Illustration talk slide 58

Illustration talk slide 59

Illustration talk slide 60

The rest of the series.

References

Sauroposeidon and friends

February 24, 2014

Sauroposeidon and kin cervicals - DRAFTAs a break from photography posts, here are four pretty big vertebrae that swirl in the same thought-space in my head. All are shown to scale, in right lateral view. These are not the biggest sauropod cervical vertebrae–Supersaurus beats them all, and there are vertebrae of Puertasaurus, Alamosaurus, and Futalognkosaurus that rival the big Sauroposeidon vert, but those are either less well preserved or still awaiting detailed description.

Incidentally, I think BYU 12867 is a C10. The centrum proportions are about right, compared to Giraffatitan, and the neural spine looks good, too, like a geometric transformation of the big Giraffatitan C8. Also, the drawn-in prezyg outline for MIWG.7306 is a little short; the actual prezyg is a monster and would have overhung the condyle by another 10cm or so. I’m pretty sure that we had a composite photograph showing this at one point, but irritatingly none of us can find it at the moment. If it turns up, I’ll update the image.

For a long time I thought Sauroposeidon was a brachiosaurid. Now it seems to be a somphospondyl (D’Emic 2012) or possibly even a basal titanosaur (Mannion et al. 2013), even if we stick just to the holotype. But if it’s not a brachiosaurid, it’s cervical vertebrae are at least coarsely brachiosaur-y in outline.

You  may recall from Naish et al. (2004) that MIWG.7306 shares several derived characters with the holotype vertebrae of Sauroposeidon. Does that mean that Angloposeidon is a somphospondyl or titanosaur as well? I dunno–as always, we need more material–but it’s an interesting possibility.

References

Illustration talk slide 19

Illustration talk slide 20

Illustration talk slide 21

Illustration talk slide 22

This whole section, including the title, is mostly swiped from Mike’s Tutorial 17.

Other posts in this series are here.

Papers referenced in these slides:

Giraffatitan skull photos

February 10, 2014

Giraffatitan skull left lateral

Let it never be said that we don’t take good care of our commenters. Heck, we’ll even degrade ourselves by blogging about theropods, if that’s what it takes to keep you all happy.

Giraffatitan skull left anterolateral

Derp dah durr

Today’s post is a response to this comment by Dean, asking for lateral view photos of the skull of Giraffatitan. Mike and I did get to spend some quality time with the T1 skull (a.k.a. “Old Toilet-Face”) when we were in Berlin in 2008.

Giraffatitan skull anterior

Unfortunately, most of our photos turned out not-so-hot. The room around the skull was not large, so we couldn’t get back very far from it. Hence our photos are plagued by perspective distortions.

Giraffatitan skull right anterolateral

Ah hurr hurr hurr

Also, we didn’t have a tripod along and the light level was fairly low, and the combination of handheld shots and long exposure times meant that most of the shots are at least a little blurry.

Giraffatitan skull right lateral

BUT. It was still a thrill to see that skull up close.

The crazy thing about Giraffatitan is that the skull looks like it’s going to be pretty sweet when you see it from the side. Because you’re thinking it’s going to be kinda narrow, like a giraffe’s head. Then you get even a partial front view and suddenly the animal’s whole skull looks like a partially-deflated whoopie cushion (whereas in life it looked like a mostly-inflated whoopie cushion). And then you have to live with the knowledge that one of the most majestic animals that ever lived on Earth was afflicted with derpty-face. I’ll bet they went extinct from shame.

Giraffatitan skull dorsal oblique

Still, there is some cool anatomy to see here, especially the snout-troughs leading down from the external nares, and the neurovascular foramina on the maxillae.

And, crucially, brachiosaurs had the good taste to hide their freakish countenances 45 feet up, where they could be safely ignored by everyone other than pterosaurs and birds. This has not escaped the notice of exhibit designers:

Giraffatitan skeleton hmmm

Go here for the unmarked original.

“Look at all the things you’ve done for me
Opened up my eyes,
Taught me how to see,
Notice every tree.”

