A bunch of stuff, loosely organized by theme.

Media

First up, I need to thank Brian Switek, who invited me to comment on Patagotitan for his piece at Smithsonian. I think he did a great job on that, arguably the best of any of the first-day major media outlet pieces. And it didn’t go unnoticed – his article was referenced at both the Washington Post and NPR (and possibly other outlets, those are the two I know of right now). I don’t think my quotes got around because they’re particularly eloquent, BTW, but rather because reporters tend to like point-counterpoint, and I was apparently the most visible counterpoint. They probably would have done the same if I’d been talking complete nonsense (which, to be fair, some people may think I was).

Paleobiology vs Records

The most commonly reproduced quote of mine is this one, originally from Brian’s piece:

I think it would be more accurate to say that Argentinosaurus, Puertasaurus and Patagotitan are so similar in size that it is impossible for now to say which one was the largest.

That may seem at odds with the, “Well, actually…[pushes glasses up nose]…Argentinosaurus was still biggest” tack I’ve taken both in my post yesterday and on Facebook. So let me elaborate a little.

There is a minor, boring point, which is that when I gave Brian that quote, I’d seen the Patagotitan paper, but not the Electronic Supplementary Materials (ESM), so I knew that Patagotitan was about the same size as the other two (and had known for a while), but I hadn’t had a chance to actually run the numbers.

The much more interesting point is that the size differences between Argentinosaurus, Puertasaurus, and Patagotitan are astonishingly small. The difference between a 2.5m femur and a 2.4m one is negligible, ditto for vertebrae with centra 59cm and 60cm in diameter. OMNH 1331, the biggest centrum bit from the giant Oklahoma apatosaur, had an intact max diameter of 49cm, making it 26% larger in linear terms than the next-largest apatosaur. The centra of these giant South American titanosaurs are more than 20% bigger yet than OMNH 1331, just in linear terms. That’s crazy.

It’s also crazy that these three in particular – Argentinosaurus, Puertasaurus, and Patagotitan – are so similar in size. Dinosaur developmental programs were ‘messy’ compared to those of mammals, both in having weird timings for things like onset of reproduction, and in varying a lot among closely related taxa. Furthermore, sauropod population dynamics should have been highly skewed toward juveniles and subadults. So is the near-equality in size among Argentinosaurus, Puertasaurus, and Patagotitan just a coincidence, or does it mean that something weird was going on? There’s really no third option. I mean, even if some kind of internal (biomechanical or physiological) or external (ecological, food or predation) constraint forced those three to the same adult body size, it’s weird then that we’re finding only or at least mostly near-max-size adults. (If the available specimens of these three aren’t near-max-size, then any hypothesis that they’re forced to the same size by constraints is out the window, and we’re back to coincidence.)

BUT

With all that said, the title of “world’s largest dinosaur” is not handed out for effort expended, number of specimens collected, skeletal completeness, ontogenetic speculation, or anything other than “the dinosaur with the largest measured elements”. And that is currently Argentinosaurus. So although for any kind of paleobiological consideration we can currently consider Argentinosaurus, Puertasaurus, and Patagotitan to all be about the same size – and Alamosaurus, Paralititan, Notocolossus, and probably others I’ve forgotten should be in this conversation – anyone wanting to dethrone Argentinosaurus needs to actually show up with bigger elements.

So, if you’re interested in paleobiology, it’s fascinating and frankly kind of unnerving that so many of these giant titanosaurs were within a hand-span of each other in terms of size. Patagotitan is one more on the pile – and, as I said yesterday, exciting because it’s so complete.

But if you want to know who holds the crown, it’s still Argentinosaurus.

Humeri

In a comment on the last post, Andrea Cau made an excellent point that I am just going to copy here entire:

Even Paralititan stromeri humerus is apparently larger than Patagotitan humerus (169 cm vs 167.5 cm). I know humerus length alone is bad proxy of body size, but at least this shows that even in that bone Patagotitan is just another big titanosaur among a well known gang of titans, not a supersized one.

That made me want to start a list of the longest sauropod humeri. Here goes – if I missed anyone or put down a figure incorrectly, I’m sure you’ll let me know in the comments.

