## The sauropod neck mass project: an experiment in open science

### March 4, 2013

Brachiosaurus sp. BYU 12866 c5? in left lateral view with CT slices, some corrected for distortion.

Last Tuesday Mike popped up in Gchat to ask me about sauropod neck masses.  We started throwing around some numbers, derived from volumetric estimates and some off-the-cuff guessing. Rather than tell you more about it, I should just paste our conversation, minimally edited for clarity and with a few hopefully helpful links thrown in.

Mike: Dud. Neck masses.
Mike: Taylor (2009:803) measured the neck of Giraffatitan by GDI as 4117 liters.
Matt: k
Mike: I didn’t convert that to a mass, but I guess density of 0.5 is as good as any, which gives us (say) 2 tonnes.
Matt: That works for me.
Mike: That’s for an 8.5 m neck. So Supersaurus at 15 …
Matt: Yep. Almost twice as long, and not much more slender, and from what I’ve seen, ASP about the same.
Mike: Is 1.76 times as long. If it was isometric with the G. neck, it would be 5.5 times as heavy, which is 11 tonnes.
Matt: Oh.
Mike: So first: yeesh. Like, that is the mass of a whole freaking Diplo. Now we surely have to say isometry is unlikely.
Matt: Prolly.
Mike: But just multiplying out by length is unrealistic too. So maybe I should guess at mass =~ l^2? If I went with that, I’d get 6410 kg, which is elephant mass.
Matt: Something just occurred to me. Like, just now. For my 2006 poster, I calculated the mass of the cervical series in Giraffatitan, by summing over the CT slices from Brachiosaurus sp. BYU 12866 and multiplying by appropriate scale factors for the rest of the verts. We could “skin” that in muscle, and actually figure this out, for various muscle thicknesses, for one sauropod.
Mike: We should totally do that … if we had some idea how heavily muscled it was.
Matt: Well, obviously the thing to do is what Hutch et al. did for the tyrannosaurs, and put on several soft tissue envelopes. Crazy skinny, our best guess, markedly unfit, OMG, etc. It’s not that much more work. In fact, that could be my SVPCA talk this year.
Mike: Sure, but that’s just how to mitigate our ignorance. All we’d be doing at this point is taking n guesses instead of one. But, yeah, we should do it. Or you should if you prefer.
Matt: Let’s make it a Wedel and Taylor. I’ll crunch the numbers, but I want your input.
Mike: Works for me!
Matt: Good. Now let’s file it until April at least.

BYU 12613, a posterior cervical probably referable to Diplodocus, in dorsal (top), left lateral (left), and posterior (right) views. It most closely resembles C14 of D. carnegii CM 84/94 (Hatcher 1901: plate 3) despite being less than half as large, with a centrum length of 270 mm compared to 642 mm for C14 of D. carnegii. From Wedel and Taylor (in press).

Matt: Oh!
Matt: Also.
Matt: You know that little Diplo cervical from BYU that we figure in our in-press paper?
Mike: I think I know the one, yeah.
Matt: I am SUCH a moron. I have CT scans of the whole thing.
Mike: Good.
Matt: I forgot that Kent and I scanned it back in 2008. Even blogged about it, fer cryin’ out loud.  So I can do the sum-over-slices, scale-for-other-verts thing for Diplodocus, too. Which is at least closer to Supes than JANGO is.
Mike: Remind me, is it from a juvenile?
Matt: Maybe, maybe not. It IS tiny, but the neural spine is fused, the internal structure is crazy complex, and it doesn’t have any obvious juvenile characters other than just being small. The ASP is about as high as it gets in diplodocids. Which, as you may remember, is not nearly as high as it gets in titanosauriforms–that’s another paper that needs writing. Damn it. To know all this stuff and not have told it yet is killing me.
Mike: PeerJ!
Matt: I know!
Mike: Bottom line, it’s nuts that no-one has ever even tried to weigh a sauropod neck.* We should definitely do it, even if we do a really crappy job, if only so that others feel obliged to rebut.
Matt: Quite. Let’s do it. For reals.
Mike: In April. Done.

* R. McNeill Alexander (1985, 1989) did estimate the mass of the neck of Diplodocus, based on the old Invicta model and assuming a specific gravity of 1.0. Which was a start, and waaay better than no estimate at all. Still, let’s pretend that Mike meant “tried based on the actual fossils and what we know now about pneumaticity”.

The stuff about putting everything off until April is in there because we have a March 31 deadline to get a couple of major manuscripts submitted for an edited thingy. And we’ve made a pact to put off all other sciencing until we get those babies in. But I want to blog about this now, so I am.

Another thing Mike and I have been talking a lot about lately is the relation between blogging and paper-writing. The mode we’ve seen most often is to blog about something and then repurpose or rewrite the blog posts as a paper. Darren paved the way on this (at least in our scientific circle–people we don’t know probably did it earlier), with “Why azhdarchids were giant storks“, which became Witton and Naish (2008). Then last year our string of posts (starting here) on neural spine bifurcation in Morrison sauropods became the guts–and most of the muscles and skin, too–of our in-press paper on the same topic.