So sings Dot in Move On, the climactic number of Stephen Sondheim’s Pulitzer Prize-winning music Sunday in the Park with George, which on the surface is about the post-impressionist painter Georges Seurat, but turns out to be a study of obsession and creativity.

xx

Un dimanche après-midi à l’Île de la Grande Jatte – 1884 [A Sunday Afternoon on the Island of La Grande Jatte – 1884]

“Taught me how to see”? What kind of talk is that? One the surface, it seems silly — we all know how to see. We do it constantly, without thinking. Yet it’s something that artists talk about all the time. And anyone who’s sat down and seriously tried to paint or draw something will have some understanding of what the phrase means. We have such strong implicit ideas of what things look like that we tend to reproduce what we “know” is there rather than what’s actually there. Like I said, we see without thinking.

In fact, the psychology of perception is complicated and sophisticated, and the brain does an extraordinary amount of filtering of the visual signals we get, to save us the bother of having to consciously process way too much data. This is a whole scientific field of its own, and I’m going to avoid saying very much about it for fear of making a fool of myself — as scientists so often do when wandering outside their own field. But I think it’s fair to say that we all have a tendency to see what we expect to see.

xx

Phylogeny of Sauropoda, strict consensus of most parsimonious trees according to Wilson (2002:fig. 13a)

In the case of sauropods, this tendency has meant that we’ve all been startlingly bad at seeing pneumaticity in the caudal vertebrae of sauropods. Because the literature has trained us to assume it’s not there. For example, in the two competing sauropod phylogenies that dominated the 2000s, both Wilson (2002) and Upchurch et al. (2004) scored caudal pneumaticity as very rare: Wilson’s character 119, “Anterior caudal centra, pneumatopores (pleurocoels)”, was scored 1 only for Diplodocus and Barosaurus; and  Upchurch et al. (2004:286) wrote that “A few taxa (Barosaurus, Diplodocus, and Neuquensaurus) have pleurocoel-like openings in the lateral surfaces of the cranial [caudal] centra that lead into complex internal chambers”. That’s all.

And that’s part of the reason that every year since World War II, a million people have walked right past the awesome mounted brachiosaur in the Museum Für Naturkunde Berlin without noticing that it has pneumatic caudals. After all, we all knew that brachiosaur caudals were apneumatic.

But in my 2005 Progressive Palaeontology talk about upper limits on the mass of land animals estimated through the articular area of limb-bone cartilage, I included this slide that shows how much bigger the acetabulum of Giraffatitan is than the femoral head that it houses:

Screenshot from 2014-01-24 17:30:30

And looking at that picture made me wonder: those dark areas on the sides of the first few caudals (other than the first, which is a very obvious plaster model) certainly look pneumatic.

Then a few years later, I was invited to give a talk at the Museum Für Naturkunde Berlin itself, on the subject “Brachiosaurus brancai is not Brachiosaurus“. (This of course was drawn from the work that became my subsequent paper on that subject, Taylor 2009) And as I was going through my photos to prepare the slides of that talk, I thought to myself: darn it, yes, it does have pneumatic caudals!

So I threw this slide into the talk, just in passing:

Screenshot from 2014-01-24 17:32:06

Those photos were pretty persuasive; and a closer examination of the specimen on that same trip was to prove conclusive.

Meanwhile …

Earlier in 2009, I’d been in Providence, Rhode Island, with my Index Data colleagues. I’d managed to carve a day out of the schedule to hope along the coast to the Yale Peabody Museum in New Haven, Connecticut. My main goal was to examine the cervicals of the mounted Apatosaurus (= “Brontosaurus“) excelsus holotype (although it was also on that same trip that I first saw the Barosaurus holotype material that we’ve subsequently published a preprint on).

The Brontosaurus cervicals turned out to be useless, being completely encased in plaster “improvements” so that you can’t tell what’s real and what’s not. hopefully one day they’ll get the funding they want to take that baby down off its scaffold and re-prep the material.