  • Giraffatitan: 213cm
  • Brachiosaurus: 203cm
  • Ruyangosaurus: 190cm (estimated from 135cm partial)
  • Turiasaurus: 179cm
  • Notocolossus: 176cm
  • Paralititan: 169cm
  • Patagotitan: 167.5cm
  • Dreadnoughtus: 160cm
  • Futlognkosaurus: 156cm

Admittedly the Patagotitan humerus is from a paratype and not from the largest individual, but that is true for some others on the list, including Giraffatitan. And we have no humeri from Argentinosaurus, Puertasaurus, and some other giants.

Dorsal Vertebrae

A couple of further thoughts on how the dorsal vertebrae of Patagotitan compare to those of Argentinosaurus. First, now that I’ve had some time to think about it, I have a hard time seeing how the dorsal polygon method used by Carballido et al. in the Patagotitan paper has any biological meaning. In their example figure, the polygon around the Puertasaurus vertebra is mostly full of bone, and the one around Patagotitan has a lot of empty space. It’s easy to imagine an alternative metric, like “area of the minimum polygon actually filled by bone”, that would lead to a different ‘winner’. But that wouldn’t mean much, either.

Something that probably does have a real and important biomechanical meaning is the surface area of the articular face of the centrum, because that’s the area of bone that has to bear the compressive load, which is directly related to the animal’s mass. The biggest Patagotitan centrum is that of MPEF-PV 3400/5, which is at least a local maximum since has smaller centra both ahead and behind. The posterior face measures 59cm wide by 42.5cm tall. Abstracted as an ellipse, which may not be perfectly accurate, those measurements give a surface area of (pi)(29.5)(21.25)=1970 cm^2. For Argentinosaurus, the largest complete centrum has a posterior face measuring 60cm wide by 47cm tall (Bonaparte and Coria 1993: p. 5), giving an elliptical surface area of (pi)(30)(23.5)=2210 cm^2. (I’d use hi-res images of the centra to measure the actual surface areas if I could, but AFAIK those images either don’t exist or at least have not yet been made public, for either taxon.) So although the Argentinosaurus dorsal seems like it is only a bit bigger in linear terms, it’s 12% larger in surface area, and that might actually be a meaningful difference.

Cervical Vertebrae

One thing I haven’t commented on yet – Patagotitan is the newest member of the “world’s longest vertebrae” club. The longest Patagotitan cervical, MPEF-PV 3400/3, is listed in the ESM as having a centrum length of 120cm, but it’s also listed as incomplete. In the skeletal recon in the paper, the centrum is colored in as present, but the neural spine is missing. So is the centrum complete in terms of length? I don’t think it’s clear right now.

Anyway, here’s the current rundown of the longest cervical centra of sauropods (and therefore, the longest vertebrae among animals):

  • BYU 9024, possibly referable to Supersaurus or Barosaurus: 137cm
  • Price River 2 titanosauriform: 129cm
  • OMNH 53062, Sauroposeidon holotype: 125cm
  • KLR1508-77-2, Ruyangosaurus giganteus referred specimen: 124cm
  • MPEF-PV 3400/3, Patagotitan holotype: 120cm (+?)
  • MPM 10002, Puertasaurus holotype: 118cm

You may be surprised to see the Price River 2 cervical in there. It was reported in an SVP abstract a few years ago (I’ll dig up that ref and update this post), and Mike and I saw it last year on the Sauropocalypse. We measured the centrum at 129cm, making it just a bit longer than the longest centrum of Sauroposeidon, and therefore the second-longest vertebra of anything ever.

Aside – I’m probably getting a reputation as a big ole meanie when it comes to debunking “world’s largest dinosaur” claims. If I’m willing to take the lead in kicking my own dinosaur down the ladder, don’t expect me to be kind to yours. I follow where the numbers lead.

Now, here’s an interesting thing – now that Sauroposeidon is coming out as a basal titanosaur, rather than a brachiosaur, it might not have been a skinny freak. The 120cm cervical of Patagotitan makes the 125cm cervical of Sauroposeidon and the 129cm cervical from Price River 2 look even more tantalizing. Maybe it’s super-giant sauropods all the way down.

