We jumped the gun a bit in asking How fat was Camarasaurus? a couple of years ago, or indeed How fat was Brontosaurus? last year. As always, we should have started with extant taxa, to get a sense of how to relate bones to live animals — as we did with neck posture.

So here we go. I give you a herd of Indian elephants, Elephas maximus (from here):

056-Elephant-c-Gehan-de-Silva-Wijeyeratne-Minneriya-2004-07-27-165-Gathering

You will notice, from this conveniently-close-to-anterior view, that their torsos bulge out sideways, much further than the limbs.

Now let’s take a look at the skeleton of the same animal in the Oxford University Museum of Natural History (downloaded from here but for some reason the photo has now gone away):

4795746797_575d1f0ce6_b

The rib-cage is tiny. It doesn’t even extend as far laterally as the position of the limb bones.

(And lest you think this is an oddity, do go and look at any mounted elephant skeleton of your choice, Indian or African. They’re all like this.)

What’s going on here?

Is Oxford’s elephant skeleton mounted incorrectly? More to the point, are all museums mounting their elephants incorrectly? Do elephants’ ribs project much more laterally in life?

Do elephants have a lot of body mass superficial to the rib-cage? If so, what is that mass? It’s hard to imagine they need a huge amount of muscle mass there, and it can’t be guts. Photos like this one, from the RVC’s televised elephant dissection on Inside Nature’s Giants, suggest the ribs are very close to the body surface:

01020162576700

I’m really not sure how to account for the discrepancy.

Were sauropods similarly much fatter than their mounted skeletons suggest? Either because we’re mounting their skeletons wrongly with the ribs too vertical, or because they had a lot of superficial body mass?

Consider this mounted Camarasaurus skeleton in the Dinosaur Hall at the Arizona Museum of Natural History (photo by N. Neenan Photography, CC-BY-SA):

Camarasaurus_skeleton

Compare the breadth of its ribcage with that of the elephant above, and then think about how much body bulk should be added.

This should encourage palaeoartists involved in the All Yesterdays movement to dramatically bulk up at least some of their sauropod restorations.

It should also make us think twice about our mass estimates.

OMNH 1331 is my new hero

March 24, 2013

Here’s an update from the road–get ready for some crappy raw images, because that’s all I have the time or energy to post (with one exception).

OMNH 1331

Here’s OMNH 1331. It’s just the slightly convex articular end off a big vertebra, collected near Kenton, Oklahoma, in 1930s by one of J. Willis Stovall’s field crews. I measured the preserved width at 45 cm using a tape measure, and at 44.5 in GIMP using the scale bar in the photo, which is up on a piece of styrofoam so it’s about the same distance from the camera as the rim of the vertebra (i.e, about 8 feet–as high as I could get and still shoot straight down). So whether your distrust runs to tape measures or scale bars in photos, I am prepared to argue that this sucker is roughly 45 cm wide.

OMNH 1331 internal structure

There’s admittedly not a ton of morphology here, but the size and the fact that the other side is hollow and has a midline bony septum show that it is a pneumatic vertebra from a sauropod, and given that the quarry it’s from was chock-full of Apatosaurus, and liberally salted with gigantic Apatosaurus, I feel pretty good about calling it Apatosaurus.

OMNH 1331 cloned and flipped

To figure out how wide the articular face was when it was intact, I duplicated the image and reversed it left-to-right in GIMP, which yields an intact max width of about 49 cm. That is friggin’ immense.

If we make the maximally conservative assumption that this is the largest centrum in the whole skeleton of a big Apatosaurus, then it has to be part of a dorsal vertebra. Here are the max diameters of the largest dorsal centra in some big mounted apatosaurs, taken from Gilmore (1936). The number in parentheses is how many percent bigger OMNH 1331 is.

  • A. louisae CM 3018 – 36.5 cm (34%)
  • A. parvus UWGM 15556 – 36.5 cm (34%)
  • A. sp. FMNH P25112 – 41 cm (20%)

OMNH 1331 lateral view

However, this might not be part of a dorsal vertebra. For one thing, it’s pretty convex, and Apatosaurus dorsals sometimes have a little bump but they’re pretty close to amphiplatyan, at least in the posterior half of the series. For another, I think that smooth lower margin on the right in the photo above is part of the rim of a big pneumatic foramen, but it’s waaay up high and pretty medial on the centrum, opening more dorsally than laterally, which I have seen a lot in anterior caudal vertebrae. Finally, Jack McIntosh went through the OMNH collections years ago and his identifications formed the basis for a lot of the catalogue IDs, and this thing is catalogued as the condyle off the back end of a proximal caudal.