But there’s another way, which is to blog parts of the science as you’re doing them, which is what Mike was doing with Tutorial 20–that’s a piece of one of our papers due on March 31.

Along the way, we’ve talked about John Hawks’ model of using his blog as a place to keep his notes. We could, and should, do more of that, instead of mostly keeping our science out of the public eye until it’s ready to deploy (which I will always favor for certain projects, such as anything containing formal taxonomic acts).

And I’ve been thinking that maybe it’s time for me–for us–to take a step that others have already taken, and do the obvious thing. Which is not to write a series of blog posts and then decide later to turn it into a paper (I wasn’t certain that I’d be writing a paper on neural spine bifurcation until I had written the second post in that series), but to write the paper as a series of blog posts, deliberately and from the outset, and get community feedback along the way. And I think that the sauropod neck mass project is perfect for that.

Don’t expect this to become the most common topic of our posts, or even a frequent one. We still have to get those manuscripts done by the end of March, and we have no shortage of other projects waiting in the wings. And we’ll still post on goofy stuff, and on open access, and on sauropod stuff that has nothing to do with this–probably on that stuff a lot more often than on this. But every now and then there will be a post in this series, possibly written in my discretionary blogging time, that will hopefully move the paper along incrementally.

References

## The Recapture Creek sauropod: the reveal

### March 3, 2013

If you’re just joining us, this post is a follow-up to this one, in which I considered the possible size and identity of the Recapture Creek femur fragment, which “Dinosaur Jim” Jensen (1987: page 604) said was “the largest bone I have ever seen”.

True to his word, Brooks Britt at BYU got back to me with measurements of the Recapture Creek femur fragment in practically no time at all:

Length 1035 mm, width 665 mm.  However, you cannot trust the measurements because Jensen put a lot of plaster on the proximal half of the bone.

Now, taking plaster off a bone is not going to make it any larger. So the plastered-up specimen is the best case scenario for the RC femur to represent a gigapod. And I know the stated width of 665 mm is the max width of the proximal end, because I sent Brooks a diagram showing the measurements I was requesting. The length is a little less than anticipated, and doesn’t quite jibe with the max proximal width–I suspect a little might have broken off from the distal end where the preservation looks not-so-hot.

Based on those measurements, it looks like Jensen got the scale bar in Figure 8 in his 1987 paper approximately right–if anything, the scale bar is a little undersized, but only by 5% or so, which is actually pretty good as these things go (scale bars without measurements are still dag-nasty evil, though). By overlapping Jensen’s photo with the femur of the Brachiosaurus altithorax holotype (FMNH P25107) to estimate the size of the element when complete, I get a total length of 2.2 meters–exactly the same size as the Brachiosaurus holotype. If the Recapture Creek femur is from a Camarasaurus, which I don’t think we can rule out, it was 2 meters long when complete, or 11% longer and 37% more massive than the big C. supremus AMNH 5761–about 35 tonnes or maybe 40 on the outside. So it’s a big bone to be sure, but it doesn’t extend the known size range of Morrison sauropods.

So, as before, caveat estimator when working from scaled illustrations of single partial bones of possibly immense sauropods.

Now, here’s a weird thing. Let’s assume for the sake of this discussion that the Recapture Creek femur is from a brachiosaur. That gives us three individual Late Jurassic brachiosaurids–the Recapture Creek animal, the Brachiosaurus altithorax holotype, and the mounted Giraffatitan brancai–that are almost exactly the same size in limb bone dimensions (although B.a. had a longer torso). But we know that brachiosaurids got bigger, as evidenced by the XV2 specimen of Giraffatitan, and based on the lack of scapulocoracoid fusion in both FMNH P25107 and the mounted Giraffatitan. So why do we keep finding these (and smaller) subadults, and so few that were XV2-sized? I know that there gets to be a preservation bias against immense animals (it’s hard to bury a 50-tonne animal all in one go), but I would not think the 13% linear difference between these subadults and XV2-class adults would be enough to matter. Your thoughts?

Reference

Jensen, J.A. 1987. New brachiosaur material from the Late Jurassic of Utah and Colorado. Great Basin Naturalist 47(4): 592-608.

## How big was the Recapture Creek sauropod?

### February 28, 2013

From Jensen (1987, page 604):

“In 1985 I found the proximal third of an extremely large sauropod femur (Figs. 8A, 12A) in a uranium miner’s front yard in southern Utah.  The head of this femur is 1.67 m (5’6″) in circumference and was collected from the Recapture Creek Member of the the Morrison Formation in Utah near the Arizona border.  It is the largest bone I have ever seen.”

Jensen included not one but two figures of this immense shard of excellence. Here they are:

Jensen 1987, Figure 8

Jensen 1987, Figure 12

The specimen was heavily reconstructed, as you can see from the big wodge of unusually smooth and light-colored material in the photo. So we can’t put much stock in that part of the specimen.