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“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.

References

Here’s a dorsal vertebra of Camarasaurus in anterior view (from Ostrom & McIntosh 1966, modified by Wilson & Sereno 1998). It is one of the most disturbing things I have ever seen in a sauropod. It makes my skin crawl.

Here’s why: the centrum and the thing we habitually call the ‘neural arch’ aren’t fully fused, and as this modified version makes clear, the ‘neural arch’ is neither neural nor an arch. Instead of being bounded ventrally by the centrum and dorsally and laterally by the neural arch, the neural canal lies entirely below the synchondrosis between the not-really-an-arch and the centrum.

Why?! WHY WOULD YOU DO THAT, CAMARASAURUS? This is not ‘Nam. This is basic vertebral architecture. There are rules.

Look at c6 of Apatosaurus CM 555 here, behaving as all good vertebrae ought to. Neural arch be archin’, as the kids say.

And if you are seeking solace in the thought that maybe the artist just drew that Cam dorsal incorrectly, forget it. I’ve been to Yale and examined the original specimen. I’ve seen things, man!

Camarasaurus isn’t the only pervert around here. Check this out:

Unfused neural arch of a caudal vertebra of a juvenile Alamosaurus from Big Bend. And I mean, this is a neural arch. This may be the most neural of all neural arches, in that it contains the entire neural canal. It’s more of a neural…ring, I guess. That’s right, this Alamosaurus caudal is batting for the opposite team from the Cam dorsal above. And it’s a team that neither you nor I play on, because we have well-behaved normal-ass vertebrae with neural arches that actually arch, and then stop, like God and Richard Owen intended.

Scientifically, my question about these vertebrae is: well, that is, I mean to say, what!? I think they have damaged me in some fundamental way.

If you have anything more intelligent to add (or even less intelligent – consider the gauntlet thrown down!), the comment thread is open.

References

  • Ostrom, John H., and John S. McIntosh. 1966. Marsh’s Dinosaurs. Yale University Press, New Haven and London. 388 pages including 65 absurdly beautiful plates.
  • Wilson, J. A. and Paul C. Sereno. 1998. Early evolution and higher-level phylogeny of sauropod dinosaurs. Society of Vertebrate Paleontology, Memoir 5: 1-68.

Enter Sarmientosaurus

April 28, 2016

journal.pone.0151661.g006

Fig 6. Cranium of Sarmientosaurus musacchioi gen. et sp. nov. (MDT-PV 2). Computed tomography-based digital visualization in right lateral (A), left lateral (B), rostral (C), caudal (D), dorsal (E), and ventral (F) views. Scale bar = 10 cm. http://dx.doi.org/10.1371/journal.pone.0151661.g006

Yesterday we got a treat: the description of a new titanosaur, Sarmientosaurus musacchioi, based on some decent cervical vertebrae and an almost absurdly nice skull from the Upper Cretaceous of Argentina (Martinez et al., 2016). It was published in PLOS ONE so it’s free to the world, including a 3D PDF of the skull and some awesome visualizations. Get all that good stuff here.

I had one day’s warning about this – Brian Switek contacted me on Monday to ask if I’d be willing to lend my thoughts on the new critter for his news article for National Geographic, which you can read here. As always, I sent more stuff than he could use, so I’m recycling the long form for the rest of this post.

Brian’s first question was about how Sarmientosaurus stands out. I wrote:

Sarmientosaurus has probably the most complete and best-preserved skull of any sauropod from South America to date. It’s kind of funny – for so long we had so few good skulls from brachiosaurs and titanosaurs, and now we’re getting them, but without much of the rest of the skeleton. In North America, unquestionably the nicest Cretaceous sauropod skull is that of the brachiosaurid Abydosaurus, but all we have with the skull is a bit of the neck. Same situation now with this new titanosaur, Sarmientosaurus. I’m not complaining – great skulls without bodies are still great skulls! – but it will be nice to someday connect heads and bodies.

Also, the authors are to be commended – I don’t think anyone has ever done such a thorough job describing the skull of a sauropod dinosaur. This paper will become the standard to which all others are compared going forward.