Here are the max diameters of the largest caudal centra in those same mounted apatosaurs, again taken from Gilmore (1936). Once again, the number in parentheses is how many percent bigger OMNH 1331 is.

  • A. louisae CM 3018 – 30 cm (63%)
  • A. parvus UWGM 15556 – 32.5 cm (51%)
  • A. sp. FMNH P25112 – 39 cm (26%)

(Aside: check out the skinny rear end on A. louisae. ‘Sup with that?)

So whatever vert it’s part of, OMNH 1331 is damn big bone from a damn big Apatosaurus. There are lots of other big Apatosaurus vertebrae in the OMNH collections, like OMNH 1670, but OMNH 1331 is the largest centrum that I know of in this museum. Which is why you’re getting a post about most of one end of a centrum in the wee hours of the morning–it’s most of one end of an awesome centrum. And it pains me when people do comparison figures of big sauropod vertebrae, or lists of the “Top 10 Largest Sauropods”, and put in stuff like Argentinosaurus and Puertasaurus and Supersaurus, but leave out Apatosaurus. It was legitimately huge, and it’s time the world realized that.

For more on the giant Oklahoma Apatosaurus, see:

Reference

Gilmore, C.W. 1936. Osteology of Apatosaurus with special reference to specimens in the Carnegie Museum. Memoirs of the Carnegie Museum 11:175-300.

supersaurus-vs-giraffe

At the top: our old friend BYU 9024 — the cervical vertebra that’s part of the Supersaurus vivianae holotype. At the bottom, C2 (the longest cervical) of Giraffa camelopardalis angolensis FMNH 34426.

The Supersaurus vertebra is 138 cm long. We don’t know which cervical it is, but there’s no reason to think it’s the longest. The giraffe vertebra is 31 cm long. Not only is the Supersaurus vertebra four times as long as that of the giraffe, it’s one of more than twice as many cervicals as the giraffe has.

Did we cheat by using an unusually small giraffe? Not really. When we articulated all seven cervicals as best we could, the sequence measured 171 cm, which is a fairly healthy 71% of the 2.4 m neck of the world-record giraffe. It’s not a monster, but it’s a decent-sized adult.

Bottom line, giraffes are just lame.

Recapture Creek comparo with measurements

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.

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 8

Jensen 1987, Figure 12

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:

Recapture Creek figs 8 and 12 comparedAs 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 brachiosaur femur fragment compared to the complete femur of the Brachiosaurus altithorax holotype FMNH P25107

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:

Recapture Creek - Camarasaurus comparo

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.

Why giraffes have short necks

September 26, 2012

Today sees the publication, on arXiv (more on that choice in a separate post), of Mike and Matt’s new paper on sauropod neck anatomy. In this paper, we try to figure out why it is that sauropods evolved necks six times longer than that of the world-record giraffe — as shown in Figure 3 from the paper (with a small version of Figure 1 included as a cameo to the same scale):

Figure 3. Necks of long-necked sauropods, to the same scale. Diplodocus, modified from elements in Hatcher (1901, plate 3), represents a “typical” long-necked sauropod, familiar from many mounted skeletons in museums. Puertasaurus modified from Wedel (2007a, figure 4-1). Sauroposeidon scaled from Brachiosaurus artwork by Dmitry Bogdanov, via commons.wikimedia.org (CC-BY-SA). Mamenchisaurus modified from Young and Zhao (1972, figure 4). Supersaurus scaled from Diplodocus, as above. Alternating pink and blue bars are one meter in width. Inset shows Figure 1 to the same scale.

This paper started life as a late-night discussion over a couple of beers, while Matt was over in England for SVPCA back in (I think) 2008. It was originally going to be a short note in PaleoBios, just noting some of the oddities of sauropod cervical architecture — such as the way that cervical ribs, ventral to the centra, elongate posteriorly but their dorsal counterparts the epipophyses do not.

As so often, the tale grew in the telling, so that a paper we’d initially imagined as a two-or-three-page note became Chapter 5 of my dissertation under the sober title of “Vertebral morphology and the evolution of long necks in sauropod dinosaurs”, weighing in at 41 1.5-spaced pages. By now the manuscript had metastatised into a comparison between the necks of sauropods and other animals and an analysis of the factors that enabled sauropods to achieve so much more than mammals, birds, other theropods and pterosaurs.

