale than the 0.93 kg/L end. And our rough estimates of ~1.7 kg/L fall about where we’d expect.

Femur of Haplocanthosaurus priscus, CM 572, modified from Hatcher (1903: fig. 14).

To convert to masses, we need to know volumes. We can use Haplocanthosaurus here – the femur of the holotype of H. priscus, CM 572, is 1275 mm long (Hatcher 1903), which is just a hair over four feet (4 ft, 2.2 in to be exact). The midshaft width is 207 mm, and the proximal and distal max widths are 353 and 309 mm, respectively. I could do a for-real GDI, but I’m lazy and approximate numbers are good enough here. Just eyeballing it, the width of the femur is about the same over most of its length, so I’m guessing the average width is about 23 cm. The average width:length ratio for the femora of non-titanosaur sauropods is 3:2 (Wilson and Carrano 1999: table 1), which would give an anteroposterior diameter of about 15 cm and an average diameter over the whole length of 19 cm. The volume would then be the average cross-section area, 3.14*9.5*9.5, multiplied by the length, 128 cm, or 36,273 cm^3, or 36.3 L. Multiplied by the ~1.7 kg/L density we estimated above, that gives an estimated mass of 62 kg, or about 137 lbs. A femur that was exactly four feet long would be a little lighter – 86.6% as massive, to be exact, or 53.4 kg (118 lbs).

I know that the PCs in RPGs are supposed to be heroes, but that seems a little extreme.

But wait! Bones dry out and they lose mass as they do so. Lawes and Gilbert (1859) reported that the dry weight of bones of healthy sheep and cattle was only 74% of the wet mass. Cows and sheep have thinner bone cortices than sauropods or elephants, but it doesn’t seem unreasonable that a dry sauropod femur might only weigh 80% as much as a fresh one. That gets us down to 43 kg – about 95 lbs – which is still well beyond what anyone is probably going to be wielding, even if they’re Conan the Cimmerian.

Picture is unrelated.

I mentioned at the top of this section that there are a couple of ways to get here. The second way is to simply see what actual elephant femora weigh, and then scale up to dinosaur size. According to Tefera (2012: table 1), a 110-cm elephant femur has a mass of 21.5 kg (47 lbs). I reckon that’s a dry mass, since the femur in question had sat in a shed for 50 years before being weighed (Tefera 2012: p. 17). Assuming isometry, a four-foot (122 cm) elephant femur would have a dry mass of 29.4 kg (65 lbs). That’s a lot lighter than the estimated mass of the sauropod femur – can we explain the discrepancy?


Femora of a horse, a cow, and an elephant (from left to right in each set), from Tefera (2012: plate 1).

I think so. Elephant femora are more slender than Haplocanthosaurus femora. Tefera (2012) reported a circumference of 44 cm for a 110-cm elephant femur. Scaling up from 110 cm to 122 cm would increase that femur circumference to 49 cm, implying a mean diameter of 15.6 cm, compared to 19 cm for the Haplo femur. That might not seem like a big difference, but it means a cross-sectional area only 2/3 as great, and hence a volume about 2/3 that of a sauropod femur of the same length. And that lines up almost eerily well with our estimated masses of 29 and 43 kg (ratio 2:3) for the four-foot elephant and sauropod femora.

A Better Weapon?

Could our hypothetical caveman do better by choosing a different dinosaur’s femur? Doubtful – the femora of ‘Sue’ are roughly the same length as the Haplo femur mentioned above, and have similar cross-sectional dimensions. Hadrosaur and stegosaur femora don’t look any better. Even if the theropod femur was somewhat lighter because of thinner cortices, how are you going to effectively grip and wield something 15-19 cm in diameter?

I note that the largest axes and sledgehammers sold by Forestry Suppliers, Inc., are about 3 feet long. Could we get our large-animal-femur-based-clubs into the realm of believability by shrinking them to 3 feet instead of 4? Possibly – 0.75 to the third power is 0.42. That brings the elephant femur club down to 12.3 kg (27 lbs) and the sauropod femur club down to 18 kg (40 lbs), only 2-3 times the mass of the largest commonly-available sledgehammers. I sure as heck wouldn’t want to lug such a thing around, much less swing it, but I can just about imagine a mighty hero doing so.

Yes, there were longer historical weapons. Among swing-able weapons (as opposed to spears, etc.), Scottish claymores could be more than four feet long, but crucially they were quite light compared to the clubs we’ve been discussing, maxing out under 3 kg, at least according to Wikipedia.

