AMNH 460 skeleton model 2

In a recent post I showed some photos of the mounted apatosaurine at the American Museum of Natural History in New York, AMNH 460, which Tschopp et al. (2015) regarded as an indeterminate apatosaurine pending further study.

A lot of museums whose collections and exhibits go back to the late 19th and early 20th centuries have scale model skeletons and sculptures that were used to guide exhibit design. I have always been fascinated by these models, partly because they’re windows into another era of scientific research and science communication, and partly because they’re just cool – basically the world’s best dinosaur toys – and I covet them. In my experience, it is very, very common to find these treasures of history buried in collections, stuck up on top of specimen cabinets, or otherwise relegated to some out-of-the-way corner where they won’t be in the way. I know that exhibit space is always limited, and these old models often reflect ideas about anatomy, posture, or behavior that we now know to be mistaken. But I am always secretly thrilled when I see these old models still on exhibit.

AMNH T rex skeleton model

The AMNH has a bunch of these things, because Henry Fairfield Osborn was crazy about ’em. He not only used 2D skeletal reconstructions and 3D model skeletons to guide exhibit design, he published on them – see for example his 1898 paper on models of extinct vertebrates, his 1913 paper on skeleton reconstructions of Tyrannosaurus, and his 1919 paper with Charles Mook on reconstructing Camarasaurus. That genre of scientific paper seems to have disappeared. I wonder if the time is right for a resurgence.

So in a glass case at the feet of AMNH 460 is a model – I’d guess about 1/12 or 1/15 scale – of that very skeleton. You can tell that it’s a model of that particular skeleton and not just some average apatosaur by looking carefully at the vertebrae. Apatosaurines weren’t all stamped from quite the same mold and the individual peculiarities of AMNH 460 are captured in the model. It’s an amazing piece of work.

AMNH 460 skeleton model

The only bad thing about it is that – like almost everything behind glass at the AMNH – it’s very difficult to photograph without getting a recursive hell of reflections. But at least it’s out where people can see and marvel at it.

Oh, and those are the cervical vertebrae of Barosaurus behind it – Mike and I spent more time trying to look and shoot past this model than we did looking at it. But that’s not the model’s fault, those Barosaurus cervicals are just ridiculously inaccessible.

So, memo to museums: at least some of us out here are nuts about your old dinosaur models, and where there’s room to put them on exhibit, they make us happy. They also give us views of the skeletons that we can’t get otherwise, so they serve a useful education and scientific purpose. More, please.

References

Osborn, H. F. (1898). Models of extinct vertebrates. Science, New Series, 7(192): 841-845.

Osborn, H.F. (1913). Tyrannosaurus, restoration and model of the skeleton. Bulletin of the American Museum of Natural History, 32: 91-92, plates 4-6.

Osborn, H. F., & Mook, C. C. (1919). Characters and restoration of the sauropod genus Camarasaurus Cope. From type material in the Cope Collection in the American Museum of Natural History. Proceedings of the American Philosophical Society, 58(6): 386-396.

AMNH 460 left anterolateral view

Apatosaurines on the brain right now.

I’ve been thinking about the question raised by Jerry Alpern, a volunteer tour guide at the AMNH, regarding the recent Tschopp et al. (2015) diplodocid phylogeny. Namely, if AMNH 460 is now an indeterminate apatosaurine, pending further study, what should the museum and its docents tell the public about it?

Geez, Apatosaurus, it’s not like we’re married!

I think it’s a genuinely hard problem because scientific and lay perspectives on facts and hypotheses often differ. If I say, “This animal is Apatosaurus“, that’s a fact if I’m talking about YPM 1860, the genoholotype of Apatosaurus ajax; it would continue to be a fact even if Apatosaurus was sunk into another genus (as Brontosaurus was for so long). We might call that specimen something else, but there would always be a footnote pointing out that it was still the holotype of A. ajax, even if the A. part was at least temporarily defunct – the scientific equivalent of a maiden name.* For every other specimen in the world, the statement, “This animal is Apatosaurus” is a hypothesis about relatedness, subject to further revision.

* This is going to sound kinda horrible, but when one partner in a marriage takes the other’s surname, that’s a nomenclatural hypothesis about the future of the relationship.

Apatosaurine cervicals are the best cervicals.

Apatosaurine cervicals are the best cervicals.