I stand by all of that. This new paper is just ridiculous in quantity and quality of descriptive detail. Do you like technicolor sauropod palates? Here, have a technicolor sauropod palate:

journal.pone.0151661.g008

Fig 8. Palate of Sarmientosaurus musacchioi gen. et sp. nov. (MDT-PV 2). Computed tomography-based digital visualization in ventral view indicating palatal bones (ectopterygoids, palatines, pterygoids, and vomers) and the right suborbital fenestra. Abbreviations see text. Scale bar = 10 cm. http://dx.doi.org/10.1371/journal.pone.0151661.g008

The next question from Brian was about the head posture and the inference drawn by Martinez et al. (2016) that Sarmientosaurus fed at ground level. My take:

It doesn’t seem unlikely to me that Sarmientosaurus had a downward-facing snout. As for being a low grazer, I am skeptical. The inner ear usually tells us something about the alert posture of an animal, not its feeding posture. Take rhinos – some of them graze from the ground, and some of them browse up higher, but they all carry their heads the same way. Most grazers have wide snouts, whereas that of Sarmientosaurus is pointed and even a little narrower than that of Giraffatitan. That’s a curious shape for a supposed grazer.

So there are three points to unpack here. First, I chose my words deliberately in saying that the inner ear tells us “something” about the alert posture, because in fact the horizontal semicircular canals (HSCCs) aren’t great even at that. As I wrote in this post seven years ago:

Where SCCs have really attracted attention in paleontology is the “more or less” horizontal orientation of the HSCCs in living animals. Some authors have argued that if you set the HSCCs level or close to level, you can figure out how the head was oriented in life.

Well, maybe. The problem is that there is a LOT of variation around level. In birds surveyed by Duijm (1951), HSCC orientation varied by 50 degrees among taxa, from 20 degrees below horizontal to 30 degrees above. Furthermore, in humans HSCC orientation varies by up to 20 degrees among individuals. Possibly humans are weirdly variable, but it seems at least equally likely that most critters are and we’ve only discovered that variation in humans because of the huge sample size.

However you slice it, those are darn big error bars around any given head posture. That doesn’t mean that HSCC orientations in dinosaurs and other extinct vertebrates are worthless for determining posture (they may also be a source of taxonomic information). Strictly speaking, it means that preserved HSCCs can get us in the 50-degree ballpark but can’t narrow things down any further. This is one of those areas in paleontology where we’re just going to have to live with a certain amount of uncertainty, at least for now.

As far as I know, that’s all still true. But I’d love to be wrong.

Second, there’s the difference between alert posture and feeding posture. Go watch horses graze – the skull is practically vertical while they’re feeding, but that’s not the orientation you get from the HSCCs. So if I’m skeptical about ignoring the error bars around HSCC orientation to determine alert posture, I’m even more skeptical about trying to infer feeding posture from them. Also, the rhino point – we have an extant group with closely related taxa where one is a grazer (white rhino, Ceratotherium) and one is a browser (black rhino, Diceros). They hold their heads about the same. So feeding preference will not necessarily be reflected in normal, non-feeding head posture.

journal.pone.0151661.g034

Fig 34. Comparison of titanosauriform sauropod dinosaur skulls in dorsal view. (A) Giraffatitan brancai (redrawn from Wilson and Sereno [103]). (B) Sarmientosaurus musacchioi gen. et sp. nov. (C) Nemegtosaurus mongoliensis (redrawn from Wilson [11]). (D) Rapetosaurus krausei (redrawn from Curry Rogers and Forster [13]). (E) Tapuiasaurus macedoi (redrawn from Zaher et al. [14]). Not to scale. http://dx.doi.org/10.1371/journal.pone.0151661.g034

Third, muzzle shape. Most grazers have wide mouths, but as I said in the email to Brian – and as this figure shows – the snout of Sarmientosaurus is narrower than that of Giraffatitan, and I don’t think anyone is seriously proposing that Giraffatitan was a grazer. So if Sarmientosaurus was more committed to low-level feeding than more basal titanosauriforms, its face was evolving in the wrong direction. Just sayin’.

