Last night, I submitted a paper for publication — for the first time since April 2013. I’d almost forgotten what it felt like. But, because we’re living in the Shiny Digital Future, you don’t have to wait till it’s been through review and formal publication to read it. I submitted to PeerJ, and at the same time, made it available as a preprint (Taylor 2014).

It’s called “Quantifying the effect of intervertebral cartilage on neutral posture in the necks of sauropod dinosaurs”, and frankly the results are weird. Here’s a taste:

Taylor (2014:figure 3). Effect of adding cartilage to the neutral pose of the neck of Apatosaurus louisae CM 3018. Images of vertebra from Gilmore (1936:plate XXIV). At the bottom, the vertebrae are composed in a horizontal posture. Superimposed, the same vertebrae are shown inclined by the additional extension angles indicated in Table 1. If the slightly sub-horizontal osteological neutral pose of Stevens and Parrish (1999) is correct, then the cartilaginous neutral pose would be correspondingly slightly lower than depicted here, but still much closer to the elevated posture than to horizontal. (Note that the posture shown here would not have been the habitual posture in life: see discussion.)

Taylor (2014:figure 3). Effect of adding cartilage to the neutral pose of the neck of Apatosaurus louisae CM 3018. Images of vertebra from Gilmore (1936:plate XXIV). At the bottom, the vertebrae are composed in a horizontal posture. Superimposed, the same vertebrae are shown inclined by the additional extension angles indicated in Table 1. If the slightly sub-horizontal osteological neutral pose of Stevens and Parrish (1999) is correct, then the cartilaginous neutral pose would be correspondingly slightly lower than depicted here, but still much closer to the elevated posture than to horizontal. (Note that the posture shown here would not have been the habitual posture in life: see discussion.)

A year back, as I was composing a blog-post about our neck-cartilage paper in PLOS ONE (Taylor and Wedel 2013c), I found myself writing down the rather trivial formula for the additional angle of extension at an intervertebral joint once the cartilage is taken into account. In that post, I finished with the promise “I guess that will have to go in a followup now”. Amazingly it’s taken me a year to get that one-pager written and submitted. (Although in the usual way of things, the manuscript ended up being 13 pages long.)

To summarise the main point of the paper: when you insert cartilage of thickness t between two vertebrae whose zygapophyses articulate at height h above the centra, the more anterior vertebra is forced upwards by t/h radians. Our best guess for how much cartilage is between the adjacent vertebrae in an Apatosaurus neck is about 10% of centrum length: the image above shows the effect of inserting that much cartilage at each joint.

And yes, it’s weird. But it’s where the data leads me, so I think it would be dishonest not to publish it.

I’ll be interested to see what the reviewers make of this. You are all of course welcome to leave comments on the preprint itself; but because this is going through conventional peer-review straight away (unlike our Barosaurus preprint), there’s no need to offer the kind of detailed and comprehensive comment that several people did with the previous one. Of course feel free if you wish, but I’m not depending on it.

References

Gilmore Charles W. 1936. Osteology of Apatosaurus, with special reference to specimens in the Carnegie Museum. Memoirs of the Carnegie Museum 11:175–300 and plates XXI–XXXIV.

Stevens, Kent A., and J. Michael Parrish. 1999. Neck posture and feeding habits of two Jurassic sauropod dinosaurs. Science 284(5415):798–800. doi:10.1126/science.284.5415.798

Taylor, Michael P. 2014. Quantifying the effect of intervertebral cartilage on neutral posture in the necks of sauropod dinosaurs. PeerJ PrePrints 2:e588v1 doi:10.7287/peerj.preprints.588v1

Taylor, Michael P., and Mathew J. Wedel. 2013c. The effect of intervertebral cartilage on neutral posture and range of motion in the necks of sauropod dinosaurs. PLOS ONE 8(10):e78214. 17 pages. doi:10.1371/journal.pone.0078214

SO close

August 21, 2014

Bipedal Diplodocus USNM 10865 - modified from Gilmore 1932 pl 6 - v2

I have often argued that given their long hindlimbs, massive tail-bases, and posteriorly-located centers of mass, diplodocids were basically bipeds whose forelimbs happened to reach the ground. I decided to see what that might look like.