Fuzzy science

Things that look fairly solid and unchanging from a distance – specifically, from the perspective of the public – often (always?) turn out to be fairly fuzzy or even arbitrary upon closer inspection. Like what is Apatosaurus (beyond the holotype, I mean) – or indeed, what is a planet.** There is no absolute truth to quest for here, only categories and hypotheses that scientists have made up so that we can have constructive conversations about the crazy spectrum of possibilities that nature presents us. We try to ground those categories and hypotheses in evidence, but there will always be edge cases, and words will always break down if you push them too hard. Those of us who work on the ragged frontier of science tend to be fairly comfortable with these inescapable uncertainties, but I can understand why people might get frustrated when they just want to know what the damned dinosaur is called.

** Triton, the largest body orbiting Neptune, is almost certainly a captured Kuiper Belt object, and it’s bigger than Pluto. Moon or planet? Probably best to say a former dwarf planet currently operating as a satellite of Neptune – but that’s a mouthful (and a mindful, if you stop to think about it), not a short, convenient, easily-digestible label. Any short label is going to omit important information. This is related to the problem of paper title length – below some threshold, making something shorter means making it incomplete.

What I would say

I suppose the short version that is most faithful to the Tschopp et al. results is:

This skeleton (AMNH 460) might be Apatosaurus or Brontosaurus or a third, new thing – scientists aren’t sure yet.

A reasonable follow-up sentence – and an answer to the inevitable “Why not?” – would be:

They have to look at 477 anatomical details for lots of skeletons and weigh all the evidence, and that takes time.

Personally, if I was talking to museum visitors I would lean in conspiratorially and say:

If you want to call it Apatosaurus or Brontosaurus, go ahead – those are both ‘live’ hypotheses, and even the world’s experts on this problem can’t tell you that you’re guessing the wrong way – at least not yet.

And if there was a kid in the group, I’d add:

Maybe you’ll be the one to figure it out!

What would you say?

My neck is fat.

My neck is fat.

P.S. I wouldn’t change the signage. It could still turn out to be Apatosaurus, and the Tschopp et al. results do not lend themselves to easy label-ification.

P.P.S. With some modification for taxonomy, all of this applies to the Field Museum diplodocid FMNH P25112 as well.

Reference

Tschopp E, Mateus O, Benson RBJ. (2015) A specimen-level phylogenetic analysis and taxonomic revision of Diplodocidae (Dinosauria, Sauropoda) PeerJ3:e857 https://dx.doi.org/10.7717/peerj.857

Look on my works, ye mighty, and despair!

DSCN0476

[Giraffatitan brancai paralectotype MB.R.2181 (formerly HMN S II), mounted skeleton in left anteroventrolateral view. Presacral vertebrae sculpted, skull scaled and 3d-printed from specimen T1. Round the decay of that colossal wreck, boundless and bare, the lone and level sands stretch far away.]

Brontosaurus, the animal formerly known as Apatosaurus, the animal formerly known as Brontosaurus.

YPM 1980: Brontosaurus excelsus, the animal formerly known as Apatosaurus excelsus, the animal formerly known as Brontosaurus excelsus.

Today is a good day for sauropod science. Since we’re not getting this up until the afternoon, you’ve probably already seen that Emanuel Tschopp and colleagues have published a monstrous specimen-level phylogenetic analysis of Diplodocidae and, among other things, resurrected Brontosaurus as a valid genus. The paper is in PeerJ so you can read it for free (here).

I’ve already been pinged by lots of folks asking for my thoughts on this. I know that the return of Brontosaurus is what’s going to catapult this paper into the spotlight, but I hope what everyone takes away from it is just what a thorough piece of work it is. I’ve never seen so many phylogenetic characters illustrated so well. It sets a new standard, and anyone who wants to overturn this had better roll up their sleeves and bring a boatload of data. I’m also very, very happy that it’s open-access so everyone in the world can see it, use it, question it, tear it apart or build on it. Getting Brontosaurus back is just gravy. Although, being pro-brontosaur enough to have named a dinosaur in honor of Brontosaurus, I’m also pretty happy about that. If you need a quick guide to who’s who now, A. ajax and A. louisae are still Apatosaurus, and B. excelsus, B. yahnahpin (formerly Eobrontosaurus), and B. parvus (originally Elosaurus) are all Brontosaurus. For more details, go read the paper.