Unfortunately, the only measurement of the specimen that Jensen gives in the paper is that circumference; there are no straight-line linear measurements, and the figures both have the dreaded scale bars. Why dreaded? Check this out:

As you can see, when the scale bars are set to the same size, the bones are way off (the scale bar in the drawing is 50 cm). This is not an uncommon problem. I make the Fig 8 version 30% bigger in max mediolateral width of the entire proximal end, and still 17% bigger in minimum diameter across the femoral head, as measured from the slight notch on the dorsal surface (on the right in this view).

Can we figure out which is more accurate based  on the internal evidence of the paper? For starters, the Fig 12 version is a drawing (1), that does not match the outline from the photo (2), and the hand-drawn scale bar (3) does not actually coincide with any landmarks (4), and that’s plenty of reasons for me not to trust it.

What about that circumference Jensen mentioned? Unfortunately, he didn’t say exactly where he took it, just that the head of the femur had a circumference of 1.67 meters. Is that for the entire proximal end, or for the anatomical head that fits in the acetabulum, er wot? I’m afraid the one measurement given in the paper is no help in determining which of the figures is more accurately scaled.

The obvious thing to do would be to see if this bone is in the BYU collections, and just measure the damn thing. More on that at the end of the post.

In the meantime, Jensen said that the shape of the Recapture Creek femur was most similar to the femur of Alamosaurus, or to that of Brachiosaurus among Morrison taxa, and he referred it to Brachiosauridae. So how does this thing–in either version–compare with the complete femur of FMNH P25107, the holotype of Brachiosaurus altithorax?

The Recapture Creek femur fragment compared to the complete femur of the Brachiosaurus altithorax holotype FMNH P25107

The first thing to notice is that the drawn outline from Figure 12 is a much better match for the Brachiosaurus altithorax femur–enough so that I wonder if Jensen drew it from the Recapture Creek specimen, or just traced the B.a. proximal femur and scaled it accordingly (or maybe not accordingly, since the scale bars don’t match).

But let’s get down to business: how long would the complete femur have been?

Using the scale bar in the photograph from Figure 8 (on the left in above image), I get a total femur length of 2.36 meters. Which is long, but only 7.7% longer than the 2.19-meter femur of FMNH P25107, and therefore only 25% more massive. So, 35 tonnes to Mike’s 28-tonne B.a., or maybe 45 tonnes to a more liberal 36-tonne B.a. Big, yeah, but not world-shattering.

Using the scale bar in the drawing from Figure 12 (on the right in the above image)–which, remember, is 50 cm, not 1 meter–I get a total femur length of about 1.9 meters, which is considerably smaller than the B.a. holotype. That is very much at odds with Jensen’s description of it as “the largest bone I have ever seen”, and given that we have many reasons for not trusting the scale bar in the drawing, it is tempting to just throw it out as erroneous.

So it would seem that unless Jensen got both scale bars too big, the Recapture Creek brachiosaur was at most only a shade bigger than the holotype specimen of Brachiosaurus altithorax.

But wait–is the Recapture Creek brachiosaur a brachiosaur at all? Jensen didn’t list any characters that pushed him toward a brachiosaurid ID, and I don’t know of any proximal femur characters preserved in the specimen that would separate Brachiosaurus from, say, Camarasaurus. And in fact a camarasaur ID has a lot to recommend it, in that Camarasaurus femora have very offset heads (the ball- or cylinder-like articular surface at the top end sticks out a big more to engage with the hip socket–see Figure 12 up near the top of the post), moreso than in many other Morrison sauropods, and that would make them better matches for the Recapture Creek femur photo. Here’s what the comparo looks like:

The Recapture Creek femur fragment compared with a complete femur of Camarasaurus.

I make that a 2.07-meter femur using the photo on the left, and a 1.66-meter femur using the drawing on the right. The one decent femur in the AMNH 5761 Camarasaurus supremus collection is 1.8 meters long, so these results are surprisingly similar to those for the B. althithorax comparison–the drawing gives a femur length shorter than the largest known specimens, and the photo gives a length only slightly longer. A camarasaur with a 2.07 meter femur would be 15% larger than the AMNH C. supremus in linear terms, and  assuming isometric scaling, 1.5 times as massive–maybe 38 tonnes to AMNH 5761′s estimated 25. A big sauropod to be sure, but not as big as the largest apatosaurs, and not nearly as big as the largest titanosaurs.

I have always been surprised that the Recapture Creek femur frag has attracted so little attention, given that “Dinosaur Jim” himself called it the biggest bone he had ever seen. But it appears that the lack of attention is justified–whether it was a brachiosaur or a camarasaur, and using the most liberal estimates the scale bars allow, it simply wasn’t that big.

Update about half an hour later: Okay, maybe I was a little harsh here. IF the photo scale bar is right, the Recapture Creek femur might still represent the largest and most massive macronarian from the Morrison Formation (Edit: only if it’s a brachiosaur and not a camarasaur; see this comment), which is something. I suppose I was particularly underwhelmed because I was expecting something up in OMNH 1670-to-Argentinosaurus territory, and so far, this ain’t it. I’ll be interested to see what the actual measurements say (read on).