T. rex FMNH PR2081 right fibula in lateral (top) and medial (bottom) views. Scale is 30 cm. From Brochu (2003: fig. 97).

If one is looking for a good dinosaur bone to wield as a club, may I suggest the fibula of a large theropod? The right (non-pathologic) fibula of ‘Sue’ is 103 cm long (3 ft, 4.5 in), has a max shaft diameter just under 3 inches – so it could plausibly be held by (large) human hands, and it probably massed something like 8-9 kg (17-20 lbs) in life, based on some quick-and-dirty calculations like those I did above. The proximal end is even expanded like the head of a war club. The length and mass are both in the realm of possibility for large, fit, non-supernaturally-boosted humans. Half-orc barbarians will love them.

And that’s my ‘expert’ recommendation as a dice-slinging paleontologist. Thanks for reading – you have Conan-level stamina if you got this far – and thanks to Shawn for letting me use his question to freewheel on some of my favorite geeky topics.



“But wait, Matt”, I hear you thinking. “Every news agency in the world is tripping over themselves declaring Patagotitan the biggest dinosaur of all time. Why are you going in the other direction?”

Because I’ve been through this a few times now. But mostly because I can friggin’ read.

Maximum dorsal centrum diameter in Argentinosaurus is 60cm (specimen MCF-PVPH-1, Bonaparte and Coria 1993). In Puertasaurus it is also 60cm (MPM 10002, Novas et al. 2005). In Patagotitan it is 59cm (MPEF-PV 3400/5, Carballido et al. 2017). (For more big centra, see this post.)

Femoral midshaft circumference is 118cm in an incomplete femur of Argentinosaurus estimated to be 2.5m long when complete (Mazzetta et al. 2004). A smaller Argentinosaurus femur is 2.25m long with a circumference of 111.4cm (Benson et al. 2014). The largest reported femur of Patagotitan, MPEF-PV 3399/44, is 2.38m long and has a circumference of either 101cm (as reported in the Electronic Supplementary Materials to Carballido et al 2017) or 110cm (as reported in the media in 2014*).

TL;DR: 60>59, and 118>111>110>101, and in both cases Argentinosaurus > Patagotitan, at least a little bit.

Now, Carballido et al (2017) estimated that Patagotitan was sliiiiightly more massive than Argentinosaurus and Puertasaurus by doing a sort of 2D minimum convex hull dorsal vertebra area thingy, which the Patagotitan vertebra “wins” because it has a taller neural spine than either Argentinosaurus or Puertasaurus, and slightly wider transverse processes than Argentinosaurus (138cm vs 128cm) – but way narrower transverse processes than Puertasaurus (138cm vs 168cm). But vertebrae with taller or wider sticky-out bits do not a more massive dinosaur make, otherwise Rebbachisaurus would outweigh Giraffatitan.

Now, in truth, it’s basically a three-way tie between Argentinosaurus, Puertasaurus, and Patagotitan. Given how little we have of the first two, and how large the error bars are on any legit size comparison, there is no real way to tell which of them was the longest or the most massive. Still, to get to the conclusion that Patagotitan was in any sense larger than Argentinosaurus you have to physically drag yourself over the following jaggedly awkward facts:

  1. The weight-bearing parts of the anterior dorsal vertebrae are larger in diameter in both Argentinosaurus and Puertasaurus than in Patagotitan. Very slightly, but still, Patagotitan is the smallest of the three.
  2. The femora of Argentinosaurus are fatter than those of Patagotitan, even at shorter length. The biggest femora of Argentinosaurus are longer, too.

So all of the measurements of body parts that have to do with supporting mass are still larger in Argentinosaurus than in Patagotitan.

Now, it is very cool that we now have a decent chunk of the skeleton of a super-giant titanosaur, instead of little bits and bobs. And it’s nice to know that the numbers reported in the media back in 2014 turned out to be accurate. But Patagotitan is not the “world’s largest dinosaur”. At best, it’s the third-largest contender among near equals.

Parting shot to all the science reporters who didn’t report the same numbers I did here: instead of getting hype-notized by assumption-laden estimates, how about doing an hour’s worth of research making the most obvious possible comparisons?

Almost immediate UPDATE: Okay, that parting shot wasn’t entirely fair. As far as I know, the measurements of Patagotitan were not available until the embargo lifted. Which is in itself odd – if someone claims to have the world’s largest dinosaur, but doesn’t put any measurements in the paper, doesn’t that make your antennae twitch? Either demand some measurements so you can make those obvious comparisons, or approach with extreme skepticism – especially if the “world’s largest dino” claim was pre-debunked three years ago!