Okay, now obviously I know that there are no trackways showing sauropods actually getting around like this. It’s just a thought experiment. But given how close the center of mass of Diplodocus is to the acetabulum, I’ll bet that this pose was achievable in life. If diplodocids had just pushed the CM a few cm farther back, they might have dispensed with forelimbs entirely, or done something different with them, like re-evolved grasping hands.

Image modified from Gilmore (1932: plate 6). Here’s a horizontal-necked bipedal Diplodocus and the original pose:

Bipedal Diplodocus USNM 10865 - modified from Gilmore 1932 pl 6

Diplodocus USNM 10865 - Gilmore 1932 pl 6 - cleaned up

UPDATE the next day: I had forgotten that Niroot had already done a bipedal Apatosaurus, and a much more convincing one than mine. Go see it.

UPDATE the next week: Well, heck. Looks like the primary value of this post was so that people would remind me of all the other places the same idea has already been covered better. As you can see from the comment thread, Mike blogged about this at the WWD site, Scott Hartman drew it, and Heinrich Mallison showed that it was plausible. Sheesh, I suck.

Reference

  • Gilmore, C. W. 1932. On a newly mounted skeleton of Diplodocus in the United States National Museum. Proceedings of the United States National Museum 81, 1-21.

Supersaurus vs Brachiosaurus - BYU 9024 and FMNH P25107

This was inspired by an email Mike sent a couple of days ago:

Remind yourself of the awesomeness of Giraffatitan:
http://svpow.files.wordpress.com/2008/11/mike-by-jango-elbow.jpeg

Now think of this. Its neck is 8.5m long. Knock of one measly meter — for example, by removing one vertebra from the middle of the neck — and you have 7.5 m.

Supersaurus’s neck was probably TWICE that long.

Holy poo.

I replied that I was indeed freaked out, and that it had given me an idea for a post, which you are now reading. I didn’t have a Giraffatitan that was sufficiently distortion-free, so I used my old trusty Brachiosaurus. The vertebra you see there next to Mike and next to the neck of Brachiosaurus is BYU 9024, the longest vertebra that has ever been found from anything, ever.

Regarding the neck length of Supersaurus, and how BYU 9024 came to be referred to Supersaurus, here’s the relevant chunk of my dissertation (Wedel 2007: pp. 208-209):

Supersaurus is without question the longest-necked animal with preserved cervical material. Jim Jensen recovered a single cervical vertebra of Supersaurus from Dry Mesa Quarry in western Colorado. The vertebra, BYU 9024, was originally referred to “Ultrasauros”. Later, both the cervical and the holotype dorsal of “Ultrasauros” were shown to belong to a diplodocid, and they were separately referred to Supersaurus by Jensen (1987) and Curtice et al. (1996), respectively.

BYU 9024 has a centrum length of 1378 mm, and a functional length of 1203 mm (Figure 4-3). At 1400 mm, the longest vertebra of Sauroposeidon is marginally longer in total length [see this post for a visual comparison]. However, that length includes the prezygapophyses, which overhang the condyle, and which are missing from BYU 9024. The centrum length of the largest Sauroposeidon vertebra is about 1250 mm, and the functional length is 1190 mm. BYU 9024 therefore has the largest centrum length and functional length of any vertebra that has ever been discovered for any animal. Furthermore, the Supersaurus vertebra is much larger than the Sauroposeidon vertebrae in diameter, and it is a much more massive element overall.

Neck length estimates for Supersaurus vary depending on the taxon chosen for comparison and the serial position assumed for BYU 9024. The vertebra shares many similarities with Barosaurus that are not found in other diplodocines, including a proportionally long centrum, dual posterior centrodiapophyseal laminae, a low neural spine, and ventrolateral flanges that connect to the parapophyses (and thus might be considered posterior centroparapophyseal laminae, similar to those of Sauroposeidon). The neural spine of BYU 9024 is very low and only very slightly bifurcated at its apex. In these characters, it is most similar to C9 of Barosaurus. However, theproportions of the centrum of BYU 9024 are more similar to those of C14 of Barosaurus, which is the longest vertebra of the neck in AMNH 6341. BYU 9024 is 1.6 times as long as C14 of AMNH 6341 and 1.9 times as long as C9. If it was built like that of Barosaurus, the neck of Supersaurus was at least 13.7 meters (44.8 feet) long, and may have been as long as 16.2 meters (53.2 feet).