(At this point we had one of our less satisfactory reviewing experiences. We sent the manuscript to a respected journal, where it wasn’t even sent out to reviewers until more than a month had passed. We then had to repeatedly prod the editor before anything else happened. Eventually, two reviews came back: one of them careful and detailed; but the other, which we’d waited five months for, dismissed our 53-page manuscript in 108 words. So two words per page, or about 2/3 of a word per day of review time. But let’s not dwell on that.)

Figure 6. Basic cervical vertebral architecture in archosaurs, in posterior and lateral views. 1, seventh cervical vertebra of a turkey, Meleagris gallopavo Linnaeus, 1758, traced from photographs by MPT. 2, fifth cervical vertebra of the abelisaurid theropod Majungasaurus crenatissimus Depéret, 1896,UA 8678, traced from O’Connor (2007, figures 8 and 20). In these taxa, the epipophyses and cervical ribs are aligned with the expected vectors of muscular forces. The epipophyses are both larger and taller than the neural spine, as expected based on their mechanical importance. The posterior surface of the neurapophysis is covered by a large rugosity, which is interpreted as an interspinous ligament scar like that of birds (O’Connor, 2007). Because this scar covers the entire posterior surface of the neurapophysis, it leaves little room for muscle attachments to the spine. 3, fifth cervical vertebra of Alligator mississippiensis Daudin, 1801, MCZ 81457, traced from 3D scans by Leon Claessens, courtesy of MCZ. Epipophyses are absent. 4, eighth cervical vertebra of Giraffatitan brancai (Janensch, 1914) paralectotype HMN SII, traced from Janensch (1950, figures 43 and 46). Abbreviations: cr, cervical rib; e, epipophysis; ns, neural spine; poz, postzygapophysis.

This work made its next appearance as my talk at SVPCA 2010 in Cambridge, under the title Why giraffes have such short necks. For the talk, I radically restructured the material into a form that had a stronger narrative – a process that involved a lot of back and forth with Matt, dry-running the talk, and workshopping the order in which ideas were presented. The talk seemed to go down well, and we liked the new structure so much more than the old that we reworked the manuscript into a form that more closely resembled the talk.

That’s the version of the manuscript that we perfected in New York when we should have been at all-you-can-eat sushi places. It’s the version that we submitted on the train from New York to New Haven as we went to visit the collections of the Yale Peabody Museum. And it’s the version that was cursorily rejected from mid-to-low ranked palaeo journal because a reviewer said “The manuscript reads as a long “story” instead of a scientific manuscript” — which was of course precisely what we’d intended.

Needless to say, it was deeply disheartening to have had what we were convinced was a good paper rejected twice from journals, at a cost of three years’ delay, on the basis of these reviews. One option would have been to put the manuscript back into the conventional “scientific paper” straitjacket for the second journal’s benefit. But no. We were not going to invest more work to make the paper less good. We decided to keep it in its current, more readable, form and to find a journal that likes it on that basis.

At the moment, the plan is to send it to PeerJ when that opens to submissions. (Both Matt and I are already members.) But that three-years-and-rolling delay really rankles, and we both felt that it wasn’t serving science to keep the paper locked up until it finally makes it into a journal — hence the deposition in arXiv which we plan to talk about more next time.

Table 3. Neck-elongation features by taxon.

In the paper, we review seven characteristics of sauropod anatomy that facilitated the evolution of long necks: absolutely large body size; quadrupedal stance; proportionally small, light head; large number of  cervical vertebrae; elongation of cervical vertebrae; air-sac system; and vertebral pneumaticity. And we show that giraffes have only two of these seven features. (Ostriches do the next best, with five, but they are defeated by their feeble absolute size.)

The paper incorporates some material from SV-POW! posts, including Sauropods were corn-on-the-cob, not shish kebabs. In fact, come to think of it, we should have cited that post as a source. Oh well. We do cite one SV-POW! post: Darren’s Invading the postzyg, which at the time of writing is the only published-in-any-sense source for pneumaticity invading cervical postzygapogyses from the medial surface.