(Incidentally, I am hugely in favor of figures like 33 and 34 in Martinez et al., 2016, which make it easy to compare the new critter to a selection of reference taxa. I wish everyone would do this all the time.)

journal.pone.0151661.g033

Fig 33. Comparison of titanosauriform sauropod dinosaur skulls in right lateral view. (A) Giraffatitan brancai (redrawn and modified from Wilson and Sereno [103]). (B) Abydosaurus mcintoshi (redrawn and modified from Chure et al. [98]). (C) Sarmientosaurus musacchioi gen. et sp. nov. (D) Nemegtosaurus mongoliensis (redrawn and modified from Wilson [11]). (E) Rapetosaurus krausei (redrawn from Curry Rogers and Forster [13]). (F) Tapuiasaurus macedoi (redrawn from Zaher et al. [14]). Not to scale. http://dx.doi.org/10.1371/journal.pone.0151661.g033

Finally (final for the purposes of the interview), Brian noted that in the media sauropods are often depicted as all being pretty much the same, and he asked what made Sarmientosaurus stand out. My response:

Until now, the skulls we’ve found of basal titanosauriforms – brachiosaurs and relatives – and more derived titanosaurs haven’t looked much alike. To me Sarmientosaurus is cool because it bridges that gap. From the top and the front the skull looks a lot like those of Brachiosaurus and Giraffatitan – really wide, pretty big teeth, long toothrow. But from the side, the smaller nostrils and long snout have obvious similarities to more derived titanosaurs like Nemegtosaurus. And they phylogenetic analysis confirms that, which is nice. But you can take one look at this thing and say, “Yeah, cool, we’ve been waiting for someone like you.”

The lateral views of titanosauriform skulls in the above figure nicely illustrate my point. If you took the Giraffatitan skull in A and the Tapuiasaurus skull in F and did a 50% morph between them, you’d get something pretty darned close to Sarmientosaurus. And about halfway between Giraffatitan and the really derived saltasaurids is where the phylogenetic analysis puts Sarmientosaurus. The gestalt of the skull nicely reflects the animal’s relationships, which does not always happen.

Oh, there are cervical vertebrae, too, and a seriously weird ossified tendon that is apparently not a cervical rib, but those will keep for another post.

The take-home here is that although I disagree with the authors on a few points of paleobiological interpretation, the Sarmientosaurus fossils are spectacular and Martinez et al. (2016) have done a tremendous job of describing and illustrating them. And the paper is free to anyone who wants it, as it should be. One of the great delights of the last few years has been watching PLOS ONE and PeerJ become the preferred outlets for high-quality descriptive work on dinosaurs.

Now if we can just find more of this thing!

Reference

Martínez RDF, Lamanna MC, Novas FE, Ridgely RC, Casal GA, Martínez JE, et al. (2016) A Basal Lithostrotian Titanosaur (Dinosauria: Sauropoda) with a Complete Skull: Implications for the Evolution and Paleobiology of Titanosauria. PLoS ONE 11(4): e0151661. doi:10.1371/journal.pone.0151661

 

 

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.

References

This just in, from Zurriaguz and Powell’s (2015) hot-off-the-press paper describing the morphology and pneumatic features of the presacral column of the derived titanosaur Saltasaurus. (Thanks to Darren for bringing this paper to my attention.)

Now, as everyone knows, titanosaurs don’t have epipophyses. In fact, they’re the one major sauropod group where Matt has not observed them.

Until today.

Zurriaguz and Powell (2015:figure 3B). Anterior cervical vertebra PVL 4017-3 of Saltasaurus loricatus, in dorsal view (rotated 90° from the paper)

Zurriaguz and Powell (2015:figure 3B). Anterior cervical vertebra PVL 4017-3 of Saltasaurus loricatus, in dorsal view (rotated 90° from the paper)

Look at the left postzygapophysis, at top left of this image. Doesn’t that look like there’s a distinct rounded eminence sticking out towards the camera?

No? Not convinced? All right, then, how about this?

Zurriaguz and Powell (2015:figure 4B). Mid-anterior cervical PVL 4017-138 of Saltasaurus loricatus in right lateral view.

Zurriaguz and Powell (2015:figure 4B). Mid-anterior cervical PVL 4017-138 of Saltasaurus loricatus in right lateral view.