Based on new material from Wyoming, Lovelace et al. (2005 [published as Lovelace et al. 2008]) noted potential synapomorphies shared by Supersaurus and Apatosaurus. BYU 9024 does not closely resemble any of the cervical vertebrae of Apatosaurus. Instead of trying to assign its serial position based on morphology, I conservatively assume that it is the longest vertebra in the series if it is from an Apatosaurus-like neck. At 2.7 times longer than C11 of CM 3018, BYU 9024 implies an Apatosaurus-like neck about 13.3 meters
(43.6 feet) long.

Supersaurus vs Diplodocus BYU 9024 and USNM 10865 - Gilmore 1932 pl 6

Bonus comparo: BYU 9024 vs USNM 10865, the mounted Diplodocus longus at the Smithsonian, modified from Gilmore 1932 (plate 6). For this I scaled BYU 9024 against the 1.6-meter femur of this specimen.

If you’d like to gaze upon BYU 9024 without distraction, or put it into a composite of your own, here you go:

Supersaurus cervical BYU 9024

References

 

Check out this beautiful Lego Diplodocus:

10954093715_c4c7fe19ec_k-crop

(Click through for the full image at full size.)

I particularly like the little touch of having of bunch of Lego Victorian gentleman scientists clustered around it, though they’re probably a bit too big for the skeleton.

This is the work of MolochBaal, and all rights are reserved. You can see five more views of this model in his Flickr gallery. I especially admire how he’s managed to get the vertebral transitions pretty smooth, the careful use of separate radius/ulna and tibia/fibula, and the use of a transparent brick in the skull to represent the antorbital fenestra.

The forefeet are wrong — their toes should not be splayed out — but you can’t blame MolochBaal for that, as he was copying the mounted CM 84/94 cast in the Madrid museum.

 

I need to be sleeping, not blogging, so here are just the highlights, with no touch-ups and minimal commentary.

IMG_4487

I don’t know what these real street signs were doing sitting on the ground when I walked to the museum this morning, but it was a good omen for the conference.

IMG_4490

Home base for this part of the conference. We head to Green River, Utah, on Friday for the Early Cretaceous half.

IMG_4505

I had never seen this on exhibit. This is not the Brachiosaurus scapulocoracoid formerly referred to “Ultrasauros”, this is the other big scap from Dry Mesa, from the giant diplodocid Supersaurus.

IMG_4982

Seems legit.

IMG_4829

This is not Dinosaur Baptist Church–it is a cathedral of an entirely different order.

IMG_4545

And that order is Sauropoda.

IMG_4548

The sauropod bones are entombed in a matrix consisting of super-hard sandstone and non-sauropod bits.

I got about 150 photos of the Wall, but only because I ran out of time. You probably already know what I’m going to attempt with them. (If not, here’s a hint.)

IMG_4860

Jim Kirkland (center left) literally walked us through the Morrison and Cedar Mountain Formations at this set of exposures north of the visitor center. The reddish stuff on the lower left is Morrison, and after that it’s CMF all the way up this ridge and next two behind it.

IMG_4916

A cast of Diplodocus carnegii at the Utah Field House of Natural History State Park Museum, signalling that we’ve come to end of this tail–er, tale.

IMG_5007

Further updates as time and opportunity allow. If you tweet about the conference, please use #MMFC14!

Christine Argot of the MNHN, Paris, drew our attention to this wonderful old photo (from here, original caption reproduced below):

© Paleontological Museum, Moscow In the beginning of XX century, the Severo-Dvinskaya gallery (named after prof. Amalitsky) became the gold basis of the exhibition hall of ancient life in the Geological Museum of St-Petersburg. The museum hall was completed with a cast of the Diplodicus carnegii skeleton presented by E.Carnegy fund in 1913, at the 300-th anniversary of the Romanovs dynasty.

© Paleontological Museum, Moscow
In the beginning of XX century, the Severo-Dvinskaya gallery (named after prof. Amalitsky) became the gold basis of the exhibition hall of ancient life in the Geological Museum of St-Petersburg. The museum hall was completed with a cast of the Diplodicus carnegii skeleton presented by E.Carnegy fund in 1913, at the 300-th anniversary of the Romanovs dynasty.