The Moral of This Story

So, if it wasn’t that big after all, and if no-one has made a stink about it being big before now, why go to all this trouble? Well, mostly just to satisfy my own curiosity. If there was a truly gigantic brachiosaur from the Morrison, it would be relevant to my interests, and it was past time I crunched the numbers to find out.

But along the way something occurred to me: this should be a cautionary tale for anyone who gets all wound up about the possible max size of Amphicoelias fragillimus. As with A. fragillimus, for the Recapture Creek critter we have part of one bone, and at least for this exercise I was working only from published illustrations with scale bars. And as with A. fragillimus, the choice of a reference taxon is not obvious, and the size estimates are all over the place, and some of them just aren’t that big.

It always amuses me when A. fragillimus comes up and people (well, trolls) accuse us of being big ole’ wet blankets that just don’t want to believe in 200-tonne sauropods. It amuses me because it’s wrong on so many levels. Believe me, when we have our sauropod fanboy hats on, we most definitely do want to believe in 200-tonne sauropods. That would rock. But when we put our scientist hats on, wanting and belief go right out the window. We have to take a cold, hard look at the data, and especially at its limitations.

Oh, the other moral is to go buy a tape measure, and use it. Sheesh!

Coda

As I said above, the obvious thing to do would be to just track down the bone and measure it. It does still exist, it’s in the BYU collections, and Brooks Britt has kindly offered to send along some measurements when he gets time. So we should have some real answers before long (and here they are). But I wanted to work through this example without them, to illustrate how much uncertainty creeps in when trying to estimate the size of a big sauropod from published images of a single partial bone.

Reference

Jensen, J.A. 1987. What I did on my holidaysNew brachiosaur material from the Late Jurassic of Utah and Colorado. Great Basin Naturalist 47(4): 592-608.

## …and that, ladies and gentlemen, is why you code Brachiosaurus and Giraffatitan separately

### January 23, 2013

D’Emic (2012: figure 5)

Now this is super-freakin’ cool, and I’ve been meaning to blog about it for a while. In Mike D’Emic’s recent titanosauriform phylogeny (D’Emic 2012), he (correctly) included Brachiosaurus and Giraffatitan as separate OTUs, and, hey, whaddayaknow, they’re not sister taxa anymore: Brachiosaurus is more closely related to a trio of Early Cretaceous North American brachiosaurids than it is to Giraffatitan.

The potential for someone to find this result was there ever since Mike broke Brachiosaurus and Giraffatitan apart, as a previously composite OTU, in his 2009 paper. It just hadn’t materialized. In fact, some authors have gone out of their way to not find this out, by keeping the old composite coding. That seems…unwise, in retrospect. Whether one agreed with Mike on the nomenclatural point of generic separation or not, not coding the two taxa as separate OTUs (especially after Mike had done that work for them) was a poor phylogenetic decision–in essence, it constrained Brachiosaurus and Giraffatitan to be sister taxa in the analysis, and outlawed any more interesting results–like the one obtained by D’Emic (2012)–before the software even started crunching trees.

So anyway, back to the coolness inherent in D’Emic’s tree. Of course, like all phylogenetic results this is just a hypothesis and it is subject to revision based on new information blah blah blah…but it is really interesting that there is now some phylogenetic support for an endemic radiation of brachiosaurids in North America (bonus goofy observation–you can’t spell ‘endemic’ without D’Emic). Or perhaps Lauriasia–I would kill to know where the British brachiosaurids (or basal titanosauriforms) fit into this story, and Lusotitan, and the apparently tiny Croatian carbonate platform brachiosaurs.

Also super-interesting that, if this tree is accurate, these endemic Early Cretaceous brachiosaurids were living alongside a giant basal somphospondyl in the form of Sauroposeidon, which came from heaven knows where. Look who it’s surrounded by–Ligabuesaurus is from Argentina, Tastavinsaurus is from Spain, and the euhelopodids are from eastern Asia. Evidently there was also a global radiation of basal somphospondyls. And why are all the Early Cretaceous North American brachiosaurids small–smaller than Brachiosaurus and Giraffatitan, anyway (at least until we find bigger individuals of the former)–while Sauroposeidon is so big? Or is that just an effect of tiny sample sizes, and one lucky strike in the form of the Sauroposeidon holotype?

So much cool stuff to think about. I don’t usually get this much enjoyment out of a tree unless it has lights and ornaments.

## 2000 AD’s flagrantly plagiarised Brontosaurus

### October 18, 2012

Another blast from the past:

Like the recent Compsognathus, this is a card from the “Flesh” card-game that was printed across several progs (issues) of the comic 2000 AD in 1977. This one is from the back cover of Prog 10. (Click through the picture for the whole back cover.)

What’s interesting about this one is how very flagrant a rip-off it is of Rudolph Zallinger’s 1960 painting of Brontosaurus being attacked by Allosaurus:

I know this painting best from Dinosaurs and other Prehistoric Reptiles, a 1966 book that I had as a boy, and which I believe is the same thing as the Giant Golden Book of Dinosaurs. Here is a high-resolution scan of my copy of that book, pages 24-25. (Click through for 5472 by 3669 version.)

