Clash of the Titans from above

Here’s the “Clash of the Titans” exhibit at the Sam Noble Oklahoma Museum of Natural History, featuring the reconstructed skeletons of the giant Oklahoma Apatosaurus – which I guess should now be called the giant Oklahoma apatosaurine until someone sorts out its phylogenetic position – and the darn-near-T. rex-sized Saurophaganax maximus, which may be Allosaurus maximus depending on who you’re reading.

Now, I love this exhibit in both concept and execution. But one thing that is more obvious in this view from the upper level balcony is that despite its impressive weaponry, a lone 3-to-5 ton Saurophaganax had an Arctic ice cap’s chance in the Anthropocene of taking down a healthy 30-meter, 40-50 ton apatosaur (which is to say, none). I like to imagine that in the photo above, the apatosaur is laughing at the pathetically tiny theropod and its delusions of grandeur.

Clash of the Titans from behind

In this shot from behind, you get a better look at the baby apatosaur standing under the big one, and it hints at a far more likely target for Saurophaganax and other large Morrison theropods: sauropods that were not fully-grown, which was almost all of them. I am hip to the fact that golden eagles kill deer, and some lions will attack elephants – as Cookie Monster says, “Sometime food, not anytime food” – but not only were smaller sauropods easier prey, they were far more numerous given the inevitable population structure of animals that started reproducing at a young age and made more eggs the bigger they got (as essentially all egg-laying animals do).

In fact, as discussed in our recent paper on dinosaur ontogeny (Hone et al. 2016), there may have been times when the number of fully-grown sauropods in a given population was zero, and the species was maintained by reproducing juveniles. The giant Oklahoma apatosaurine is a unique specimen today – by far the largest apatosaurine we have fossils of – but it may also have been an anomaly in its own time, the rare individual that made it through the survivorship gauntlet to something approaching full size.

Amazingly enough, there is evidence that even it was not fully mature, but that’s a discussion for another day. Parting shot:

Oklahoma Apatosaurus neck and head


Notocolossus is a beast

January 20, 2016

Notocolossus skeletal recon - Gonzalez Riga et al 2016 fig 1

(a) Type locality of Notocolossus (indicated by star) in southern-most Mendoza Province, Argentina. (b) Reconstructed skeleton and body silhouette in right lateral view, with preserved elements of the holotype (UNCUYO-LD 301) in light green and those of the referred specimen (UNCUYO-LD 302) in orange. Scale bar, 1 m. (González Riga et al. 2016: figure 1)

This will be all too short, but I can’t let the publication of a new giant sauropod pass unremarked. Yesterday Bernardo González Riga and colleagues published a nice, detailed paper describing Notocolossus gonzalezparejasi, “Dr. Jorge González Parejas’s southern giant”, a new titanosaur from the Late Cretaceous of Mendoza Province, Argentina (González Riga et al. 2016). The paper is open access and freely available to the world.

As you can see from the skeletal recon, there’s not a ton of material known from Notocolossus, but among giant sauropods it’s actually not bad, being better represented than Argentinosaurus, Puertasaurus, Argyrosaurus, and Paralititan. In particular, one hindfoot is complete and articulated, and a good chunk of the paper and supplementary info are devoted to describing how weird it is.

But let’s not kid ourselves – you’re not here for feet, unless it’s to ask how many feet long this monster was. So how big was Notocolossus, really?

Well, it wasn’t the world’s largest sauropod. And to their credit, no-one on the team that described it has made any such superlative claims for the animal. Instead they describe it as, “one of the largest terrestrial vertebrates ever discovered”, and that’s perfectly accurate.

Notocolossus limb bones - Gonzalez Riga et al 2016 fig 4

(a) Right humerus of the holotype (UNCUYO-LD 301) in anterior view. Proximal end of the left pubis of the holotype (UNCUYO-LD 301) in lateral (b) and proximal (c) views. Right tarsus and pes of the referred specimen (UNCUYO-LD 302) in (d) proximal (articulated, metatarsus only, dorsal [=anterior] to top), (e) dorsomedial (articulated), and (f) dorsal (disarticulated) views. Abbreviations: I–V, metatarsal/digit number; 1–2, phalanx number; ast, astragalus; cbf, coracobrachialis fossa; dpc, deltopectoral crest; hh, humeral head; ilped, iliac peduncle; of, obturator foramen; plp, proximolateral process; pmp, proximomedial process; rac, radial condyle; ulc, ulnar condyle. Scale bars, 20 cm (a–c), 10 cm (d–f). (Gonzalez Riga et al 2016: figure 4)

Any discussions of the size of Notocolossus will be driven by one of two elements: the humerus and the anterior dorsal vertebra. The humerus is 176 cm long, which is shorter than those of Giraffatitan (213 cm), Brachiosaurus (204 cm), and Turiasaurus (179 cm), but longer than those of Paralititan (169 cm), Dreadnoughtus (160 cm), and Futalognkosaurus (156 cm). Of course we don’t have a humerus for Argentinosaurus or Puertasaurus, but based on the 250-cm femur of Argentinosaurus, the humerus was probably somewhere around 200 cm. Hold that thought.