I found a different version of what seems to be the same photo (greyscaled, lower resolution, but showing more of the surrounding area) here:

1932-jyosqjdogynshijh rp cpodtegqnhjimtgalwjo

What we have here is a truly bizarre mount of Diplodocus – almost certainly one of the casts of the D. carnegii holotype CM 84 — with perfectly erect, parasagittal hind-limbs, but bizarrely everted elbows.

There are a few mysteries here.

First, where and when was this photo taken? Christine’s email described this as a “picture of a Diplodocus cast taken in St. Petersburg around 1920″, and the caption above seems to confirm that location; but then why is it copyright the Paleontological Museum, Moscow? Since the web-site in question is for a Swedish museum, it’s not forthcoming.

The second photo is from the web-site of the Borisyak Paleontological Institute in Moscow, but that site unfortunately provides no caption. The juxtaposition with two more modern Diplodocus-skeleton photos that are from its own gallery perhaps suggest that the modern mount shown in the more recent photographs is a re-pose of the old mount in the black-and white photo. If so, that might mean that the skeleton was actually in Moscow all along rather than St. Petersburg, or perhaps that it was moved from St. Petersburg to Moscow and  remounted there.

Does anyone know? Has anyone out there visited the St. Petersburg museum recently and seen whether there is still a Diplodocus skeleton there? If so, is it still mounted in this bizarre way? Better yet, do you have photos?

Tornier's sprawling, disarticulated reconstruction of Diplodocus, modified from Tornier (1909, plate II).

Tornier’s sprawling, disarticulated reconstruction of Diplodocus, modified from Tornier (1909, plate II).

The second question of course is why was this posture used? This pose makes no sense for several reasons — one of which is that even if Diplodocus could attain this posture it would only serve to leave the forefeet under the torso in the same position as erect forelimbs would have them. The pose only makes any kind of sense at all if you imagine the animal lowering its torso to drink; but given that it had a flexible six-meter-long neck, that hardly seems necessary.

Of course Diplodocus does have a history of odd postures: because of the completeness of the D. carnegii holotype, it became the subject of the Sauropod Posture Wars between Tornier, Hay and Holland in the early 20th Century. Both Tornier (1909) and Hay (1910) favoured a sprawling posture like that of lizards (see images above and below), and were soundly refuted by Holland

The form and attitudes of Diplodocus. Hay (1910: plate 1)

The form and attitudes of Diplodocus. Hay (1910: plate 1)

But the Tornier and Hay postures bear no relation to that of the mounted skeleton in the photographs above: they position the forefeet far lateral to the torso, and affect the hindlimbs as well as the forelimbs. So whatever the Russian mount was doing, I don’t think it can have been intended as a representation of the Tornier/Hay hypothesis.

But it gets even weirder. Christine tells me that “I’m aware of [...] the tests that Holland performed on the Russian cast to get rid of the hypothesis suggesting a potential lizard-like posture. So I think that he would have never allowed such a posture for one of the casts he mounted himself.” Now I didn’t know that Holland had executed the mounting of this cast. Assuming that’s right, it makes it even more inexplicable that he would have allowed such a posture.

Or did he?

Christine’s email finishes by asking: “What do you think? do you think that somebody could have come behind Holland to change the position? do you know any colleague or publication who could mention this peculiar cast and comment its posture?”

Can anyone help?

References

  • Hay, Oliver. P. 1910. On the manner of locomotion of the dinosaurs, especially Diplodocus, with remarks on the origin of birds. Proceedings of the Washington Academy of Sciences 12(1):1-25.
  • Holland, W. J. 1910. A review of some recent criticisms of the restorations of sauropod dinosaurs existing in the museums of the United States, with special reference to that of Diplodocus carnegiei in the Carnegie museum. American Naturalist 44:259-283.
  • Nieuwland, Ilja. 2010. The colossal stranger. Andrew Carnegie and Diplodocus intrude European Culture, 1904–1912. Endeavour 34(2):61-68.
  • Tornier, Gustav. 1909. Wie war der Diplodocus carnegii wirklich gebaut? Sitzungsbericht der Gesellschaft naturforschender Freunde zu Berlin 4:193– 209.

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.
Follow

Get every new post delivered to your Inbox.

Join 410 other followers