Well, who knew? There I was posting images of “Pelorosaurusbecklesi‘s humerus, radius and ulna, and skin impression. There I was saying that this beast is due a proper description, and warrants its own generic name. And what should come out today but a new paper by Paul Upchurch, Phil Mannion and, oh yes, me, which does exactly that.

Screen Shot 2015-06-03 at 19.05.12

The headline news is the long-overdue establishment of a new genus name for this species — something that we’ve known was needed at least since Upchurch’s (1993) dissertation. Paul and Phil came up with the name Haestasaurus, from “Haesta”, the name of the putative pre-Roman chieftain whose people apparently settled the area of Hastings and gave the town its name. It’s nice that I can finally stop typing the scare-quotes around the no-longer-relevant old genus name “Pelorosaurus“!

Upchurch et al. 2015: figure 2. Left humerus of Haestasaurus becklesii (NHMUK R1870). A, anterior view; B, posterior view; Abbreviations: af, anconeal fossa; dp, deltopectoral crest; hh, humeral head; ltf, lateral triceps fossa; mtf, medial triceps fossa.

Upchurch et al. (2015: figure 2). Left humerus of Haestasaurus becklesii (NHMUK R1870). A, anterior view; B, posterior view; Abbreviations: af, anconeal fossa; dp, deltopectoral crest; hh, humeral head; ltf, lateral triceps fossa; mtf, medial triceps fossa.

(As you can see, the photography is rather better than in my own illustrations, which I made independently some years ago.)

Of course Paul has had an eye on this work, on and off, since the early 1990s. Then in the late 2000s, when I was working on Xenoposeidon and other Wealden sauropods, I started work independently on a redescription — which of course is why I prepared the figures that have appeared in the last few posts. But that work petered out as I started working more on other specimens and on the problems of the sauropod neck. More recently, Paul and Phil hunkered down and got the nitty-gritty descriptive work done.

Once they had a complete draft manuscript, they very graciously invited me onto the authorship — not something they had to do, but they chose to based on my previous interest in the specimen. My contribution was minor: I provided two of the illustrations, tidied up the early versions of several others, and did an editing pass on the text.

Upchurch et al. (2015: figure 1). Map showing England and Wales, with boundaries for English counties. The magnified inset shows the Isle of Wight and East and West Sussex in more detail, marking the positions of selected major towns/cities and the fossil localities mentioned in the main text. Based on

Upchurch et al. (2015: figure 1). Map showing England and Wales, with boundaries for English counties. The magnified inset shows the Isle of Wight and East and West Sussex in more detail, marking the positions of selected major towns/cities and the fossil localities mentioned in the main text. Based on “English ceremonial counties 1998″ by Dr. Greg, http://en.wikipedia.org/wiki/File:English_ceremonial_counties_1998.svg. CC By-SA 3.0.

(This map is one of the two illustrations that I provided; the other is the multi-view photograph of the Pelorosaurus conbeari humerus.)

I’m grateful to Paul and Phil, both for inviting me onto this project, and for taking into account my strong preference for an open-access venue. It’s largely because of the latter that the paper now appears in PLOS ONE, where the glorious colour illustrations appear at full resolution and may be re-used for any purpose subject to attribution.

So: what actually is Haestasaurus? Is it the early titanosaur that we’ve all been assuming? The unexciting answer is: we don’t really know. Our paper contains three phylogenetic hypotheses (all of them Paul and Phil’s work, I can’t take any credit). These results are from adding Haestasaurus to the Carballido and Sander (2014) matrix, to the Mannion et al. (2013) standard discrete matrix and to the Mannion et al. (2013) continuous-and-discrete matrix. Only the last of these recovers Haestasaurus as a titanosaur — as sister to Diamantinasaurus and then Malawisaurus, making it a lithostrotian well down inside Titanosauria.

