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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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.

So, this happened today

March 28, 2013

Big Bend Alamosaurus cervical J with Matt for scale

Mounted Alamosaurus in Dallas 1

Next week I’m going to visit the Perot Museum of Nature and Science in Dallas, Texas, to see their big Alamosaurus (these photos were kindly provided by Ron Tykoski of the Perot Museum, with permission to post). See that sweet string of cervical vertebrae in front of the mounted skeleton? A photo of those same vertebrae when they were still in the ground was featured in the post “How big was Alamosaurus?” three and a half years ago. Happily now they are out of the ground, prepped, and on display, and Tony Fiorillo and Ron Tykoski are working on getting them and some other new Alamosaurus material described.

Mounted Alamosaurus in Dallas 2

Here’s another view of that mount. You may be wondering, first, how legit is it, and second, how big is it? Happily, I have answers for you. In email messages with permission to cite, Ron Tykoski wrote,

The Alamosaurus skeletal mount by RCI  in the photos is based upon scaling the Smithsonian and UT Austin material to match the size of our cervicals here in Dallas.  There were enough overlapping parts between the pieces at the three institutions to get the proportions pretty nicely supported.

I ran across your SV-POW thread on ‘How big was Alamosaurus?’ back when you first posted it in ‘09.  You ought to be pleased to know that you came remarkably close to the eventual size of the skeleton we wound up with.  The full skeleton RCI generated (again, based off scaling to the Dallas verts) is 84ft long, about 16ft at the shoulder (I dropped a tape measure from the 1st dorsal neural spine to the floor during skeleton construction and got 480cm-490cm), and a neck + head of about 25ft.  The overall length and neck length were provided by RCI after fabrication and assembly.   That shoulder height is a bit suspect though based on the positioning of the pectoral girdle in the mount, relative to the ribcage and vert column.   I think the head currently is posed about 25ft or so off the floor, but I can’t verify that (I didn’t get into the scissor-lift to check that at the time).  This skeleton actually played a role in determining the size of the hall in which it is installed.  We decided early in the planning phase for the building that this skeleton would be the centerpiece for the hall.  As a result, the ceilings for this floor had to be made extra-high, and the mid-room support pillars designed out to accommodate the skeleton and still clear all the HVAC, sprinkler heads, and other necessities.

That’s all pretty fantastic–both that we have enough of Alamosaurus to do a pretty rigorous full skeletal mount, and that the beast was legitimately pretty darned big. Ron goes on:

One correction to the story on SV-POW, the Dallas cervical series consists of only 9 verts, not 10.  There may have been frags or something that made folks think there was a 10th at the anterior end of the series when first found, but I’ve never seen evidence of it in our collection.  This may be supported by the fact that the verts were given letter designations in the field (that we still use), and are identified as verts B through J, from anterior to posterior.

I later learned from Tony Fiorillo that the vertebrae were labelled B through J in the field in case anything anterior to B turned up, but nothing did, so the ‘A’ placeholder went unused. That reminds me of the search in the mid-1800s for the hypothetical planet Vulcan (not the one you’re thinking of) between Mercury and the Sun, which I bring up for no reasons other than that hypothetical planets are cool, and if you’re exploring, it’s worth keeping an open mind about what might yet turn up.

There’s more to say about the size of Alamosaurus–we haven’t even covered the big material described by Fowler and Sullivan (2011) yet–but I’m not going to say a whole lot right now, since I’m going to see the Big Bend material in Dallas in just a few days. Watch this space.

Reference

Fowler, D.W. and Sullivan, R.M. 2011. The first giant titanosaurian sauropod from the Upper Cretaceous of North America. Acta Palaeontologica Polonica 56 (4): 685–690.

I was cruising the monographs the other night, looking for new ideas, when the humerus of Opisthocoelicaudia stopped me dead in my tracks. I think you’ll agree it is an arresting sight:

Opisthocoelicaudia right humerus in lateral, anterior, medial, and posterior views, from Borsuk-Bialynicka (1977: figure 7)

Opisthocoelicaudia right humerus in medial, anterior, lateral, and posterior views, from Borsuk-Bialynicka (1977: figure 7)

I’d seen it before, but somehow I had never grokked its grotesque fatness. I mean, damn, Opisthocoelicaudia, you really let yourself go. Especially compared to the slenderness and grace of this juvenile Alamosaurus humerus:

Alamosaurus left humerus in anterior and posterior views, from Lehman and Coulson (2002: figure 7).

Alamosaurus left humerus in anterior and posterior views, from Lehman and Coulson (2002: figure 7).

Now, I realize that part of the slenderness of this Alamosaurus humerus might be because it’s a juvenile–other alamosaur humeri are a bit more robust–but it’s still a striking contrast. I couldn’t help but superimpose them, scaled to the same midshaft width:

Alamosaurus and Opisthocoelicaudia humeri superimposed

I flipped the Alamosaurus humerus left-to-right to match that astonishing lump of Opisthocoelicaudia. The result reminds me of one of Abrell and Thompson’s Actual Facts:

If you put Woodrow Wilson inside William Howard Taft, he would have stuck out by a good 18 inches.