And while I’m here, I may as well throw in my scan of the “Brachiosaurus” (i.e. Giraffatitan)on pages 20-21. (Click through for 5431 by 3162 version.)

I will leave it to others to point out which other classic piece of sauropod art this one plagiarises.

## The sacrum of Brachiosaurus

### May 12, 2012

In my 2009 brachiosaur paper, I gave rather short shrift to the sacrum of Brachiosaurus — in part because there is no really good sacrum of Giraffatitan to compare it to. Also my own photos of the sacrum, taken back before I figured out how to photograph big bones, are all pretty terrible.

Happily, Phil Mannion took some much better photos and gave us permission to use them. So I prepared this multi-view figure, which we plan to use in a forthcoming paper about another sacrum:

At the bottom, we have the sacrum in left lateral view; above it, in dorsal view. To the left is the anterior view (with dorsal to the right) and the right is the posterior view (with dorsal to the left). The idea of this composition is that you could print the image out, and cut and fold it into a cuboid.  (In fact I might just do that.)

As usual with these things, click through for the much more impressive full-resolution version (3809 x 3157 pixels).

## MOLD-A-RAMA

### May 3, 2012

I’m in Chicago, visiting the Field Museum, which means two things: Brachiosaurus (see below), and Mold-A-Rama. Downstairs from the great hall, on the ground floor, they have Mold-A-Rama machines, and I cannot resist their siren song.

The Mold-A-Rama is the king of novelty souvenirs. You can keep your stamped pennies, little pewter spoons, hand-painted bells, and refrigerator magnets. None of them is worthy to sully the presence of the awesome Mold-A-Rama. You put in two dollars, and then you get to watch as this hissing, clanking 1950s machine with tubes and wires and lights actually makes your item right in front of your eyes. A few minutes later, BAM, you’re holding a hot, smelly hunk of probably carcinogenic plastic that is so fresh from the mold that it is still a bit soft. You can’t buy that kind of authenticity–except from a Mold-A-Rama.

See that red thing, newly made, still in the bowels of the machine? That is MY T. rex!

This is my first Mold-A-Rama Triceratops. I already have a T. rex from my last visit, way back in 2005, which I can now pass on to my son. I also have a Stegosaurus, a Brontosaurus (shown but not commented on here), and a Trachodon. Yeah, yeah, I know the real animals are known as Apatosaurus and Edmontosaurus these days, but I’m not talking about the real animals. I’m talking about Mold-A-Rama, and trust me, the Mold-A-Rama critters are Brontosaurus and Trachodon. They drag their tails, they live in swamps, they’re cold-blooded and they died out from racial senescence (in about 1975, I think).

Finally, because Mike will straight-up murder me if I post from Chicago without it, here’s your friendly local not-quite-fully-mature mounted holotype specimen of Brachiosaurus:

## Neural spine bifurcation in sauropods, Part 6: more reasons why Haplocanthosaurus is not a juvenile of a known diplodocid

### April 15, 2012

Last time, we saw why Haplocanthosaurus couldn’t be a juvenile of Apatosaurus or Diplodocus, based on osteology alone.  But there’s more:

# Ontogenetic status of Haplocanthosaurus

Here is where is gets really surreal.  Woodruff and Fowler (2012) blithely assume that Haplocanthosaurus is a juvenile of something, but the type specimen of the type species — H. priscus CM 572 — is an adult.  As Hatcher (1903:3) explains:

The type No. 572 of the present genus consists of the two posterior cervicals, ten dorsals, five sacrals, nineteen caudals, both ilia, ischia and pubes, two chevrons, a femur and a nearly complete series of ribs, all in an excellent state of preservation and pertaining to an individual fully adult as is shown by the coössified neural spines and centra.

So far as I can see, Woodruff and Fowler are confused because the second species that Hatcher describes, H. utterbacki, is based on the subadult specimen CM 879.  Where possible in the previous post, I have used illustrations of the adult H. priscus, so that the comparisons are of adult with adult.  The exceptions are the two anterior cervicals and the first dorsal, which are known only from H. utterbacki.  And sure enough, if you look closely at the illustrations, you can see that in these vertebrae and only these vertebrae, Hatcher had the neurocentral junction illustrated — because it wasn’t yet fused.

Haplocanthosaurus posterior, mid and anterior cervical vertebrae, C14, C9 and C4, in right lateral view. C14 of adult H. priscus (from Hatcher 1903:plate I); C9 and C4 of H. utterbacki (from plate II). Red ellipses highlight neurocentral sutures.

As it happens, the difference in ontogenetic status between these two specimens is nicely illustrated by Wedel (2009), although he was only in it for the pneumaticity:

Neurocentral fusion in Haplocanthosaurus. A, B. Posterior cervical vertebra C?12 of sub-adult H. utterbacki holotype CM 879: A, X-ray in right lateral view; B, coronal CT slice showing separate ossificaton of centrum and neural arch. C, D. Mid-dorsal vertebra D6 of adult H. priscus holotype CM 572: X-rays in (A) right lateral and (B) anterior view, showing fully fused neural arch. Wedel (2009:fig. 6)

So H. utterbacki CM 879 certainly is an immature form of something; and that something is Haplocanthosaurus, most likely H. priscus.  (The characters which Hatcher used to separate the two species are not particularly convincing.)