A louisae from Wikipedia - full

Apatosaurus lousiae CM 3018: still Apatosaurus. Photo from Wikipedia.

My personal feelings aside, a lot of people are asking how solid is this generic re-separation. I haven’t read the entire paper yet – it’s 299 pages long, for crying out loud – but the separation of Brontosaurus and Apatosaurus seems solid enough. Tschopp et al. didn’t do it lightly, they justify their decision in detail. I don’t hold with the idea that just because two taxa are sisters, means that they cannot be separated generically. As usual in phylogenetic taxonomy, it comes down to what we decide as a community constitutes “diagnosably distinct”. Tschopp et al. have actually put some thought into what that might mean here, and whether you agree with them or not, they’ve at least made all of their evidence and reasoning explicit. That’s both an opportunity and a challenge for critics: an opportunity to pin down exactly where and why you may disagree, and a challenge to do exactly that. You can’t just sit back and say, “I think the analysis is flawed” or “I wouldn’t have coded that character that way” (well, you can, but if that’s all you say, no-one is obliged to take that kind of lazy, drive-by criticism seriously). There are 477 characters here, most of them illustrated, for 81 OTUs, and a lot of post-hoc discussion of the results. So whether you agree with the authors or not, in whole or in part, both fans and critics should dig in and build on this work. Is it the last word on diplodocid taxonomy? Of course not. But it does move the field forward significantly, and the Tschopp et al. should be applauded for that.

There’s a lot more in there than just bringing back Brontosaurus. “Diplodocus” hayi is elevated to its own genus, Galeamopus. Neither of those things are super surprising. There have been rumors since the 90s at least that Brontosaurus might be coming back, and everyone has known for a while that D. hayi was a bit wonky. I was also not surprised to see Australodocus returned to Diplodocidae – when I saw the type material in 2011, it looked diplodocid to me (based on some characters I’ll have to unpack in some other post). More surprising to me are the sinking of Dinheirosaurus into Supersaurus, the finding that Tornieria is not particularly close to Diplodocus, and the uncertain positions of AMNH 460, the American Museum mount, which is an indeterminate apatosaurine pending further study, of FMNH 25112, the Field Museum “Apatosaurus”, which might not even be an apatosaurine at all(!). In several cases, Tschopp et al. come right out and say that X is going to need further study, so if you want to work on sauropods and you’re stuck for project ideas, go see what needs doing.

AMNH mounted Apatosaurus with Taylor for scale

AMNH 460: we don’t know who this is anymore.

As I was scanning the paper again while composing the last paragraph, I almost fell down the rabbit hole. So much interesting stuff in this paper. Even if all you care about is morphology, the hundred or so figures illustrating the phylogenetic characters ought to keep you happy for a very long time. I look forward to reading through the vertebral characters in detail and seeing what I’ve been missing all these years.

I’m contractually obliged to point out that the authors chose to publish the complete peer-review history of the paper, so you can see what the editor (Andy Farke) and reviewers had to say. As always, I think this transparency (and credit for the reviewers) is great for science, and I can’t wait until it’s the norm at more journals.

FMNH 25112 formerly Apatosaurus

FMNH 25112: what even IS that thing?

In addition to the paper, there’s also an interview with lead author Emanuel Tschopp on the PeerJ blog, and a nice shout-out for SV-POW!

Parting shot: why did Tschopp et al. get different results than anyone had previously? Because they used more specimens and more taxa – more data full stop. That’s also why their paper warrants serious consideration. It’s serious work. Let’s go stand on their shoulders.

Reference

Tschopp E, Mateus O, Benson RBJ. (2015) A specimen-level phylogenetic analysis and taxonomic revision of Diplodocidae (Dinosauria, Sauropoda) PeerJ 3:e857 https://dx.doi.org/10.7717/peerj.857

In 2012, Matt and I spent a week in New York, mostly working at the AMNH on Apatosaurusminimus and a few other specimens that caught our eye. But we were able to spend a day at the Yale Peabody Museum up in New Haven, Connecticut, to check out the caudal pneumaticity in the mounted Apatosaurus (= “Brontosaurus“) excelsus, YPM 1980, and the bizarrely broad cervicals of the Barosaurus lentus holotype YPM 429.

While we there, it would have been churlish not to pay some attention to the glorious and justly famous Age of Reptiles mural, painted by Rudolph F. Zallinger from 1944-1947.