None of that probably signifies anything more than that I am easily amused. And also,  Opisthocoelicaudia is Just Plain Wrong. You hear me, Opisthocoelicaudia? Don’t make me make you cry mayonnaise!

References

Sorry for the very short post. We have some longer stuff planned, but we’ve been too busy to kick it out this week, and I wanted to leave you with something cool to ponder over the weekend. Here’s the ilium of Giraffatitan overlaid on that of Brontomerus, scaled to the same acetabulum diameter (Giraffatitan is HMN J1, left ilium, modified from Janensch 1961: pl. E, fig. 2; Brontomerus is of course OMNH 66430 from Taylor et al. 2011:fig. 2).

And here’s the same thing comparing Rapetosaurus and Brontomerus (Rapetosaurus is holotype FMNH PR 2209, left ilium, modified from Curry Rogers 2009: fig. 39B). This one was tricky to scale because the ilial margin of the acetabulum is so different in the two taxa.

Here is the same trick performed with the ilium of the canonical pretty basal neosauropod Camarasaurus — specifically, Camarasaurus supremus AMNH 5761 Il. 1, left ilium, modified from Osborn and Mook (1921: fig. 87).  In this case, the proportions are so very different that it’s hard to make a meaningful superimposition: we tried to scale to equal acetabulum size, but probably that of the Camarasaurus was proportionally larger than in the other taxa illustrated in this post.  Still, here it is:

Finally, in response to Paul Barrett’s comment on a subsequent article, here is a superimposition of the ilium of Alamosaurus on that of Brontomerus:

(Sorry about the poor quality of this one, but the only figure I could find of a complete Alamosaurus ilium was the line-drawing in Lehman and Coulson (2002:fig. 8) — none of the standard descriptive works seem to illustrate a complete or near-complete ilium.)

We had a figure like these in an early draft of the paper, but we ditched it because we felt that doing a broader comparative figure would be more valuable. But I like the kick in the brainpan that these overlays provide.

References

A section of the cotyle of a presacral vertebra of Alamosaurus (Woodward and Lehman 2009:fig. 6A).

The last time we talked about Alamosaurus, I promised to explain what the arrow in the above image is all about. The image above is a section through the cotyle (the bony socket of a ball-and-socket joint) at the end of one of the presacral vertebra. The external bone surface would have been over on the left; it was either very thin (which happens) or a bit eroded, or both. The arrow is pointing at something weird–a plate of bone inside the vertebra that forms a sort of shadow cotyle deep to the articular surface.

This is weird for a couple of reasons. First, once camellate (small-chambered) vertebrae get above a certain level of complexity, it’s hard to make any sense of the orientation of individual bony struts. Possibly I haven’t seen enough vertebrae, or played with enough 3D models, to figure it out. You would certainly expect that the struts would be oriented to resist biomechanical loads, just like the struts in the long bones of your limbs; the fact that sauropod verts were filled with air whereas your long bones are filled with marrow shouldn’t make any difference. Back in the day, Kent Sanders–who is second author on that super-important paper on unidirectional air flow in croc lungs that you’ve probably heard about (Farmer and Sanders 2010)–speculated to me that the complex of laminae we see in the vertebrae of most sauropods are still there in the inflated-looking vertebrae of titanosaurs and birds, they’re just incarnated in internal struts rather than external laminae. Cool hypothesis for somebody to test.

The other reason that this is weird is that the plate of bone is parallel to the articular surface. One place where I have seen some regularity in terms of strut orientation is in zygapophyses, where in both camerae and camellate vertebrae the internal struts are oriented at right angles to the articular surfaces of the zygs, like beams propping up a wall. In this Alamosaurus section, there are indeed smaller struts that run at right angles to both the cotyle and the internal plate, but I have no idea why they’re so wimpy and the plate is so thick; a priori I would have expected the reverse.

It turns out that this isn’t even the first time that an internal “shadow” of the cotyle has been figured–check out this figure that I redrew from Powell’s (1992:fig. 16) Saltasaurus osteology. But don’t credit me with the discovery. I’d looked at this section a hundred times and even drawn it and never noticed the shadow cotyle, until it was pointed out by Woodward and Lehman (2009)–another reason to read that paper if you haven’t yet. Kudos to Holly Woodward for spotting this and making the connection.

Now that I’ve drawn attention to the weirdness and given credit where it’s due, this is one of those times I’m going to throw up my hands in confusion and open the floor for comments.

References

  • Farmer, C.G., and Sanders, K. 2010. Unidirectional airflow in the lungs of alligators. Science 327:338-340.
  • Powell, J.E. 1992. Osteologia de Saltasaurus loricatus (Sauropoda – Titanosauridae) del Cretacico Superior del noroeste Argentino; pp. 165-230 in J.L. Sanz and A.D. Buscalioni (editors), Los Dinosaurios y Su Entorno Biotico: Actas del Segundo Curso de Paleontologia in Cuenca. Institutio Juan de Valdes.
  • Woodward, H.N.,  and Lehman, T.M. 2009. Bone histology and microanatomy of Alamosaurus sanjuanensis (Sauropoda: Titanosauria) from the Maastrichtian of Big Bend National Park, Texas. Journal of Vertebrate Paleontology 29(3):807-821.