With that out the way, we can move on to …

# Phylogenetic analysis

A simple way to evaluate the parsimony or otherwise of a synonymy is to use a phylogenetic analysis. In their abstract, Woodruff and Fowler claim that “On the basis of shallow bifurcation of its cervical and dorsal neural spines, the small diplodocid Suuwassea is more parsimoniously interpreted as an immature specimen of an already recognized diplodocid taxon”.  Without getting into the subject of Suuwassea again — Matt pretty much wrapped that up in part 4 — the point here is that the word “parsimony” has a particular meaning in studies of evolution: it refers to minimising the number of character-state changes.  And we have tools for measuring those.

So let’s use parsimony to evaluate the hypothesis that Haplocanthosaurus is one of the previously known diplodocids.  Pretending for the moment that Haplocanthosaurus really was known only from subadults, how many additional steps would we need to account for if ontogeny were to change its position to make it group with one of the diplodocids?

You don’t need to be a cladistics wizard to do this.  (Which is handy, since I am not one.)  Here’s the method:

• Start with an existing matrix, add constraints, re-run it, and see how the tree-length changes.  Since I am familiar with it, I started with the matrix from my 2009 paper on brachiosaurs.
• Re-run the matrix to verify that you get the same result as in the published paper based on it.  This gives you confidence that you’re running it right.  In this case, I got a minimum tree length of 791 steps, just as in Taylor (2009).
• Add extra instructions to the run-script defining and imposing constraints.  Note that you do not have to mess with the characters, taxa or codings to do this.
• Run the matrix again, with the constraint in place, and see how the tree-length changes.
• Repeat as needed with other constraints to evaluate other phylogenetric hypotheses.

(This is how we produced the part of the Brontomerus paper (Taylor et al. 2011:89) where we said “One further step is sufficient to place Brontomerus as a brachiosaurid, a basal (non−camarasauromorph) macronarian, a basal (non−diplodocid) diplodocoid or even a non−neosauropod. Three further steps are required for Brontomerus to be recovered as a saltasaurid, specifically an opisthocoelicaudiine”.  And that’s why we weren’t at all dogmatic about its position.)

Anyway, going through this exercise with Haplocanthosaurus constrained in turn to be the sister taxon to Apatosaurus, Diplodocus, etc., yielded the following results:

• (no constraint) –  791 steps
• Apatosaurus — 817 (26 extra)
• Diplodocus — 825 (34 extra)
• Barosaurus — 815 (24 extra)
• Camarasaurus — 793 (2 extra)
• Brachiosaurus — 797 (6 extra)

(I threw in the other well-known Morrisson-Formation sauropods Camarasaurus and Brachiosaurus, even though Woodruff and Fowler don’t mention them, just because it was easy to do and I was interested to see what would happen.  And when I say Brachiosaurus, I mean B. altithorax, not Giraffatitan.)

I hope you’re as shocked as I am to see that for Haplocanthosaurus to emerge as the sister taxon of any diplodocid needs a minimum of 24 additional steps — or an incredible 34 for it to be sister to Diplodocus.  In other words, the hypothesis is grossly unparsimonious.  Of course, that doesn’t in itself mean that it’s false: but it does render it an extraordinary claim, which means that it needs extraordinary evidence.  And while “the simple spines of Haplocanthosaurus might bifurcate when it grows up” is extraordinary evidence, it’s not in the way that Carl Sagan meant it.

In short, running this simple exercise — it took me about a hour, mostly to remember how to do constraints in PAUP* — would have given Woodruff and Fowler pause for thought before dragging Haplocanthosaurus into their paper.

Oh, and it’s ironic that placing Haplo as sister to Brachiosaurus requires only a quarter as many steps as the closest diplodocid, and as sister to Camarasaurus requires only two steps.  If you really want to synonymise Haplocanthosaurus, Camarasaurus is the place to start.  (But don’t get excited, it’s not Camarasaurus either.  It’s Haplocanthosaurus.)

[By the way, anyone who'd like to replicate this experiment for themselves is welcome: all the files are available on my web-site.  You only really need the .nex file, which you can feed to PAUP*, but I threw in the log-file, the generated tree files and the summary file, too.  Extra Credit: run this same exercise to evaluate the parsimony of Suuwassea as a subadult of one of these other genera.  Report back here when you're done to earn SV-POW! points.]

# Conclusion

It’s a truism that we stand on the shoulders of giants.  In the case of sauropod studies, those giants are people like J. B. Hatcher, Charles Gilmore, Osborn and Mook and — bringing it up to date — John McIntosh, Paul Upchurch, Jeff Wilson and Jerry Harris.  When Hatcher described Haplocanthosaurus as a new genus rather than a subadult Diplodocus, he wasn’t naive.  He recognised the effects of ontogeny, and he was aware that one of his two specimens was adult and the other subadult.  He was also probably more familiar with Diplodocus osteology than anyone else has ever been before or since, having written the definitive monograph on that animal just two years previously (Hatcher 1901).