So here it is, with the Brontosaurus neck for scale:

IMG_0501-zallinger-mural

Click through for high resolution (3552 × 2664).

And here is a close-up of the most important, charismatic, part of the mural:

IMG_0500-zallinger-mural

Again, click through for high resolution (3552 × 2664).

That’s your lot for now. We’ve long promised a proper photo post of the Brontosaurus mount itself, and I’ll try to get that done soon. For now, it’s just scenery.

Go to Google and do a picture search for “natural history museum”. Here are the results I get. (I’m searching the UK, where that term refers to the British museum of that name — results in the USA may very.)

google-search-for-nhm

In the top 24 images, I see that half of them are of the building itself — rightly so, as it’s a beautiful and impressive piece of architecture that would be well worth visiting even if it was empty. Of the rest, ten are of specimens inside the museum: and every single one of them is of the Diplodocus in the main hall. (The other two photos are from the French natural history museum, so don’t really belong in this set. Not coincidentally, they are both primarily photos of the French cast of the same Diplodocus.)

The NHM’s Diplodocus — I can’t bring myself to call it “Dippy” is the icon of the museum. It’s what kids go to see. It’s what the museum used as the basis of the logo for the 2005 SVPCA meeting that was held there. It’s essentially the museum mascot — the thing that everyone thinks of when they think of the NHM.

And rightly so: it’s not just a beautiful specimen, it’s not just sensational for the kids. As the first cast ever made of the Carnegie specimen CM 84, it’s a historically important object in its own right. It was the first mounted Diplodocus ever, being presented in 1905 before the the original material was even on display in Pittsburgh.

diplodocus_nocopyright

As a matter of fact, this cast was the very first mounted sauropod to be publicly displayed: that honour is usually given to the AMNH Apatosaurus, but as museum-history expert Ilja Nieuwland points out:

The London ‘Dippy’ was in fact the first sauropod on public display, if only for three days in early July of 1904, in the Pittsburgh Exposition Society Hall.

There you have the Natural History Museum Diplodocus: the symbol of the museum, an icon of evolution, a historical monument, a specimen of great scientific value and unparalleled symbolism.

So naturally the museum management want to tear it down. They want to convert the Diplodocus hall into a blue whale hall. Because the museum doesn’t already have a blue whale hall.

Or, no — wait — it does already have a blue whale hall. That’s it. That’s what I meant to say. And very impressive it is, too.

16222408

I don’t mind admitting that the whale hall is my second favourite room in the museum. Whenever I go there as a tourist (rather than as a scientist, when I spend all my time in the basement), I make sure I see it. It’s great.

The thing is, it’s already there. A museum with a whale hall does not need another whale hall.

Obviously anticipating the inevitable outcry, the museum got all its ducks in a row on this. They released some admittedly beautiful concept artwork, and arranged to have opinion pieces written in support of the change — some by people who I would have expected to know better.

One of the more breathtaking parts of this planned substitution is the idea that Diplodocus is no longer relevant. The NHM’s director, Sir Michael Dixon says the change is “about asking real questions of contemporary relevance”. He says “going forward we want to tell more of these stories about the societally relevant research that we do”. This “relevance” rhetoric is everywhere. The museum “must move with the times to stay relevant”, writes Henry Nicholls in the Guardian.

There was a time when Diplodocus was relevant, you know: waaay back in the 1970s. But time has moved on, and now that’s 150,000,035 years old, it’s become outdated.

Conversely, the rationale for the whale seems to be that they want to use it as a warning about extinction. But could there ever be a more powerful icon of extinction than a dinosaur?

The thing is, the right solution is so obvious. Here’s what they want to do:

2528769B00000578-2930638-image-a-19_1422525497076

Clearly the solution is, yes, hang the whale from the ceiling — but don’t remove the Diplodocus. Because, seriously, what could be a better warning about extinction than the juxtaposition of a glorious animal that we lost with one that we could be about to lose?

All this argument about which is better, a Diplodocus or a blue whale: what a waste of energy. Why should we have to choose? Let’s have both.

I’ve even had an artist’s impression made, at great expense, to show how the combination exhibit would look. Check it out.

2528769B00000578-2930638-image-a-19_1422525497076-art

(If anyone would like to attempt an even better rendering, please by my guest. Let me know, and I’ll add artwork to this page.)