Notocolossus and Puertasaurus dorsals compared

Top row: my attempt at a symmetrical Notocolossus dorsal, made by mirroring the left half of the fossil from the next row down. Second row: photos of the Notocolossus dorsal with missing bits outlined, from Gonzalez Riga et al (2016: fig. 2). Scale bar is 20 cm (in original). Third row: the only known dorsal vertebra of Puertasaurus, scaled to about the same size as the Notocolossus vertebra, from Novas et al. (2005: fig. 2).

The anterior dorsal tells a similar story, and this is where I have to give González Riga et al. some props for publishing such detailed sets of measurements in the their supplementary information. They Measured Their Damned Dinosaur. The dorsal has a preserved height of 75 cm – it’s missing the tip of the neural spine and would have been a few cm taller in life – and by measuring the one complete transverse process and doubling it, the authors estimate that when complete it would have been 150 cm wide. That is 59 inches, almost 5 feet. The only wider vertebra I know of is the anterior dorsal of Puertasaurus, at a staggering 168 cm wide (Novas et al. 2005). The Puertasaurus dorsal is also quite a bit taller dorsoventrally, at 106 cm, and it has a considerably larger centrum: 43 x 60 cm, compared to 34 x 43.5 cm for Notocolossus (anterior centrum diameters, height x width).

Centrum size is an interesting parameter. Because centra are so rarely circular, arguably the best way to compare across taxa would be to measure the max area (or, since centrum ends are also rarely flat, the max cross-sectional area). It’s late and this post is already too long, so I’m not going to do that now. But I have been keeping an informal list of the largest centrum diameters among sauropods – and, therefore, among all Terran life – and here they are (please let me know if I missed anyone):

  • 60 cm – Argentinosaurus dorsal, MCF-PVPH-1, Bonaparte and Coria (1993)
  • 60 cm – Puertasaurus dorsal, MPM 10002, Novas et al. (2005)
  • 51 cm – Ruyangosaurus cervical and dorsal, 41HIII-0002, Lu et al. (2009)
  • 50 cm – Alamosaurus cervical, SMP VP−1850, Fowler and Sullivan (2011)
  • 49 cm – Apatosaurus ?caudal, OMNH 1331 (pers. obs.)
  • 49 cm – Supersaurus dorsal, BYU uncatalogued (pers. obs.)
  • 46 cm – Dreadnoughtus dorsal, MPM-PV 1156, Lacovara et al. (2014: Supplmentary Table 1) – thanks to Shahen for catching this one in the comments!
  • 45.6 cm – Giraffatitan presacral, Fund no 8, Janensch (1950: p. 39)
  • 45 cm – Futalognkosaurus sacral, MUCPv-323, Calvo et al. (2007)
  • 43.5 cm – Notocolossus dorsal, UNCUYO-LD 301, González Riga et al. (2016)

(Fine print: I’m only logging each taxon once, by its largest vertebra, and I’m not counting the dorsoventrally squashed Giraffatitan cervicals which get up to 47 cm wide, and the “uncatalogued” Supersaurus dorsal is one I saw back in 2005 – it almost certainly has been catalogued in the interim.) Two things impress me about this list: first, it’s not all ‘exotic’ weirdos – look at the giant Oklahoma Apatosaurus hanging out halfway down the list. Second, Argentinosaurus and Puertasaurus pretty much destroy everyone else by a wide margin. Notocolossus doesn’t seem so impressive in this list, but it’s worth remembering that the “max” centrum diameter here is from one vertebra, which was likely not the largest in the series – then again, the same is true for Puertasaurus, Alamosaurus, and many others.