By the same token, people like Upchurch and Wilson have done us all a huge favour by making the hard yards in sauropod phylogenetics.  If we’re going to go challenging the standard consensus phylogeny, it’s just good sense to go back to their work (or the more recent work of others, such as Whitlock 2011), re-run the analyses with our pet hypotheses encoded as constraints, and see what they tell us.

So in the end, my point is this: let’s not waste our giants.  Let’s take the time to get up on their shoulders and survey the landscape from up there, rather than staying down at ground level and seeing how high we can jump from a standing start.

## The rest of the series

Links to all of the posts in this series:

and the post that started it all:

## References

• Hatcher, J.B. 1901. Diplodocus (Marsh): its osteology, taxonomy, and probable habits, with a restoration of the skeleton. Memoirs of the Carnegie Museum 1:1-63.
• Hatcher, J.B. 1903. Osteology of Haplocanthosaurus with description of a new species, and remarks on the probable habits of the Sauropoda and the age and origin of the Atlantosaurus beds; additional remarks on Diplodocus. Memoirs of the Carnegie Museum 2:1-75.
• Taylor, M.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.
• Taylor, M.P., Wedel, M.J. and Cifelli, R.L. 2011. A new sauropod dinosaur from the Lower Cretaceous Cedar Mountain Formation, Utah, USA. Acta Palaeontologica Polonica 56(1):75-98. doi:10.4202/app.2010.0073
• Wedel, M.J. 2009. Evidence for bird-like air sacs in saurischian dinosaurs. Journal of Experimental Zoology 311A:611-628.
• Whitlock, J.A. 2011. A phylogenetic analysis of Diplodocoidea (Saurischia: Sauropoda). Zoological Journal of the Linnean Society 161(4):872-915. doi: 10.1111/j.1096-3642.2010.00665.x
• Woodruff, D.C, and Fowler, D.W. 2012. Ontogenetic influence on neural spine bifurcation in Diplodocoidea (Dinosauria: Sauropoda): a critical phylogenetic character. Journal of Morphology, online ahead of print.

## How big was Brachiosaurus‘s forelimb?

### June 1, 2011

I recently happened upon a picture of the late Jim Jenson standing beside the huge front leg of “Ultrasauros”, which leads me to ask a few questions. Did he really find a complete forelimb? Was the leg from Brachiosaurus altithorax? If that leg is valid at actual size how tall/long was the whole animal? It looks to be about 40% to 50% taller than the berlin Giraffatitan, I am guessing the leg is a constructed representation of how the leg would look rather than a cast of the actual leg because if the whole front leg was found they would probably be the most talked about sauropod bones in the world and the fact is I’ve heard pretty much nothing about these remains for years.

I answered this in a followup comment, but because the answer involved a few nice images, I thought it ought to be promoted into a post of its own.  So here it is, in expanded form.

I believe I know the picture Peter was talking about: it was either the one on the right, of Jensen working on the limb in the lab, or the one below of the same limb, again with Jensen, this time out in the desert.

As an aside: based on a post by ReBecca Hunt-Foster (scroll down to the 12th picture), it looks like this forelimb may have ended up in the New Mexico Museum of Natural History and Stuff (NMMNHS).

Anyway, the bad news is that, no, this is not a complete forelimb fossil. The worse news is that the limb is not even partly cast from real material: it’s a pure sculpture, based presumably on the forelimb of Giraffatitan brancai, but scaled up according to Jensen’s idea of how big “Ultrasauros” was. The only part of the model that probably was cast from real material is not part of the limb proper, but the scapulocoracoid — which is the only real brachiosaur element that Jensen found and described from the Dry Mesa quarry.  In fact, the scap in these photos (and in ReBecca’s) does look very much like BYU 9462, the element that Jensen meant to designate as the “Ultrasauros” holotype, but didn’t, instead plumping for … ah, you all know the story.

"Ultrasauros" scapulocoracoid BYU 9462 (almost certainly a cast), with Graeme Elliott for scale.

But in fact, the scapulocoracoid in the whole-forelimb pictures above looks much too small in comparison with the other elements; or to put it the right way round, since only the scap is based on an actual fossil, all the other elements are too big — which suggests that Jensen exaggerated the sizes of the sculpted limb bones well beyond what the scapulocoracoid warranted.  (In any case, the idea that this scap represents a much larger brachiosaurid than any previously known specimen was shown by Curtice et al. (1996) to be mistaken — it’s from an animal pretty similar in size to, and probably a little smaller than, the largest known Tendaguru specimens.)

But the good news is, Peter’s sense of awe is not misplaced. Real brachiosaur forelimbs are actually not much less impressive than this. See for example me next to the right forelimb of the Berlin Giraffatitan mount, which is real bone — as shown in our classic post Shedloads Of Awesome:

Or here I am again, this time with the Chicago Brachiosaurus mount. (The Chicago mount is a cast, based on a hybrid of real Brachiosaurus elements, some bits of Giraffatitan, and some sculptures, but the scaling is good.)