So that’s my solution. Keep the museum’s iconic, defining centrepiece — and add some more awesome instead of exchanging it. Everyone wins.

A while back, Ben Miller reminded me that when I posted about the old Yale “Brontosaurus” skull, I promised:

So how did the YPM come to make such a monstrosity? What was it based on? Tune in next time for the surprising details!

I told him at the time that I’d soon get around to writing a post. But before I did, he wrote a post on this himself: Bully for Camarasaurus. And it’s excellent. Go and read it!

I don’t have a lot to add to what Ben has written, except regarding this:

What Marsh had instead [when restoring the skull for his 1891 “Brontosaurus” reconstruction] were a few fragmentary bits of Camarasaurus cranial material, plus a snout and jaw (USNM 5730) now considered to be Brachiosaurus.

Here’s what Marsh came up with:

Marsh1891-plateXVI-Apatosaurus-skull-UNREVERSED

But what of the supposed Brachiosaurus skull that he used as a reference? It was finally described 107 years later by Carpenter and Tidwell (1998), in a paper that helpfully also lays out the history behind it. Here’s how it looks:

CarpenterTidwell1998-fig1

The skull was found by a crew under the supervision of M. P. Felch in the western part of his Quarry 1, Garden Park, Colorado. Felch reported it to O. C. Marsh in a letter of 8 September 1883. It was found by a meter-long cervical vertebra that probably belonged to Brachiosaurus “which was destroyed during attempts to collect it” (McIntosh and Berman 1975:196). [Of course, Felch and Marsh could hardly have been expected to identify this vertebra correctly, as Brachiosaurus would not be discovered and named for another twenty years (Riggs 1903), and the nature of its neck would not become apparent until Janensch (1914) described the related brachiosaurid Giraffatitan (= “Brachiosaurus“) brancai.]

The Felch skull, along with other material from the quarry, was shipped to Marsh at Yale in October of that year, and was initially assigned the specimen number YPM 1986. At that time it was only partially prepared, hence the rather poor resemblance between the restored version above and Marsh’s hypothetical “Brontosaurus” [= Apatosaurus] skull that was based on it.

It’s notable that Holland (1915) was quite certain that this was not a skull of Brontosaurus, and that a Diplodocus-like skull found with the A. louisae holotype belonged to it. It’s worth reading the skull section of his paper to see just how solid his reasoning was. And it’s extraordinary to think that Osborn’s power, all the way over in New York, was so great that he was able to successfully bully Holland, 370 miles away in Pittsburgh, into not putting the evidently correct skull on the Carnegie Museum’s Apatosaurus mount. That mount remained sadly headless until after Holland’s death.

Aaanyway, YPM 1986 was pretty much ignored after Marsh’s abuse of it as a reference for the Brontosaurus reconstruction’s skull. After Marsh’s death in 1899, much of the material collected by Felch was transferred to the Smithsonian (US National Museum of Natural History). The skull was among these specimens, and so was re-catalogued as USNM 5730.

As so often, it was Jack McIntosh who rediscovered this skull and recognised its true affinities. Some time after his tentative identification of the skull as pertaining to Brachiosaurus (presumably on the basis of its resemblance to that of Giraffatitan), Carpenter borrowed the skull, had it more fully prepared, wrote the description, and had a restored model constructed from casts of the preserved elements and models of the missing ones.

Carpenter and Tidwell (1998:fig. 2) also handily showed the restored Felch quarry skull alongside those of other sauropods:

CarpenterTidwell1998-fig2

By re-ordering the top row, we can see what a neat intermediate it is between the skulls of Camarasaurus (left) and Giraffatitan (= “Brachiosaurus” of their usage):

CarpenterTidwell1998-fig2-top-row-reordered

I provisionally accepted USNM 5730 as belonging to Brachiosaurus in my re-evaluation of 2009, and included it in my reconstruction (Taylor 2009:fig. 7):

Taylor (2007: figure 7). Skeletal reconstruction of Brachiosaurus altithorax. White bones represent the elements of the holotype FMNH P 25107. Light grey bones represent material referred to B. altithorax: the Felch Quarry skull USNM 5730, the cervical vertebrae BYU 12866 (C?5) and BYU 12867 (C?10), the "Ultrasauros" scapulocoracoid BYU 9462, the Potter Creek left humerus USNM 21903, left radius and right metacarpal III BYU 4744, and the left metacarpal II OMNH 01138. Dark grey bones modified from Paul's (1988) reconstruction of Giraffatitan brancai. Scale bar equals 2 m.