As for the non-extended epipophyses that kicked the whole project off: we did illustrate how they could look, and discussed why they would seem to make mechanical sense:

Figure 10. Real and speculative muscle attachments in sauropod cervical vertebrae. 1, the second through seventeenth cervical vertebrae of Euhelopus zdanskyi Wiman, 1929 cotype specimen PMU R233a-δ(“Exemplar a”). 2, cervical 14 as it actually exists, with prominent but very short epipophyses and long cervical ribs. 3, cervical 14 as it would appear with short cervical ribs. The long ventral neck muscles would have to attach close to the centrum. 4, speculative version of cervical 14 with the epipophyses extended posteriorly as long bony processes. Such processes would allow the bulk of both the dorsal and ventral neck muscles to be located more posteriorly in the neck, but they are not present in any known sauropod or other non-avian dinosaur. Modified from Wiman (1929, plate 3).

But we found and explained some good reasons why this apparently appealing arrangement would not work. You’ll need to read the paper for details.

Sadly, we were not able to include this slide from the talk illustrating the consequences:

Anyway, go and read the paper! It’s freely available, of course, like all arXiv depositions, and in particular uses the permissive Creative Commons Attribution (CC BY) licence. We have assembled related information over on this page, including full-resolution versions of all the figures.

In the fields of maths, physics and computer science, where deposition in arXiv is ubiquitous, standard practice is to go right ahead and cite works in arXiv as soon as they’re available, rather than waiting for them to appear in journals. We will be happy for the same to happen with our paper: if it contains information that’s of value to you, then feel free to cite the arXiv version.

Reference

  • Taylor, Michael P., and Mathew J. Wedel. 2012. Why sauropods had long necks; and why giraffes have short necks. arXiv:1209.5439. 39 pages, 11 figures, 3 tables. [Full-resolution figures]

In the recent post on OMNH 1670, a dorsal vertebra of a giant Apatosaurus from the Oklahoma panhandle, I half-promised to post the only published figure of this vertebra, from Stovall (1938: fig. 3.3). So here it is:

And in the second comment on that post, I promised a sketch from one of my notebooks, showing how much of the vertebra is reconstructed. Here’s a scan of the relevant page from my notebook. Reconstructed areas of the vert are shaded (confusingly, using strokes going in opposite directions on the spine and centrum, and the dark shaded areas on the front of the transverse processes are pneumatic cavities), and measurements are given in mm.

Next item: is this really a fifth dorsal vertebra?

Apatosaurus louisae CM 3018 D4 and D5, in anterior (top), left lateral, and posterior views, from Gilmore (1936: plate 25).

Here are D4 and D5 of A. louisae CM 3018. They sort of bracket OMNH 1670 in terms of morphology. D4 has a broader spine, and D5 has a narrower one. The spine of D5 lacks the slight racquet-shaped expansion seen in OMNH 1670, but the overall proportions of the spine are more similar. On the other hand, the transverse processes of D4 taper a bit in anterior and posterior view, as in OMNH 1670, and unlike the transverse processes of D5 with their more parallel dorsal and ventral margins. But honestly, neither of these verts is a very good match (and the ones on either side, D3 and D6, are even worse).

Apatosaurus parvus UWGM 15556 (formerly A. excelsus CM 563) D4 (left) and D3 (right) in anterior (top), right lateral, and posterior views, from Gilmore (1936: plate 32).

Here are D3 and D4 of A. parvus UWGM 15556. D3 is clearly a poor match as well–it is really striking how much the vertebral morphology changes through the anterior dorsals in most sauropods, and Apatosaurus is no exception. D3 looks like a dorsal in lateral view, but in anterior or posterior view it could almost pass for a posterior cervical. If I was going to use the term “cervicodorsal”, indicating one of the vertebrae from the neck/trunk transition, I would apply it as far back as D3, but not to D4. That thing is all dorsal.

And it’s a very interesting dorsal from the perspective of identifying OMNH 1670. It has fairly short, tapering transverse processes. The neural spine is a bit shorter and broader, but it has a similar racquet-shaped distal expansion. I’m particularly intrigued by the pneumatic fossae inscribed into the anterior surface of the neural spine–in Gilmore’s plate they make a broken V shapen, like so \ / (or maybe devil eyes). Now, OMNH 1670 doesn’t have devil eyes on its spine, but it does have a couple of somewhat similar pneumatic fossae cut into the spine just below the distal racquet–perhaps a serially modified iteration of the same pair of fossae as in the A. parvus D4. It’s a right sod that D5 from this animal has its spine blown off–but it still has its transverse processes, and they are short and tapering as in OMNH 1670.