But since I had the privilege of spending quality time with such an iconic specimen, it would have been churlish not to look at the rest of it. And lo and behold, what did I see when I looked at the tail but more pneumaticity that we thought we knew wasn’t there!

Wedel and Taylor (2013b: Figure 10).

An isolated pneumatic fossa is present on the right side of caudal vertebra 13 in Apatosaurus excelsus holotype YPM 1980. The front of the vertebra and the fossa are reconstructed, but enough of the original fossil is visible to show that the feature is genuine. (Wedel and Taylor 2013b: Figure 10).

What does this mean? Do other Giraffatitan and Apatosaurus specimens have pneumatic tails? How pervasive is the pneumaticity? What are the palaeobiological implications?

Stay tuned! All will be revealed in Matt’s next post (or, if you can’t wait, in our recent PLOS ONE paper, Wedel and Taylor 2013b)!

References

I just found out — thanks to a tweet from abertonykus — that this exists:

sauropod_vertebra_picture_adventure__by_classicalguy-d6ssfil

That’s me on top of the Giraffatitan, Matt to the right, and Darren swinging from its wattle.

It’s the work of classicalguy on Deviant Art. He provides a poem and some brief commentary along with the original. There also one for the Tetrapod Zoology podcats, and one for Tom Holtz.

As I mentioned a few days ago, Matt and I have a couple of papers in the new PLOS ONE Sauropod Gigantism collection. We were each lead author on one and second author on the other, so for convenience’s sake we’ll refer to them as my paper (Taylor and Wedel 2013c on neck cartilage) and Matt’s paper (Wedel and Taylor 2013b on caudal pneumaticity.)

Mine is very simple in concept (although it ended up at 17 pages and 23 figures). It’s all about addressing one of the overlooked variables in reconstructing the postures of the necks of sauropods (and indeed of all tetrapods). That is, the spacing between consecutive vertebrae, and the effect this has on “neutral pose”.

The concept of “neutral pose” goes back to the DinoMorph work of Stevens and Parrish (1999). They defined it (p. 799) as follows: “We determined the neutral poses for each animal, wherein the paired articular facets of the postzygapophyses of each cervical vertebra were centered over the facets of the prezygapophyses of its caudally adjacent counterpart.”

x

Taylor and Wedel (2013c: Figure 3). Articulated sauropod vertebrae. Representative mid-cervical vertebra of Giraffatitan brancai, articulating with its neighbours. The condyle (ball) on the front of each vertebra’s centrum fits into the cotyle (socket) at the back of the preceding one, and the prezygapophyses articulate with the preceding vertebra’s postzygapophyses. These vertebrae are in Osteological Neutral Pose, because the pre- and postzygapophyseal facets overlap fully.

One of the more fundamental flaws in Stevens and Parrish (1999) is the assumption that animals habitually rest their necks in neutral pose — an assumption that is unsupported by evidence and, as it turns out, false (Vidal et al. 1986, Taylor et al. 2009). But let’s leave that aside for the moment, and consider what neutral pose actually represents.

The fact that there is even such a thing as neutral articulation between two consecutive vertebrae is due to there being three points of contact between those vertebra: as with the legs of a tripod, three points is the minimum number you need to fix an object in three-dimensional space. Two of these points are at the zygapophyses, as noted in the original definition above. The third point is the articulation between the centra.

The centrum has been curiously overlooked in discussions of neutral pose, but needless to say its length is crucial in establishing what is neutral. In the image above, if the centrum was longer, then the angle between the consecutive vertebrae would need to be raised in order to keep the zygapophyses articulated.