This time, look at the right postzyg (again at top left in the image). Doesn’t that look like there are two separate bony structures up there separated by a notch? A postzygapophyseal facet below, and an epipophysis above? Right?

Huh? What’s that? Just damage, you say?

All right. Let’s bring out the smoking gun.

Zurriaguz and Powell (2015:figure 5). Last anterior cervical vertebra (PVL 4017-5) of Saltasaurus loricatus in right lateral view. (Ignore the inset square for our purposes: it's in the original.)

Zurriaguz and Powell (2015:figure 5). Last anterior cervical vertebra (PVL 4017-5) of Saltasaurus loricatus in right lateral view. (Ignore the inset square for our purposes: it’s in the original.)

Again up at top left, we seem to have a clear case of a ventrally directed postzygapophyseal facet surmounted by a separate eminence which can only be an epipophysis. It even seems to be roughened for tendon attachment.

What does this mean? Only the same thing we said last time: The more we look for epipophyses, the more we find them. Amazing how often that turns out to be true of various things.

We seem to be headed towards the conclusion that epipophyses, while never ubiquitous, pop up in all sorts of places scattered all across the ornithodiran tree, encompassing birds, other theropods, sauropods, prosauropods, several groups of ornithischians, and both pterodactyloid and “rhamphorhynchoid” pterosaurs.

But what about outside Ornithodira?

Can we find epipophyses even out there, in the wilderness?

Stay tuned!

References

In a comment on the last post, on the mass of Dreadnoughtus, Asier Larramendi wrote:

The body mass should be considerably lower because the reconstructed column don’t match with published vertebrae centra lengths. 3D reconstruction also leaves too much space between vertebrae. The reconstruction body trunk is probably 15-20% longer than it really was. Check the supplementary material: http://www.nature.com/srep/2014/140904/srep06196/extref/srep06196-s1.pdf

So I did. The table of measurements in the supplementary material is admirably complete. For all of the available dorsal vertebrae except D9, which I suppose must have been too poorly preserved to measure the difference, Lacovara et al. list both the total centrum length and the centrum length minus the anterior condyle. Centrum length minus the condyle is what in my disseration I referred to as “functional length”, since it’s the length that the vertebra actually contributes to the articulated series, assuming that the condyle of one vertebra sticks out about as far as the cotyle is recessed on the next vertebra. Here are total lengths/functional lengths/differences for the seven preserved dorsals, in mm:

  • D4 – 400/305/95
  • D5 – 470/320/150
  • D6 – 200/180/20
  • D7 – 300/260/40
  • D8 – 350/270/80
  • D9 – 410/ – / –
  • D10 – 330/225/105

The average difference between functional length and total length is 82 mm. If we apply that to D9 to estimate it’s functional length, we get 330mm. The summed functional lengths of the seven preserved vertebrae are then 1890 mm. What about the missing D1-D3? Since the charge is that Lacovara et al. (2014) restored Dreadnoughtus with a too-long torso, we should be as generous as possible in estimating the lengths of the missing dorsals. In Malawisaurus the centrum lengths of D1-D3 are all less than or equal to that of D4, which is the longest vertebra in the series (Gomani 2005: table 3), so it seems simplest here to assign D1-D3 functional lengths of 320 mm. That brings the total functional length of the dorsal vertebral column to 2850 mm, or 2.85 m.

At this point on my first pass, I was thinking that Lacovara et al. (2014) were in trouble. In the skeletal reconstruction that I used for the GDI work in the last post, I measured the length of the dorsal vertebral column as 149 pixels. Divided by 36 px/m gives a summed dorsal length of 4.1 m. That’s more than 40% longer than the summed functional lengths of the vertebrae calculated above (4.1/2.85 = 1.44). Had Lacovara et al. really blown it that badly?