Notocolossus phylogeny - Gonzalez Riga et al 2016 fig 5

(a) Time-calibrated hypothesis of phylogenetic relationships of Notocolossus with relevant clades labelled. Depicted topology is that of the single most parsimonious tree of 720 steps in length (Consistency Index = 0.52; Retention Index = 0.65). Stratigraphic ranges (indicated by coloured bars) for most taxa follow Lacovara et al.4: fig. 3 and references therein. Additional age sources are as follows: Apatosaurus[55], Cedarosaurus[58], Diamantinasaurus[59], Diplodocus[35], Europasaurus[35], Ligabuesaurus[35], Neuquensaurus[60], Omeisaurus[55], Saltasaurus[60], Shunosaurus[55], Trigonosaurus[35], Venenosaurus[58], Wintonotitan[59]. Stratigraphic ranges are colour-coded to also indicate geographic provenance of each taxon: Africa (excluding Madagascar), light blue; Asia (excluding India), red; Australia, purple; Europe, light green; India, dark green; Madagascar, dark blue; North America, yellow; South America, orange. (b–h) Drawings of articulated or closely associated sauropod right pedes in dorsal (=anterior) view, with respective pedal phalangeal formulae and total number of phalanges per pes provided (the latter in parentheses). (b) Shunosaurus (ZDM T5402, reversed and redrawn from Zhang[45]); (c) Apatosaurus (CM 89); (d) Camarasaurus (USNM 13786); (e) Cedarosaurus (FMNH PR 977, reversed from D’Emic[32]); (f) Epachthosaurus (UNPSJB-PV 920, redrawn and modified from Martínez et al.[22]); (g) Notocolossus; (h) Opisthocoelicaudia (ZPAL MgD-I-48). Note near-progressive decrease in total number of pedal phalanges and trend toward phalangeal reduction on pedal digits II–V throughout sauropod evolutionary history (culminating in phalangeal formula of 2-2-2-1-0 [seven total phalanges per pes] in the latest Cretaceous derived titanosaur Opisthocoelicaudia). Abbreviation: Mya, million years ago. Institutional abbreviations see Supplementary Information. (González Riga et al. 2016: figure 5)

As for the estimated mass of Notocolossus, González Riga et al. (2016) did their due diligence. The sections on mass estimation in the main text and supplementary information are very well done – lucid, modest, and fair. Rather than try to summarize the good bit, I’ll just quote it. Here you go, from page 7 of the main text:

The [humeral] diaphysis is elliptical in cross-section, with its long axis oriented mediolaterally, and measures 770 mm in minimum circumference. Based on that figure, the consistent relationship between humeral and femoral shaft circumference in associated titanosaurian skeletons that preserve both of these dimensions permits an estimate of the circumference of the missing femur of UNCUYO-LD 301 at 936 mm (see Supplementary Information). (Note, however, that the dataset that is the source of this estimate does not include many gigantic titanosaurs, such as Argentinosaurus[5], Paralititan[16], and Puertasaurus[11], since no specimens that preserve an associated humerus and femur are known for these taxa.) In turn, using a scaling equation proposed by Campione and Evans[20], the combined circumferences of the Notocolossus stylopodial elements generate a mean estimated body mass of ~60.4 metric tons, which exceeds the ~59.3 and ~38.1 metric ton masses estimated for the giant titanosaurs Dreadnoughtus and Futalognkosaurus, respectively, using the same equation (see Supplementary Information). It is important to note, however, that subtracting the mean percent prediction error of this equation (25.6% of calculated mass[20]) yields a substantially lower estimate of ~44.9 metric tons for UNCUYO-LD 301. Furthermore, Bates et al.[21] recently used a volumetric method to propose a revised maximum mass of ~38.2 metric tons for Dreadnoughtus, which suggests that the Campione and Evans[20] equation may substantially overestimate the masses of large sauropods, particularly giant titanosaurs. Unfortunately, however, the incompleteness of the Notocolossus specimens prohibits the construction of a well-supported volumetric model of this taxon, and therefore precludes the application of the Bates et al.[21] method. The discrepancies in mass estimation produced by the Campione and Evans[20] and Bates et al.[21] methods indicate a need to compare the predictions of these methods across a broad range of terrestrial tetrapod taxa[21]. Nevertheless, even if the body mass of the Notocolossus holotype was closer to 40 than 60 metric tons, this, coupled with the linear dimensions of its skeletal elements, would still suggest that it represents one of the largest land animals yet discovered.

So, nice work all around. As always, I hope we get more of this critter someday, but until then, González Riga et al. (2016) have done a bang-up job describing the specimens they have. Both the paper and the supplementary information will reward a thorough read-through, and they’re free, so go have fun.


I just gave an answer to this question on Quora, and it occurred to me that I ought to also give it a permanent home here. So here it is.

This is a great example of a question that you’d think would have a simple, clear answer, but doesn’t. In fact, as a palaeontologist specialising in dinosaur gigantism, I have an abiding fear of being asked this question in a pub quiz, and not being able to produce the name that’s written on the quizmaster’s answer sheet.