* From this article in the Boston Globe:

Paleobiologist Paul Upchurch of University College London believes size estimates are more reliable when extrapolated from the circumference of bones.

He said this femur is a whopping 43.3 inches around, about the same as the Argentinosaurus’ thigh bone.

‘‘Whether or not the new animal really will be the largest sauropod we know remains to be seen,’’ said Upchurch, who was not involved in this discovery but has seen the bones first-hand.

Some prophetically appropriate caution from Paul Upchurch there, who has also lived through a few of these “biggest dinosaur ever” bubbles.


What would the world look like if, as proposed by the Max Planck Institute, the scholarly world flipped from being dominated by subscriptions to Gold open access? I think there are three things to say.

First, incentives. A concern is sometimes expressed that when publishers are paid per paper published, they will have an incentive to want more papers to be published. Would this exacerbate the existing publish-or-perish culture where we are flooded by quantity of publications, sometimes at the expense of quality?

It’s certainly true that in a Gold OA world, the publishers would like to see more papers (and monographs) published. But whether we the academic community respond to that desire by publishing more is not a decision that the publishers get to make. This — like so many issues — comes back to the problem of what incentives apply in academia. While scholars gains rewards like promotion and tenure by publishing many papers (for example because committees evaluate people based on their H-index), it is inevitable that those scholars will seek to publish many papers — and this would be true whether in a subscription-based or Gold OA-based system. Thus I think the problem of publishing quantity rather than quality is quite independent from the problem of how we pay for publications.

Second, costs. I sometimes hear a concern is that a flip to Gold OA would create an environment where funds are tied up, and resources are not sufficient of fund new and innovative journals.

I’m sure these numbers are not new to regular readers, but it seems pretty clear that a flipped world would have much lower total costs than the present system. Here are the numbers:

The STM Report for 2015, page 6, reports total publisher income in the STM field as $10 billion for 2013, and says that about 2.5 million papers were published that year. That gives an average income per paper of $4000. (We can probably assume a broadly similar figure for non-STM papers, too.) By contrast, the Wellcome Trust’s recent report on its APC spending in 2013-14 shows an average APC of £1837, currently about $2634. This is slightly less than 2/3 what the world at large is paying per paper.

In other words, even using the relatively high APCs paid by the Wellcome Trust, the world’s 2.5 million papers per year could be published for $6.6 billion — saving $3.4 billion to spent elsewhere.

Third, markets. This one is a question, and I think it’s crucial for the prospects of a Gold-OA ecosystem: will we get an efficient market in APCs? If we do, then prices will be forced down until they are very close to costs — which publishers like Hindawi, Ubiquity Press and PeerJ have shown can be in the $400-500 range, almost literally an order of magnitude less than the world presently pays for publication. But if no true market emerges, prices will not fall — indeed publishers may have the leverage to raise APCs at rates greater than inflation, as they have been doing for subscriptions.

That is why I believe that, however tempting “APC Big Deals” are to individual libraries or consortia, they should be strenuously resisted. As with subscription Big Deals, the short-term savings (while real) would be absolutely dwarfed by the long-term losses.

If I’m right about this, then we face a tragedy of the commons during this phase of transition from subscriptions to Gold OA: it will be in the short-term interests of each library to accept a Big Deal on APCs; but again the interests of the community. We will need to communicate well, and function as a global community, to avoid falling into this trap.

[I first wrote this post as an email to a list for delegates of the OSI2016 conference. Then I realised that it’s of broader interest, and edited it into the form seen here.]

I was a bit disappointed to hear David Attenborough on BBC Radio 4 this morning, while trailing a forthcoming documentary, telling the interviewing that you can determine the mass of an extinct animal by measuring the circumference of its femur.

We all know what he was alluding to, of course: the idea first published by Anderson et al. (1985) that if you measure the life masses of lots of animals, then measuring their long-bone circumferences when they’ve died, you can plot the two measurements against each other, find a best-fit line, and extrapolate it to estimate the masses of dinosaurs based on their limb-bone measurements.


This approach has been extensively refined since 1985, most recently by Benson et al. (2014). but the principle is the same.

But the thing is, as Anderson et al. and other authors have made clear, the error-bars on this method are substantial. It’s not super-clear in the image above (Fig 1. from the Anderson et al. paper) because log-10 scales are used, but the 95% confidence interval is about 42 pixels tall, compared with 220 pixels for an order of magnitude (i.e. an increment of 1.0 on the log-10 scale). That means the interval is 42/220 = 0.2 of an order of magnitude. That’s a factor 10 ^ 0.2 = 1.58. In other words you could have two animals with equally robust femora, one of them nearly 60% heavier than the other, and they would both fall within the 95% confidence interval.