My rule of thumb, based on a lot of posing for photos around the Chicago mount, is that if I stand next to the forelimb and reach up, I can just rest my hand on the top of the ulna without stretching.  I’m about six feet tall, if that helps.

Jim Jensen was 4% taller than me — 6’3″.  Bearing that in mind, and looking at the second photograph above (the first one is useless because of the forced perspective), Jensen’s inability to reach close to the top of the ulna suggests that his model is inflated by maybe 30%.  Which means that it represents an animal about 1.3^3 = 2.2 times as voluminous and heavy as it should be.  But let’s not forget that among the Giraffatitan material in Berlin is the isolated fibula XV 2, which at 134 cm in length is 12.6% longer than the 119 cm tibia of S II.  So that is from animal about half way between S II and Jensen’s Imaginary Monster in size.

So.  Real brachiosaurs are awesome enough.

## Early Brachiosaurus art

### April 8, 2010

Most people think of Janensch’s (1950b) plate VIII as being the first skeletal reconstruction of “Brachiosaurus” (although Janensch’s species “Brachiosaurusbrancai is now referred to the separate genus Giraffatitan).  And it certainly is a classic:

"Brachiosaurus" brancai (i.e. Giraffatitan) classic skeletal reconstruction (Janensch 1950b: plate VIII)

But the reconstruction published in 1950 is modelled on the physical mount of the specimen HMN SII, which not only was constructed much earlier, but was even published as a photograph in Janensch’s (1938) earlier paper on the mass of his species.  Comparing the drawing (above) with the photograph (below), it’s easy to see how closely they resemble each other, not only in proportions but in pose:

"Brachiosaurus" brancai (i.e. Giraffatitan) mounted skeleton in the Humboldt Museum fur Naturkunde, Berlin; photograph from ﻿﻿Janensch (1938: fig 1)

Yet less well known is that when the mount was completed, shortly before the start of World War II, it was unveiled against a backdrop of Nazi banners.  I have not been able to find a photograph of this (and if anyone has one, please do let me know), but I do have this drawing of the event, taken from an Italian magazine and dated 23rd December 1937.  Since the date stamp is marked “Zoolog. Museum Berlin”, I guess that is the date that a museum artist executed the drawing, or maybe when a copy was released by the museum to the magazine.  Once again, I don’t know, and would welcome clarification.  Anyway, here it is:

So we have a published photograph and a published drawing of a brachiosaur skeleton that predate Janensch’s (1950b) reconstruction, but was there an earlier actual skeletal reconstruction?  Indeed there was: Matthew’s (1915) popular book included what I believe was the first ever brachiosaur reconstruction, and here it is:

Composite Brachiosaurus skeletal reconstruction, from Matthew (1915: fig. 24)

Matthew’s caption to this figure says that it is “from specimens in the Field Museum in Chicago and the Natural History Museum in Berlin”, i.e. it incorporates elements from both Brachiosaurus proper (B. altithorax) and the Tanzanian species “Brachiosaurusbrancai.  And if you’re familiar with the fossils in question, that’s evidently the case: for example, the scapula is clearly based on HMN Sa 9, and the posterior dorsals are unmistakably those of FMNH P25107.  [The inclusion of those dorsals fulfils our weekly sauropod-vertebra picture mandate, in case you were wondering.]

This is pretty impressive work, especially given that it was published one year after Janensch’s (1914) preliminary short paper on the Tendaguru Formation’s fossils.  Since that initial report did not figure the scapula Sa 9, it’s tempting to imagine that Matthew or his artist must have visited Berlin and seen the material in person; but as this was in the middle of World War I, that seems unlikely.  Does anyone know the story here?

And finally, we come to what is probably the first life restoration of Brachiosaurus or any brachiosaur.  It’s the work of Abel, and I found it in Young (1975: page 4):

Abel's restoration of Brachiosaurus, undated, from Young (1975) page 4

Infuriatingly, Young does not say anything whatsoever about the provenance of this restoration — for all I know, it might have been done in 1974 by a talentless artist who ignored the previous sixty years’ work.  But it seems more likely that it’s very early work, and therefore of great historical importance.  Once more (and believe me, I am getting embarrassed at how often I’ve said this), I welcome any further information.

### And in other news …

Many of you will have used PDFs downloaded from the O. C. Marsh archive at http://sauroposeidon.net/marsh.html.  That address will become inoperative at the end of this month, and the archive is now hosted at http://marsh.dinodb.com/ – Please update your bookmarks, links, etc.

## References

• 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), pp. 81-110.
• Janensch, Werner.  1938.  Gestalt und Grösse von Brachiosaurus und anderen riesenwüchsigen Sauropoden.  Biologe 7: 130-134, 2 figs.
• Janensch, Werner.  1950b.  Die Skelettrekonstruktion von Brachiosaurus brancai.  Palaeontographica (Suppl. 7) 3: 97-103.
• Matthew, W. D.  1915.  Dinosaurs, with special reference to the American Museum collections.  American Museum of Natural History, New York.  164 pages.
• Young, D.  1975.  Brachiosaurus, the biggest dinosaur of them all. Field Museum of Natural History Bulletin 46(1):3-9.