And of course it was longer in life, because of the cartilage in between the consecutive centra. (The use of the more specific term “osteological neutral pose” goes some way to recognising that tissues other than bone have been overlooked, but the problem has not really been addressed or even properly acknowledged in published works before our paper.)

xx

Taylor and Wedel (2013c: Figure 5). Intervertebral gaps in camel necks. Head and neck of dromedary camels. Top: UMZC H.14191, in right lateral view, posed well below habitual posture, with apparently disarticulated C3/C4 and C4/C5 joints. Photograph taken of a public exhibit at University Museum of Zoology, Cambridge, UK. Bottom: OUMNH 17427, in left lateral view, reversed for consistency with Cambridge specimen. Photograph taken of a public exhibit at Oxford University Museum of Natural History, UK. Inset: detail of C4 of the Oxford specimen, showing articulations with C3 and C5. The centra are separated by thick pads of artificial ‘‘cartilage’’ to preserve spacing as in life.

You simply can’t ignore cartilage when modelling neck postures and expect to get anything resembling a meaningful result. That is, presumably, the reason why the habitual posture of rabbits in life exceeds the most extended posture we were able to obtain when manipulating dry vertebrae of a hare: compare Vidal et al. (1986: fig. 4) with Taylor et al. (2009: fig. 1).

How big is the effect? That depends on the thickness of the cartilage and the height of the zygapophyses above the center of rotation. Here is an illustration that we should have put in the paper, but which inexplicably neither of us thought of:

figNEW-angle-at-zygs

Influence of intervertebral cartilage on vertebral articulation angle. Consider the posterior vertebra (black) as fixed. The blue vertebra represents neutral pose of the preceding vertebra with centra abutting and zygapophyseal facets maximally overlapped. The red vertebra indicates neutral pose once intervertebral cartilage is added between the vertebra (where else?) The green lines show the angle by which the more anterior vertebra must be inclined in order to accommodate the cartilage, and the magenta line shows the height of the zygapophyseal articulation above the center of rotation between the two vertebrae.

Here’s some elementary trigonometry. Suppose the intervertebral cartilage is x distance thick at mid-height of the centra, and that the height of the zygs above this mid-height point (the magenta line) is y. The triangle between the middle of the condyle of the posterior vertebra, the middle of the cotyle of the anterior one and the zygapophyseal articulation is near enough a right-angled triangle as makes no odds.

Consider the angle θ between the green lines. Sin(θ) = opposite/hypotenuse = x/y, and by similarity, the additional angle of inclination of the anterior vertebra is also θ.

But for small angles (and this is generally a small angle), sin(θ) ≈ θ. So the additional inclination in radians = cartilage thickness divided by zygapophyseal height. For example, in vertebrae where the zygs are 23 cm above the mid-height of the centra, adding 4 cm of intervertebral cartilage adds about 4/23 = 0.174 radians = 10 degrees of extra inclination. (That’s pretty similar to the angle in the illustration above. Eyeballing the cartilage thickness and zyg height in the illustration suggests that 23:4 ratio is about right, which is a nice sanity-check of this method.)

millionaire-stupid-contestant4

At this point, I am cursing my own stupidity for not putting this diagram, and the very simple calculation, into the paper. I guess that can happen when something is written in a hurry (which to be honest this paper was). The formula is so simple — and accurate enough within tolerances of inevitable measurement error — that we really should have used it all over the place. I guess that will have to go in a followup now.

Anyway — next time, we’ll address this important related question: how thick, in fact, was the cartilage between the cervicals of sauropods?

References

This is an exciting day: the new PLOS Collection on sauropod gigantism is published to coincide with the start of this year’s SVP meeting! Like all PLOS papers, the contents are free to the world: free to read and to re-use.  (What is a Collection? It’s like an edited volume, but free online instead of printed on paper.)

There are fourteen papers in the new Collection, encompassing neck posture (yay!), nutrition (finally putting to bed the Nourishing Vomit Of Eucamerotus hypothesis), locomotion, physiology and evolutionary ecology. Lots every sauropod-lover to enjoy.

x

Taylor and Wedel (2013c: Figure 12). CT slices from fifth cervical vertebrae of Sauroposeidon. X-ray scout image and three posterior-view CT slices through the C5/C6 intervertebral joint in Sauroposeidon OMNH 53062. In the bottom half of figure, structures from C6 are traced in red and those from C5 are traced in blue. Note that the condyle of C6 is centered in the cotyle of C5 and that the right zygapophyses are in articulation.