I’m surprised that someone as experienced and knowledgeable as Attenborough would perpetuate the idea that you can measure mass with any precision in this way (even more so when using only a femur, rather than the femur+humerus combo of Anderson et al.)

More: when the presenter told him that not all scientists buy the idea that the new titanosaur is the biggest known, he said that came as a surprise. Again, it’s disappointing that the documentary researchers didn’t make Attenborough aware of, for example, Paul Barrett’s cautionary comments or Matt Wedel’s carefully argued dissent. Ten minutes of simple research would have found this post — for example, it’s Google’s fourth hit for “how big is the new argentinian titanosaur”. I can only hope that the actual documentary, which screens on Sunday 24 January, doesn’t present the new titanosaur’s mass as a known and agreed number.

(To be clear, I am not blaming Attenborough for any of this. He is a presenter, not a palaeontologist, and should have been properly prepped by the researchers for the programme he’s fronting. He is also what can only be described as 89, so should be forgiven if he’s not quite as quick on his feel when confronted with an interviewer as he used to be.)

Update 1 (the next day)

Thanks to Victoria Arbour for pointing out an important reference that I missed: it was Campione and Evans (2012) who expanding Anderson et al.’s dataset and came up with the revised equation which Benson et al. used.

Update 2 (same day as #1)

It seems most commenters are inclined to go with Attenborough on this. That’s a surprise to me — I wonder whether he’s getting a free pass because of who he is. All I can say is that as I listened to the segment it struck me as really misleading. You can listen to it for yourself here if you’re in the UK; otherwise you’ll have to make do with this transcript:

“It’s surprising how much information you can get from just one bone. I mean for example that thigh bone, eight feet or so long, if you measure the circumference of that, you will be able to say how much weight that could have carried, because you know what the strength of bone is. So the estimate of weight is really pretty accurate and the thought is that this is something around over seventy tonnes in weight.”

(Note also that the Anderson et al./Campione and Evans method has absolutely nothing to do with the strength of bone.)

Also if interest was this segment that followed immediately:

How long it was depends on whether you think it held its neck out horizontaly or vertically. If it held it out horizontally, well then it would be about half as big again as the Diplodocus, which is the dinosaur that’s in the hall of the Natural History Museum. It would be absolutely huge.

Interviewer: And how tall, if we do all the dimensions?

Ah well that is again the question of how it holds its neck, and it could have certainly reached up about to the size of a four or five storey building.

Needless to say, the matter of neck posture is very relevant to our interests. I don’t want to read too much into a couple of throwaway comments, but the implication does seem to be that this is an issue that the documentary might spend some time on. We’ll see what happens.


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:

Arambourgiania vs giraffe vs the Disacknowledgement redux Witton ver 2 low res

(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.

Arambourgiania vs giraffe vs the Disacknowledgement redux Witton ver 2 low res

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?


  • 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.

Last night, I submitted a paper for publication — for the first time since April 2013. I’d almost forgotten what it felt like. But, because we’re living in the Shiny Digital Future, you don’t have to wait till it’s been through review and formal publication to read it. I submitted to PeerJ, and at the same time, made it available as a preprint (Taylor 2014).

It’s called “Quantifying the effect of intervertebral cartilage on neutral posture in the necks of sauropod dinosaurs”, and frankly the results are weird. Here’s a taste:

Taylor (2014:figure 3). Effect of adding cartilage to the neutral pose of the neck of Apatosaurus louisae CM 3018. Images of vertebra from Gilmore (1936:plate XXIV). At the bottom, the vertebrae are composed in a horizontal posture. Superimposed, the same vertebrae are shown inclined by the additional extension angles indicated in Table 1. If the slightly sub-horizontal osteological neutral pose of Stevens and Parrish (1999) is correct, then the cartilaginous neutral pose would be correspondingly slightly lower than depicted here, but still much closer to the elevated posture than to horizontal. (Note that the posture shown here would not have been the habitual posture in life: see discussion.)