First, what do we mean by “biggest”? Diplodocus was longer than Apatosaurus, but Apatosaurus was heavier. Giraffatitan was taller than either. Let’s simplify and assume we want to know the heaviest dinosaur.

Second, estimating the masses of extinct animals is incredibly hard even when we have a pretty complete skeleton. For example, the gigantic mounted brachiosaur skeleton in Berlin (which used to be called “Brachiosaurus” brancai but is now recognised as the separate genus Giraffatitan) has been subject to at least 14 estimates in the published scientific literature, as summarised here. They vary from 13,618 kg to 78,258 kg — a factor of 5.75 for the same individual. That’s like looking at a human skeleton and not knowing whether its from Kate Moss or Arnold Schwazenegger. (There are reasons for this and I urge you to read the linked article.)

Third, the big dinosaurs tend to be very poorly represented. Giraffatitan is probably the heaviest dinosaur known from a more or less complete skeleton (though even that is put together from several different individuals) so I could give that as the answer to the hypothetical pub-quiz — though the answer sheet would probably be out of date and call it Brachiosaurus.

Fourth, which individual of a given species do we mean? I said Giraffatitan is known from a more or less complete skeleton. And my best guess is that that individual massed, say, 30,000 kg. But an isolated fibula of the same species is known that’s 12.6% longer than the one in the skeletal mount. That suggest an animal that masses 1.126^3 = 1.43 times as massive as the mounted skeleton — say 43,000 kg. There might be yet bigger Giraffatitan individuals. On the other hand, there is some evidence that Apatosaurus, which is usually thought of as not being so big, might have got even bigger.

Fifth, the very biggest specimens tend to be known from only a handful of bones. A good example here is the titanosaur Argentinosaurus, which is known from several vertebrae and a few limb bones, but not all from the same individual. It’s a good bet that it massed 60-70 tonnes — so maybe about twice as much as Giraffatitan, but much less than the often-cited 100 tonnes. Other, more recently discovered, titanosaurs seem to be in the same size class: Puertasaurus, Futalognkosaurus, Dreadnoughtus and more. They they are hard to compare directly due to the paucity of overlapping material, or at least described overlapping material. (Scientists are working on getting more of this stuff properly described in the literature, which will help.)

But, sixth, the very biggest dinosaurs tend to be apocryphal. There’s Amphicoelias fragillimus, known only from E. D. Cope’s drawing of the upper half of a single vertebra. This may have been 50 m long and massed 80 tonnes; but other published estimates say 58 m and 122 tonnes. We really can’t say from the very poor remains.

So if you get asked this question in a pub quiz, your best bet is to roll a dice, pick an answer, close your eyes and hope. Roll 1 for Giraffatitan, 2 for Brachiosaurus, 3 for Apatosaurus, 4 for Argentinosaurus, 5 for Dreadnoughtus and 6 for Amphicoelias fragillimus. Good luck!


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.

I was contacted recently by David Goldenberg (, a journalist who’s putting together a piece on the biggest dinosaurs. He asked me a few questions, and since I’d taken the time to write answers I thought I may as well post them here.

1) Do you think that we will ever know what the largest dinosaur (by mass) was?

In principle, we can never know that we’ve found the largest dinosaur. All we can know (and we probably can’t really know even this, as we’ll see below) is that we’ve found the largest so far. If we were dealing with animals where there’s a good sample size, there would be statistical techniques that we could use to figure out the likely size-range. But most giant dinosaur species are known only a handful of specimens — sometimes only a single one. How big did Puertasaurus get? We can’t possibly say: the best we can do is estimate how big the one known specimen of Puertasaurus was.

That said, we can sort of get a feel for size classes. There are quite a few sauropods that seem to come in at around 30-40 tonnes — Brachiosaurus, Giraffatitan, Supersaurus, Dreadnoughtus — which suggests there might be some kind of a limit there. But there are bigger titanosaurs (Argentinosaurus, Puertasaurus, Futalognkosaurus) which show that if the barrier exists at all, it’s a “soft” one. And of course the tantalising hints of super-giant sauropods.

There are at least three of these: Amphicoelias fragillimus, a diplodocid known from a drawing of a vertebral arch which has since been lost or destroyed, which could well have massed 100 tonnes. Bruhathkayosaurus, a giant titanosaur known from a two-meter tibia, since destroyed, which could conceivably have massed twice that; and the Broome Sandstone track-maker, known only from footprints, which might have been somewhere in between.

Any one of those, we might write off and say it’s too good to be true — all three stories are pretty vague as to evidence and require a lot of guesswork in the inferences. But the fact that we have all three of these makes me feel pretty certain that there were indeed sauropods out there in the 100-200 tonne range (i.e. the size of big whales). I only hope we find solid, verifiable, curated evidence for them some time soon.