Both both of the other analyses find Haestasaurus as a very basal macronarian — outside of Titanosauriformes. Here is the result of the analysis based on Carballido and Sander’s Europasaurus matrix:

Upchurch et al. (2105: figure 15). Strict consensus tree (CSM). A strict consensus tree based on the 28 most parsimonious trees generated by analysis of the Carballido and Sander [19] data matrix with the addition of Haestasaurus and six new characters (Tendaguria excluded a priori). GC values (multiplied by 100) are shown in square brackets for all nodes where these values are greater than 0. The monophyletic Diplodocoidea has been collapsed to a single branch in order to reduce figure size. Abbreviation: Brc, Brachiosauridae.

Upchurch et al. (2105: figure 15). Strict consensus tree (CSM). A strict consensus tree based on the 28 most parsimonious trees generated by analysis of the Carballido and Sander [19] data matrix with the addition of Haestasaurus and six new characters (Tendaguria excluded a priori). GC values (multiplied by 100) are shown in square brackets for all nodes where these values are greater than 0. The monophyletic Diplodocoidea has been collapsed to a single branch in order to reduce figure size. Abbreviation: Brc, Brachiosauridae.

As you can see, Haestasaurus is here a camarasaurid, making it (along with Camarasaurus itself) the most basal of all macronarians. In the second analysis — the one using discrete characters only from Mannion et al.’s Lusotitan paper — Haestasaurus is again in the most basal macronarian clade, but this time as sister to Janenschia and then Tehuelchesaurus. (In this topology, Camarasaurus is the next most basal macronarian after that three-taxon clade.)

So it looks like Haestasaurus is either a very basal macronarian or a pretty derived titanosaur. We don’t know which.

But, hey, at least it has a proper name now!

Acknowledgements

It’s Matt’s birthday today. I’d like to dedicate a sauropod to him, but I don’t have the authority to do that. So instead, I dedicate this blog-post to him, and declare it the Mathew J. Wedel Memorial Blog Post.

References

We’ve seen the humerus of the Wealden-supergroup putative titanosaur “Pelorosaurusbecklesi. We’ve seen the bones of the forearm, the radius and ulna. That’s it for bony remains: no other bones have been found.

But there is one other fossil that’s part of the same specimen: this skin impression:

Skin impression of

Skin impression of “Pelorosaurusbecklesii holotype NHMUK R1868. (Note that the other elements of this specimen are all catalogued as R1870.)

As you can see, the body — or at least this part of the body — was covered with roughly hexagonal tessellating (non-overlapping) scales, of about 1-2 cm diameter. But what part of the body is it from? The initial — extremely brief — description of this specimen, by Mantell (1852:143) hardly mentions the skin impression at all. All it says is:

A portion of the scaly cuirass which covered the limbs and is composed of hexagonal plates, was exhibited.

Still, this does at least suggest that the skin impression was from a limb — hardly surprising given then the left forelimb was the only part of the skeleton recovered. Upchurch et al. (2004:295) were more specific:

This skin impression was found in close association with the elbow region of the forelimb of Pelorosaurus becklesii.

But I don’t know whether this assertion is based on something written earlier, or is just a surmise.

Assuming that the skin impression is indeed from the elbow, and putting it all together, here’s what we know of “Pelorosaurus” becklesii:

Schematic of

Schematic of “Pelorosaurusbecklesi holotype NHMUK R1870, showing the preserved humerus, radius, ulna, and skin impression of the elbow. Based on Scott Hartman’s skeletal reconstruction of Alamosaurus. This is not a skeletal reconstruction of “Pelorosaurusbecklesi.

It’s not much, but it’s enough to be diagnostic.

But what actually is this beast? A titanosaur, as often assumed? A more basal macronarian? Something else entirely? Who can tell? Someone really ought to get onto that.

References

  • Mantell, Gideon A. 1852. On the structure of the Iguanodon, and on the fauna and flora of the Wealden Formation. Notices of the proceedings at the meetings of the members of the Royal Institution, with abstracts of the discourses delivered at the evening meetings 1:141-146.
  • Upchurch, Paul, Paul M. Barrett and Peter Dodson. 2004. Sauropoda. pp. 259-322 in D. B. Weishampel, P. Dodson and H. Osmólska (eds.), The Dinosauria, 2nd edition. University of California Press, Berkeley and Los Angeles. 861 pp.