Matt and I are particularly excited that we have two papers in this collection: Taylor and Wedel (2013c) on intervertebral cartilage in necks, and Wedel and Taylor (2013b) on pneumaticity in the tails of (particularly) Giraffatitan and Apatosaurus. So we have both ends of the animal covered. It also represents a long-overdue notch on our bed-post: for all our pro-PLOS rhetoric, this is the first time either of has had a paper published in a PLOS journal.

Wedel and Taylor (2013b: Figure 4). Giraffatitan brancai tail MB.R.5000 (‘Fund no’) in right lateral view. Dark blue vertebrae have pneumatic fossae on both sides, light blue vertebrae have pneumatic fossae only on the right side, and white vertebrae have no pneumatic fossae on either side. The first caudal vertebra (hatched) was not recovered and is reconstructed in plaster.

It’s a bit of a statistical anomaly that after a decade of collaboration in which there was never a Taylor & Wedel or Wedel & Taylor paper, suddenly we have five of them out in a single year (including the Barosaurus preprint, which we expect to eventually wind up as Taylor and Wedel 2014). Sorry about the alphabet soup.

Since Matt is away at SVP this week, I’ll be blogging mostly about the Taylor and Wedel paper this week. When Matt returns to civilian life, the stage should be clear for him to blog about pneumatic caudals.

Happy days!

References

Let’s take another look at that Giraffatitan cervical. MB.R.2180:C5, from a few days ago:

FigureA-Giraffatitan-SI-C5

That’s a pretty elongate vertebra, right? But how elongate, exactly? How can we quantify whether it’s more or less elongate than some other vertebra?

The traditional answer is that we quantify elongation using the elongation index, or EI. This was originally defined by Upchurch (1998:47) as “the length of a vertebral centrum divided by the width across its caudal face”. Measuring from the full-resolution version of the image above, I make that 1779/529 pixels, or 3.36.

But then those doofuses Wedel et al. (2000:346) came along and said:

When discussing vertebral proportions Upchurch (1998) used the term elongation index (EI), defined as the length of the centrum divided by the width of the cotyle. Although they did not suggest a term for the proportion, Wilson & Sereno (1998) used centrum length divided by the height of the cotyle as a character in their analysis. We prefer the latter definition of this proportion, as the height of the cotyle is directly related to the range of motion of the intervertebral joint in the dorsoventral plane. For the purposes of the following discussion, we therefore redefine the EI of Upchurch (1998) as the anteroposterior length of the centrum divided by the midline height of the cotyle.

Since then, the term EI has mostly been used in this redefined sense — but I think we all agree now that it would have been better for Wedel et al to have given a new name to Wilson and Sereno’s ratio rather than apply Upchurch’s name to it.

Aaaanyway, measuring from the image again, I give that vertebra an EI (sensu Wedel et al. 2000) of 1779/334 = 5.33. Which is 58% more elongate than when using the Upchurch definition! This of course follows directly from the cotyle being 58% wider than tall (529/334 pixels).

So one of principal factors determining how elongate a vertebra seems to be is the shape of its cotyle. And that’s troublesome, because the cotyle is particularly subject to crushing — and it’s not unusual for even consecutive vertebrae from the same column to be crushed in opposite directions, giving them (apparently) wildly different EIs.

Here’s an example (though not at all an extreme one): cervicals 4 and 6 of the same specimen, MB.R.2180 (formerly HM SI), as the multi-view photo above:

DSCN5527-5535-SI-c4-and-c6-posterior

Measuring from the photos as before, I make the width:height ratio of C4 683/722 pixels = 0.95, and that of C6  1190/820 pixels = 1.45. So these two vertebrae — from the same neck, and with only one other vertebrae coming in between them — differ in preserved cotyle shape by a factor of 1.53.

And by the way, this is one of the best preserved of all sauropod neck series.