Apatosaurus sp. FMNH P25112, dorsal vertebrae 1-10 and sacrals 1 and 2, Riggs (1903: plate 46)

Here are all the dorsals and the first couple sacrals of FMNH P25112, which was originally described as A. excelsus but in the specimen-level analysis of Upchurch et al. 2005) comes out as the sister taxon to the A. ajax/A. parvus/A. excelsus clade. Note the striking similarity of the D5 here with D4 of the A. parvus specimen in Gilmore’s plate (until the careful phylogenetic work up Upchurch et al. 2005, that A. parvus specimen, once CM 563 and now UWGM 15556, was considered to represent A. excelsus as well). But  also notice the striking similarity of D6 to OMNH 1670. It’s not quite a dead ringer–the transverse processes are longer and have weird bent-down “wingtips” (XB-70 Valkyrie, anyone?)–but it’s pretty darned close, especially in the shape of the neural spine.

So what does this all mean? First, that trying to specify the exact serial position of an isolated vertebra is nigh on to impossible, unless it’s something that is one-of-a-kind like an axis. Second, after doing all these comparos I think it’s unlikely that OMNH 1670 is a D4–those are a bit too squat across the board–but it could plausibly be either a D5 or a D6. Third, I’m really happy that it doesn’t seem to match any particular specimen better than all the rest. What I don’t want to happen is for someone to see that this vertebra looks especially like specimen X and therefore decide that it must represent species Y. As I said in the comments of the previous post, what this Oklahoma Apatosaurus material needs is for someone to spend some quality time seeing, measuring, and photographing all of it and then doing a phylogenetic analysis. That sounds like an ambitious master’s thesis or the core of a dissertation, and I hope an OU grad student takes it on someday.

If you were intrigued by my suggestion that the big Oklahoma Apatosaurus rivalled Supersaurus in size, and wanted to see a technical comparison of the two, I am happy to report that Scott Hartman has done the work for you. Here’s one of his beautiful Apatosaurus skeletal reconstructions, scaled to the size of OMNH 1670, next to his Supersaurus silhouette. This is just a small teaser–go check out his post on the subject for a larger version and some interesting (and funny) thoughts on how the two animals compare.

References

  • Gilmore, C.W. 1936. Osteology of Apatosaurus with special reference to specimens in the Carnegie Museum. Memoirs of the Carnegie Museum 11:175-300.
  • Riggs, E.S. 1903. Structure and relationships of opisthocoelian dinosaurs, part I: Apatosaurus Marsh. Field Columbian Museum Publications, Geological Series 2(4): 165–196.
  • Stovall, J.W. 1938. The Morrison of Oklahoma and its dinosaurs. Journal of Geology 46:583-600.

Left: the Queen of England, 163 cm.  Middle, the Oklahoma apatosaur dorsal, 135 cm.  Right, classic “big Apatosaurus” dorsal, 106 cm.  To scale.

Something I’ve always intended to do but never gotten around to is posting on some of the immense Apatosaurus elements from the Oklahoma panhandle. Here’s one of the most impressive, OMNH 1670, an isolated dorsal. Notice that the tip of the neural spine is ever-so-shallowly bifurcated, which in Apatosaurus indicates a D4, D5, or D6. The low parapophyses and fat transverse processes are similar to D4, but Apatosaurus D4s usually have somewhat broader spines, so I’m guessing this thing is a D5. These things vary and I could easily be off by a position in either direction.

Next to it is D5 of CM 3018, the holotype specimen of Apatosaurus louisae (from Gilmore 1936: plate 25), which has served as the basis for many of the published mass estimates of the genus Apatosaurus. OMNH 1670 is 135 cm tall, compared to 106 cm for D5 of CM 3018. If the rest of the animal scaled the same way, it would have been 1.27^3 = 2 times as massive. Mass estimates for CM 3018 are all over the map, from about 18 tons up to roughly twice that, so the big Oklahoma Apatosaurus was probably in Supersaurus territory, mass-wise, and may have rivaled some of the big titanosaurs (Update: see the two giant diplodocids square off in a cool follow-up post by Supersaurus wrangler Scott Hartman). Here’s a fun rainy-day activity: take any skeletal reconstruction of Apatosaurus, clone it in Photoshop or GIMP, scale it up by 27%, and park it next to the original. It looks a lot bigger. So I’m continually surprised that Apatosaurus is so rarely mentioned in the various roundups of giant sauropods, both in the technical literature and in popular articles online. This vertebra was figured by Stovall (1938)–if I get inspired, I’ll dig up that figure and post it another day (hey, look, I did!).