Taylor (2007: figure 7). Skeletal reconstruction of Brachiosaurus altithorax. White bones represent the elements of the holotype FMNH P 25107. Light grey bones represent material referred to B. altithorax: the Felch Quarry skull USNM 5730, the cervical vertebrae BYU 12866 (C?5) and BYU 12867 (C?10), the “Ultrasauros” scapulocoracoid BYU 9462, the Potter Creek left humerus USNM 21903, left radius and right metacarpal III BYU 4744, and the left metacarpal II OMNH 01138. Dark grey bones modified from Paul’s (1988) reconstruction of Giraffatitan brancai. Scale bar equals 2 m.

But as noted by Carpenter and Tidwell (1998:82), the lack of comparable parts between the Felch skull and the Brachiosaurus holotype (which remains the only definitive Brachiosaurus material) means that the assignment has to remain tentative.

What we really need is a more complete Brachiosaurus specimen: one with both a skull and good postcervical elements that let us refer it definitively to Brachiosaurus altithorax by comparison with the holotype. And heck, while we’re at it, let’s have a specimen with a good neck, too!

The real question remains: how did Marsh, using a brachiosaur skull as his basis, come up with this?

Marsh1891-plateXVI-Apatosaurus-skull-UNREVERSED

 

And stranger still, how someone at the Yale Peabody Museum — we don’t know who — used it, or more likely Marsh’s reconstruction, as a basis for this sculpture:

IMG_0517

 

The Yale mount didn’t go up until 1931 — the last of the Big Four Apatosaurus mounts after the AMNH, Carnegie and Field Museum, which is surprising as it was the first of those specimens to be found. So by the time the skull was sculpted, sauropod skulls were actually reasonably well known. It’s not clear quite how anyone working from a decent reconstruction of, say, a Camarasaurus skull — the one in Osborn and Mook (1921:figure 30), say — could come up with this monster.

The last thing to say is this: it does credit to the YPM that they display this historically important sculpture rather than hiding it away and pretending it never happened. For me, part of the fascination of palaeontology is seeing not just how organisms evolved through prehistory but how ideas evolved through history. It’s great that we can still see important mistakes, alongside their corrections (i.e. the new and lovely skull on the YPM Apatosaurus mount.)

 

References

  • Carpenter, Kenneth, and Virginia Tidwell. 1998. Preliminary description of a Brachiosaurus skull from Felch Quarry 1, Garden Park, Colorado. Modern Geology 23:69-84.
  • Holland, William J. 1915. Heads and tails: a few notes relating to the structure of the sauropod dinosaurs. Annals of the Carnegie Museum 9:273-278.
  • Janensch, Werner. 1914. Ubersicht uber der Wirbeltierfauna der Tendaguru-Schichten nebst einer kurzen Charakterisierung der neu aufgefuhrten Arten von Sauropoden. Archiv fur Biontologie, Berlin III, 1(1):81-110.
  • Marsh, O. C. 1891. Restoration of Triceratops (with plates XV and XVI). American Journal of Science, 3rd series 41(244):339-342.
  • McIntosh, John S., and David, S. Berman. 1975. Description of the palate and lower jaw of the sauropod dinosaur Diplodocus (Reptilia: Saurischia) with remarks on the nature of the skull of Apatosaurus. Journal of Paleontology 49(1):187-199.
  • Osborn, Henry Fairfield, and Charles C. Mook. 1921. Camarasaurus, Amphicoelias and other sauropods of Cope. Memoirs of the American Museum of Natural History, n.s. 3:247-387, and plates LX-LXXXV.
  • Riggs, Elmer S. 1903. Brachiosaurus altithorax, the largest known dinosaur. American Journal of Science 15(4):299-306.
  • Taylor, Michael P. 2009. A re-evaluation of Brachiosaurus altithorax Riggs 1903 (Dinosauria, Sauropoda) and its generic separation from Giraffatitan brancai (Janensch 1914). Journal of Vertebrate Paleontology 29(3):787-806.

 

Follow

Get every new post delivered to your Inbox.

Join 492 other followers