Before we can rule on that, we have to estimate how much cartilage separated the dorsal vertebrae. This is a subject of more than passing interest here at SV-POW! Towers–the only applicable data I know of are the measurements of intervertebral spacing in two juvenile apatosaurs that Mike and I reported in our cartilage paper last year (Taylor and Wedel 2013: table 3, and see this post). We found that the invertebral cartilage thickness equaled 15-24% of the length of the centra.* For the estimated 2.85-meter dorsal column of Dreadnoughtus, that means 43-68 cm of cartilage (4.3-6.8 cm of cartilage per joint), for an in vivo dorsal column length of 3.28-3.53 meters. That’s still about 15-20% shorter than the 4.1 meters I measured from the skeletal recon–and, I must note, exactly what Asier stated in his comment. All my noodling has accomplished is to verify that his presumably off-the-cuff estimate was spot on. But is that a big deal?

Visually, a 20% shorter torso makes a small but noticeable difference. Check out the original reconstruction (top) with the 20%-shorter-torso version (bottom):

Dreadnoughtus shortened torso comparison - Lacovara et al 2014 fig 2

FWIW, the bottom version looks a lot more plausible to my eye–I hadn’t realized quite how weiner-dog-y the original recon is until I saw it next to the shortened version.

In terms of body mass, the difference is major. You’ll recall that I estimated the torso volume of Dreadnoughtus at 32 cubic meters. Lopping off 20% means losing 6.4 cubic meters–about the same volume as a big bull elephant, or all four of Dreadnoughtus‘s limbs put together. Even assuming a low whole-body density of 0.7 g/cm^3, that’s 4.5 metric tons off the estimated mass. So a ~30-ton Dreadnoughtus is looking more plausible by the minute.

For more on how torso length can affect the visual appearance and estimated mass of an animal, see this post and Taylor (2009).

* I asked Mike to do a review pass on this post before I published, and regarding the intervertebral spacing derived from the juvenile apatosaurs, he wrote:

That 15-24% is for juveniles. For the cervicals of adult Sauroposeidon we got about 5%. Why the differences? Three reasons might be relevant: 1, taxonomic difference between Sauroposeidon and Apatosaurus; 2, serial difference between neck and torso; 3, ontogenetic difference between juvenile and adult. By applying the juvenile Apatosaurus dorsal measurement directly to the adult Dreadnoughtus dorsals, you’re implicitly assuming that the adult/juvenile axis is irrelevant (which seems unlikely to me), that the taxonomic axis is (I guess) unknowable, and that the cervical/dorsal distinction is the only one that matter.

That’s a solid point, and it deserves a post of its own, which I’m already working on. For now, it seems intuitively obvious to me that we got a low percentage on Sauroposeidon simply because the vertebrae are so long. If the length-to-diameter ratio was 2.5 instead of 5, we’d have gotten 10%, unless cartilage thickness scales with centrum length, which seems unlikely. For a dorsal with EI of 1.5, cartilage thickness would then be 20%, which is about what I figured above.

Now, admittedly that is arm-waving, not science (and really just a wordy restatement of his point #2). The obvious thing to do is take all of our data and see if intervertebral spacing is more closely correlated with centrum length or centrum diameter. Now that it’s occurred to me, it seems very silly not to have done that in the actual paper. And I will do that very thing in an upcoming post. For now I’ll just note three things:

  1. As you can see from figure 15 in our cartilage paper, in the opisthocoelous anterior dorsals of CM 3390, the condyle of the posterior vertebra is firmly engaged in the cotyle of the anterior one, and if anything the two vertebrae look jammed together, not drifted apart. But the intervertebral spacing as a fraction of centrum length is still huge (20+4%) because the centra are so short.
  2. Transferring these numbers to Dreadnoughtus only results in 4.3-6.8 cm of cartilage between adjacent vertebrae, which does not seem unreasonable for a 30- or 40-ton animal with dorsal centra averaging 35 cm in diameter. If you asked me off the cuff what I thought a reasonable intervertebral spacing was for such a large animal, I would have said 3 or 4 inches (7.5 to 10 cm), so the numbers I got through cross-scaling are actually lower than what I would have guessed.
  3. Finally, if I’ve overestimated the intervertebral spacing, then the actual torso length of Dreadnoughtus was even shorter than that illustrated above, and the volumetric mass estimate would be smaller still. So in going with relatively thick cartilage, I’m being as generous as possible to the Lacovara et al. (2014) skeletal reconstruction (and indirectly to their super-high allometry-derived mass estimate), which I think is only fair.

References