Fun fact: in Apatosaurus the tallest (most posterior) dorsals are 1.3-1.5 times as tall as D5 (Gilmore 1936: 201). So D10 from this individual was probably between 1.7 and 2 meters tall–not quite in Amphicoelias fragillimus territory but getting closer than I’ll bet most people suspected.

NB: if you try to use the scale bar lying on the centrum of OMNH 1670 to check my numbers, you will get a wonky answer. The problem is that the vertebra is so large that it is almost impossible to get far enough back from it (above it, in this case, since it is lying on a padded pallet) to get a shot free from distortion due to parallax. For this shot, the pallet with the vert was on the floor, and I was standing on top of the tallest ladder in the OMNH collections, leaning out over the vert to get centered over the prezygapophyses, and shooting straight down–in other words, I had done everything possible to minimize the visual distortion. But it still crept in. Anyway, trust the measurements, which I–and presumably Gilmore–made with a good old reliable tape measure.

References

  • Gilmore, C.W. 1936. Osteology of Apatosaurus with special reference to specimens in the Carnegie Museum. Memoirs of the Carnegie Museum 11:175-300.
  • Stovall, J.W. 1938. The Morrison of Oklahoma and its dinosaurs. Journal of Geology 46:583-600.

Vanessa Graff and I spent yesterday working in the herpetology and ornithology collections at the Natural History Museum of Los Angeles County (LACM). The herpetology collections manager, Neftali Comacho, pointed us to this skull of Alligator mississippiensis. It’s not world’s biggest gator–about which more in a second–but it’s the biggest I’ve seen in person. Normally it lives in a big rubbermaid tub in the collections area, but this Sunday it will be out on display for Reptile and Amphibian Appreciation Day (RAAD) at the LACM. RAAD will include guest talks, tours of the collections, and live animal demonstrations. If you’re in SoCal and you’re into herps–or have kids, grandkids, nephews or nieces that are into herps–it will be well worth checking out. While you’re there, don’t neglect the newly renovated Age of Dinosaurs and Age of Mammals halls, which are frankly phenomenal: spacious, well-lit, loads of actual material on display, skeletons you can walk all the way around, informative but unobtrusive signage, tasteful integration with existing architecture…I could go on. Better if you just go and see for yourself.

About that gator. First the bad news.  It came to the LACM from another collection, and has no data–no locality, no date collected, nothing. The skull is also missing all of its teeth, the left retroarticular process, the back end of the braincase and the occipital condyle. I think the latter losses were probably caused by a foramen of Winchester.*

Now, the awesome news. The length from the snout tip to the end of the articulars was 680mm and from the snout to the end of the quadrates was 590mm. Irritatingly I did not get a dorsal head length, which is the gold standard for comparative croc skull measurements, because I only reread Darren’s giant croc skull post after I got home last night. Going from the photos, I think the dorsal head length was right around 50 cm (beware, the yardstick in the photos is marked off in inches).

Darren’s post led me to this one, which has some very useful measurements (yay!) of giant croc skulls. The table at the end of that post lists alligator skulls with dorsal head lengths of 58, 60, and 64 cm, so the big LACM gator is nowhere near being the world’s largest. In fact, the 64 cm skull would be a quarter again as large, which is a truly horrifying thought. Still, it’s a big damn skull from a big damn gator.

You might get the impression that here in the Wedel lab we are shamelessly obsessed with giant saurians. And that is in fact true. But we also look at tiny ones, too. Here I’m playing with the skull of a little Tomistoma, the false gharial. Tomistoma is notable because another individual of the genus produced the longest skull of any known extant crocodilian–a whopping 84 cm dorsal head length (photos of this monster are in both of the giant croc skull posts linked above).

The moral of the story? If the sign says don’t go swimming, don’t go swimming. Go to RAAD instead, and see the giant alligator skull, and a ton of other cool stuff besides. And if you’re into gator skulls or just like geeking out on awesome anatomy, check out the 3D Alligator Skull site, a joint project of the Holliday lab and Witmer lab. Have fun!

* bullet hole

In a new comment on an oldish post, Peter Adlam asked:

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.

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