2) What bones do you need to have before you can make an accurate measurement?

You can’t ever make an accurate measurement. Consider even a really well represented, essentially complete specimen such as MB.R.2181 (previously known as HM S II), the giant mounted skeleton in the Museum für Naturkunde Berlin. Peer-reviewed published estimates of the mass of that one individual have varied between 13,618 and 78,258 kg — a factor of 5.75. Even if you discard these obvious outlier estimates, recent and credible estimates vary from 23,337 to 38,000 kg, which is still a factor of 1.63.

And this is not completely crazy. Two humans with essentially identical skeletons can weigh 70 and 114 kg, after all. Soft tissue is essentially impossible to predict.

3) What do you make of the fact that so many different species have been given the title? Is that the fault of the media or scientists or what?

A big part of is that it depends on what you count. That Berlin brachiosaur is the biggest dinosaur known from an essentially complete skeleton, so Giraffatitan is a legitimate holder of the crown. (Confusing matters further, it used to be thought to be a species of Brachiosaurus). But there were definitely bigger sauropods than that — just not known from such complete specimens. Argentinosaurus was certainly bigger, for example. But there’s no way to put a meaningful whole-body mass estimate on it.

But yes, there is also an understandable tendency towards sensationalism, both from scientists and the press. There have been plenty of new discoveries that can legitimately be described as “could be the biggest yet”.

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

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

Huanghetitan ruyangensis sacrum comparison

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

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

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

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



How bigsmall was Aquilops?

December 12, 2014

Handling Aquilops by Brian Engh

Life restoration of Aquilops by Brian Engh (CC-BY).

If you’ve been reading around about Aquilops, you’ve probably seen it compared in size to a raven, a rabbit, or a cat. Where’d those comparisons come from? You’re about to find out.

Back in April I ran some numbers to get a rough idea of the size of Aquilops, both for my own interest and so we’d have some comparisons handy when the paper came out.

Archaeoceratops skeletal reconstruction by Scott Hartman. Copyright Scott Hartman, 2011, used here by permission.

Archaeoceratops skeletal reconstruction by Scott Hartman. Copyright Scott Hartman, 2011, used here by permission.

I started with the much more completely known Archaeoceratops. The measurements of Scott Hartman’s skeletal recon (shown above and on Scott’s website – thanks, Scott!) match the measurements of the Archaeo holotype given by Dodson and You (2003) almost perfectly. The total length of Archaeoceratops, including tail, is almost exactly one meter. Using graphic double integration, I got a volume of 8.88L total for a 1m Archaeoceratops. That would come down to 8.0L if the lungs occupied 10% of body volume, which is pretty standard for non-birds. So that’s about 17-18 lbs.

Archaeoceratops and Aquilops skulls to scale

Aquilops model by Garrett Stowe, photograph by Tom Luczycki, copyright and courtesy of the Sam Noble Oklahoma Museum of Natural History.

Archaeoceratops has a rostrum-jugal length of 145mm, compared to 84mm in Aquilops. Making the conservative assumption that Aquilops = Archaeoceratops*0.58, I got a body length of 60cm (about two feet), and volumes of 1.73 and 1.56 liters with and without lungs, or about 3.5 lbs in life. The internet informed me that the common raven, Corvus corax, has an adult length of 56-78 cm and a body mass of 0.7-2 kg. So, based on this admittedly tall and teetering tower of assumptions, handwaving, and wild guesses, Aquilops (the holotype individual, anyway) was about the size of a raven, in both length and mass. But ravens, although certainly well-known, are maybe a bit remote from the experience of a lot of people, so we wanted a comparison animal that more people would be familiar with. The estimated length and mass of the holotype individual of Aquilops also nicely overlap the species averages (60 cm, 1.4-2.7 kg) for the black-tailed jackrabbit, Lepus californicus, and they’re pretty close to lots of other rabbits as well, hence the comparison to bunnies.

Of course, ontogeny complicates things. Aquilops has some juvenile characters, like the big round orbit, but it doesn’t look like a hatchling. Our best guess is that it is neither a baby nor fully grown, but probably an older juvenile or young subadult. A full-grown Aquilops might have been somewhat larger, but almost certainly no larger than Archaeoceratops, and probably a meter or less in total length. So, about the size of a big housecat. That’s still pretty darned small for a non-avian dinosaur.

Although Aquilops represents everything I normally stand against – ornithischians, microvertebrates, heads – I confess that I have a sneaking affection for our wee beastie. Somebody’s just gotta make a little plush Aquilops, right? When and if that happens, you know where to find me.



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