Taylor (2014:figure 3). Effect of adding cartilage to the neutral pose of the neck of Apatosaurus louisae CM 3018. Images of vertebra from Gilmore (1936:plate XXIV). At the bottom, the vertebrae are composed in a horizontal posture. Superimposed, the same vertebrae are shown inclined by the additional extension angles indicated in Table 1. If the slightly sub-horizontal osteological neutral pose of Stevens and Parrish (1999) is correct, then the cartilaginous neutral pose would be correspondingly slightly lower than depicted here, but still much closer to the elevated posture than to horizontal. (Note that the posture shown here would not have been the habitual posture in life: see discussion.)

A year back, as I was composing a blog-post about our neck-cartilage paper in PLOS ONE (Taylor and Wedel 2013c), I found myself writing down the rather trivial formula for the additional angle of extension at an intervertebral joint once the cartilage is taken into account. In that post, I finished with the promise “I guess that will have to go in a followup now”. Amazingly it’s taken me a year to get that one-pager written and submitted. (Although in the usual way of things, the manuscript ended up being 13 pages long.)

To summarise the main point of the paper: when you insert cartilage of thickness t between two vertebrae whose zygapophyses articulate at height h above the centra, the more anterior vertebra is forced upwards by t/h radians. Our best guess for how much cartilage is between the adjacent vertebrae in an Apatosaurus neck is about 10% of centrum length: the image above shows the effect of inserting that much cartilage at each joint.

And yes, it’s weird. But it’s where the data leads me, so I think it would be dishonest not to publish it.

I’ll be interested to see what the reviewers make of this. You are all of course welcome to leave comments on the preprint itself; but because this is going through conventional peer-review straight away (unlike our Barosaurus preprint), there’s no need to offer the kind of detailed and comprehensive comment that several people did with the previous one. Of course feel free if you wish, but I’m not depending on it.


Gilmore Charles W. 1936. Osteology of Apatosaurus, with special reference to specimens in the Carnegie Museum. Memoirs of the Carnegie Museum 11:175–300 and plates XXI–XXXIV.

Stevens, Kent A., and J. Michael Parrish. 1999. Neck posture and feeding habits of two Jurassic sauropod dinosaurs. Science 284(5415):798–800. doi:10.1126/science.284.5415.798

Taylor, Michael P. 2014. Quantifying the effect of intervertebral cartilage on neutral posture in the necks of sauropod dinosaurs. PeerJ PrePrints 2:e588v1 doi:10.7287/peerj.preprints.588v1

Taylor, Michael P., and Mathew J. Wedel. 2013c. The effect of intervertebral cartilage on neutral posture and range of motion in the necks of sauropod dinosaurs. PLOS ONE 8(10):e78214. 17 pages. doi:10.1371/journal.pone.0078214

Gender balance at SVPCA

September 17, 2014

I’ve always thought of SVPCA as a pretty well gender-balanced conference: if not 50-50 men and women, then no more than 60-40 slanted towards men. So imagine my surprise when I ran the actual numbers.

1. Delegates. I went through the delegate list at the back of the abstracts book, counting the men and women. Those I knew, or whose name made it obvious, I noted down; the half-dozen that I couldn’t easily categorise, I have successfully stalked on the Internet. So I now know that there were 39 women and 79 men — so that women made up 33% of the delegates, almost exactly one third.

Official conference photo, SVPCA 2014, York, UK.

Official conference photo, SVPCA 2014, York, UK.

2. Presentations. There were a total of 50 presentations in the three days of SVPCA: 18 on days 1 and 3, and 14 on day 2, which had a poster session in place of the final session of four talks. I counted the presenters (which were usually, but not always, the lead authors). Here’s how the number of talks by women broke down:

Day one: 2 of 18
Day two: 8 of 14
Day three: 3 of 18

In total, this gives us 13 of 50 talks by women, or 26%.

3. Presenter:delegate ratios. Since 37 of the 79 attending men gave talks, that’s 47% of them; but only 13 of the 39 attending women gave talks, which is 33%. On other words, a man attending SVPCA was 40% more likely to give a talk than a woman.

I’m not sure what to make of all this. I was shocked when I found that only one ninth of the first day’s talks were by women. It’s a statistical oddity that women actually dominated day two, but day three was nearly as unbalanced as day one.

Since SVPCA accepts pretty much every submitted talk, the conference itself can’t be blamed for the imbalance. (At least, not unless you think the organisers should turn down talks by men just because they’re men, leaving blank spots in the program.) It seems that the imbalance more likely reflects that of the field in general. Maybe more disturbing is that the proportion of women giving talks was rather less than the proportion attending (26% vs. 33%) which suggests that perhaps women feel more confident about attending than about presenting.

It would be interesting to know how these numbers compare with SVP’s.