Let’s take a look at the canonical well-preserved sauropod neck: the Carnegie Diplodocus, CM 84. Here are the adjacent cervicals 13 and 14, in posterior view, from Hatcher (1901: plate VI):

Hatcher1901-plate-VI-C13-C14-posterior

For C14 (on the left), I get a width:height ratio of 342/245 pixels = 1.40. For C13 (on the right), I get 264/256 pixels = 1.03. So C14 is apparently 35% broader than its immediate predecessor. I absolutely don’t buy that this represents how the vertebrae were in life.

FOR EXTRA CREDIT: what does this tell us about the reliability of computer models that purport to tell us about neck posture and flexibility, based on the preserved shapes of their constituent vertebrae?

So what’s to be done?

The first thing, as always in science, is to be explicit about what statements we’re making. Whenever we report an elongation index, we need to clearly state whether it’s EI sensu Upchurch 1998 or EI sensu Wedel et al. 2000. Since that’s so cumbersome, I’m going propose that we introduce two new abbreviations: EIH (Elongation Index Horizonal), which is Upchurch’s original measure (length over horizontal width of cotyle) and EIV (Elongation Index Vertical), which is Wilson and Sereno’s measure (length over vertical height of cotyle). If we’re careful to report EIH and EIV (or better still both) rather than an unspecified EI, then at least we can avoid comparing apples with oranges.

But I think we can do better, by combining the horizontal and vertical cotyle measurements in some way, and dividing the length by the that composite. This would give us an EIA (Elongation Index Average), which we could reasonably expect to preserve the original cotyle size, and so to give a more reliable indication of “true” elongation.

The question is, how to combine the cotyle width and height? There are two obvious candidates: either take the arithmetic mean (half the sum) or the geometric mean (the square root of the product). Note that for round cotyles, both these methods will give the same result as each other and as EIH and EIV — which is what we want.

Which mean should we use for EIA? to my mind, it depends which is best preserved when a vertebra is crushed. If a 20 cm circular cotyle is crushed vertically to 10cm, does it tend to smoosh outwards to 30 cm (so that 10+30 = the original 20+20) or to 40 cm (so that 10 x 40 = the original 20 x 20)? If the former, then we should use arithmetic mean; if the latter, then geometric mean.

Does anyone know how crushing works in practice? Which of these models most closely approximates reality? Or can we do better than either?

Update (8:48am): thanks for Emanuel Tschopp for pointing out (below) what I should have remembered: that Chure et al.’s (2010) description of Abydosaurus introduces “aEI”, which is the same as one of my proposed definitons of EIA. So we should ignore the last four paragraphs of this post and just use aEI. (Their abbreviation is better, too.)

 

References

  • Hatcher, Jonathan B. 1901. Diplodocus (Marsh): its osteology, taxonomy and probable habits, with a restoration of the skeleton. Memoirs of the Carnegie Museum 1:1-63 and plates I-XIII.
  • Upchurch, Paul. 1998. The phylogenetic relationships of sauropod dinosaurs. Zoological Journal of the Linnean Society 124:43-103.
  • Wedel, Mathew J., Richard L. Cifelli and R. Kent Sanders. 2000b. Osteology, paleobiology, and relationships of the sauropod dinosaur Sauroposeidon. Acta Palaeontologica Polonica 45(4):343-388.
  • Wilson, J. A. and Paul C. Sereno. 1998. Early evolution and higher-level phylogeny of sauropod dinosaurs. Society of Vertebrate Paleontology, Memoir 5:1-68.

I thought I’d done a decent job of illustrating MB.R.2180:C5 last time, but Wedel was not satisfied, demanding ventral and right-lateral views as well as the provided right lateral, anterior, posterior and dorsal.

All right then: here you go!

FigureA-Giraffatitan-SI-C5

Here once more, for comparison, is Janensch’s (1950) illustration of the same vertebra:

Janensch1950-figs-23-25

 

As you’ll see, I changed the composition of my version, now that I have a right lateral view, to more closely match the composite of Janensch’s figures. The third row of mine is now exactly the same composition as I used for his illustrations, so it’s easier to compare the two.

 

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