Yesterday, we looked at (mostly) the humerus of the Wealden sauropod “Pelorosaurusbecklesii, which you will recall is known from humerus, radius, ulna and a skin impression, and — whatever it might be — is certainly not a species of Pelorosaurus.

Now let’s look at the radius and ulna.

Left forearm of

Left forearm of “Pelorosaurusbecklesii holotype NHMUK R1870, articulated, in anterior view, with proximal to the left: radius in front, ulna behind.

They fit together pretty neatly: the proximal part of the radius is a rounded triangular shape, and it slots into the triangular gap between the anteromedial and anterolateral processes of the proximal part of the ulna.

Left forearm of “Pelrosaurus” becklesii holotype NHMUK R1870 in proximal view, with anterior to the right. The arms of the ulna enclose the radius.

Left forearm of “Pelorosaurusbecklesii holotype NHMUK R1870 in proximal view, with anterior to the right. The “arms” of the ulna enclose the radius.

Let’s take a closer look at the ulna:

Left ulna of

Left ulna of “Pelorosaurusbecklesii holotype NHMUK R1870. Top row: proximal view, with anterior to the bottom. Middle row, from left to right: medial, anterior, lateral and posterior views. Bottom row: distal view, with anterior to top.

And the radius:

Left radius of

Left radius of “Pelorosaurus” becklesii holotype NHMUK R1870. Top row: proximal view, with anterior to the bottom. Middle row, from left to right: medial, anterior, lateral and posterior views. Bottom row: distal view, with anterior to top.

As you can see, it’s pretty well preserved: there’s no evidence of significant crushing in any of the bones, and the 3d shape is apparent.

In short, it’s a really sweet specimen. Someone really ought to get around to describing it properly, and giving it the new generic name that it clearly warrants.

It’s an oddity that in eight years of SV-POW!, we’ve never written about one of the best of all the Wealden-formation sauropod specimens: the forelimb and associated skin impression NHMUK R1870 that is known as “Pelorosaurusbecklesii.

Let’s fix that. Here is all the bony material (i.e. everything except the skin patch) in a photo taken in the basement of the Natural History Museum back in 2007:

Left forelimb material of

Left forelimb material of “Pelorosaurusbecklesii holotype NHMUK R1870. Left: humerus, in posterior view. Right, from top to bottom: ulna in anterior view; radius in anterior view. Yes, I should have turned the humerus over before taking this photo. What can I tell you? I was young and stupid then.

As you can see, the two lower-limb bones were broken back then (though I believe they have since been repaired), but the breaks are very clean, and it’s actually quite interesting to see inside the bones:

Breakage in bones of the lower left forelimb of

Breakage in bones of the lower left forelimb of “Pelorosaurusbecklesii holotype NHMUK R1870. Left: proximal part of radius in distal view. Right: proximal part of ulna in distal view.

I wish I knew enough about mineralisation to comment intelligently on what we can see there. If anyone has thoughts, do leave them in the comments.

We can look in more detail at those lower-limb bones in a subsequent post, but for now, here’s the humerus:

Pelorosaurusbecklesii holotype NHMUK R1870, left humerus. Top row: proximal view, with anterior to the bottom. Middle row, from left to right: medial, anterior, lateral and posterior views. Bottom row: distal view, with anterior to the top.

As you can see it’s in really nice shape, and pretty distinctive. Way back in my 2007 Progressive Palaeo talk (Taylor 2007), I coded up the humerus (alone, without the other elements) in the Harris-based phylogenetic matrix that I’ve used repeatedly in other projects. It came out as the sister taxon to the titanosaur Malawisaurus (which in that matrix comes out fairly basal within Titanosauria): in fact, it could hardly do anything else, since the coding was exactly the same as that of Malawisaurus.

And indeed it’s been pretty widely accepted that “P.” becklesii is a titanosaur — one of the earliest known, and the only name-bearing one from the Wealden Supergroup, unless you count the extremely indeterminate Iuticosaurus, which predictably enough is based on a single eroded partial mid-caudal centrum. Still, the titanosaurian identity of “P.” becklesii has never been convincingly demonstrated — only inferred by non-cladistic means.

Pelorosaurusbecklesii holotype NHMUK R1870, left humerus in anterodistal view (anterior to the left).

So why the quotes around the genus name “Pelorosaurus“? Because it’s long been recognised that, whatever this specimen might be, it ain’t Pelorosaurus, which is based on the Cetiosaurusbrevis caudals and a much more slender humerus.

Here’s that humerus, so you can see how different it is from that of “Pelorosaurusbecklesii:

Right humerus of Pelorosaurus conybeari holotype NHMUK 28626. Top row: distal view, anterior to bottom. Middle row, left to right: lateral, anterior and medial views. Bottom row: distal, anterior to top. Missed parts reconstructed from the humerus of Giraffatitan brancai (Janensch 1961: Beilage A)

Right humerus of Pelorosaurus conybeari holotype NHMUK 28626. Top row: distal view, with anterior to bottom. Middle row, left to right: lateral, anterior and medial views. Bottom row: distal view, with anterior to top. Missing parts reconstructed from the humerus of Giraffatitan brancai (Janensch 1961: Beilage A)

Paul Upchurch recognised the generic distinctness of “Pelorosaurusbecklesii way back in his (1993) dissertation. But because of Cambridge University’s policy of only making copies of dissertations available for £65, that work is effectively unknown. (Perhaps we should all chip in a fiver, buy a copy and “liberate” it. Or maybe 22 years on, Paul would rather leave it in obscurity and let his reputation continue to rest on his impressive body of later work.)

What has happened to this specimen in the last 22 years? Very little has been published about it. It got a mention in the systematic review of sauropods in Dinosauria II (Upchurch et al. 2004), but the only mention that is more than in passing, as far as I’m aware, is that of see Upchurch’s first published (1995) phylogenetic analysis. From page 380:

The only reliable Lower Cretaceous titanosaurid material, apart from Malawisaurus, comes from Europe, especially England. The earliest of these forms may be represented by the forelimb of ‘Pelorosaurus becklesii‘ (Mantell 1852) from the Valanginian of Sussex. This specimen was considered to be Sauropoda incertae sedis by McIntosh (1990b). However, a skin impression shows polygonal plates of a similar shape and size to those found in Saltasaurus (Bonaparte & Powell 1980). The ulna and radius are robust and the ulna bears the typical concavity on its anteromedial proximal process. Upchurch (1993) therefore argued that this form should be provisionally included within the Titanosauridae.

[Update: as Darren points out in the comment below, Upchurch et al. (2011) figure the specimen in colour and devote three pages to it. They leave it as Titanosauria, and “refrain from naming a new taxon until more comparative data are available” (p. 501).]

Given my interest in the Wealden, it’s surprising that we’ve never blogged about “Pelorosaurusbecklesii before, but it’s true: I’ve mentioned it three times in comments, but never in a post. It’s good to finally fix that!

Next time: the radius and ulna.

References

  • Janensch, Werner. 1961. Die Gliedmaszen und Gliedmaszengurtel der Sauropoden der Tendaguru-Schichten. Palaeontographica (Suppl. 7) 3:177-235.
  • Taylor, Michael P. 2007. Diversity of sauropod dinosaurs from the Lower Cretaceous Wealden Supergroup of southern England. p. 23 in Graeme T. Lloyd (ed.), Progressive Palaeontology 2007, Thursday 12th-Saturday 14th April, Department of Earth Sciences, University of Bristol. 38 pp.
  • Upchurch, Paul. 1993. The Anatomy, Phylogeny and Systematics of Sauropod Dinosaurs. Ph.D dissertation, University of Cambridge, UK. 489 pages.
  • Upchurch, Paul. 1995. The evolutionary history of sauropod dinosaurs. Philosophical Transactions of the Royal Society of London Series B, 349:365-390.
  • Upchurch, Paul, Paul M. Barrett and Peter Dodson. 2004. Sauropoda. pp. 259-322 in D. B. Weishampel, P. Dodson and H. Osmólska (eds.), The Dinosauria, 2nd edition. University of California Press, Berkeley and Los Angeles. 861 pages.
  • Upchurch, Paul, Philip D. Mannion and Paul M. Barrett. 2011. Sauropod dinosaurs. pp. 476-525 in: Batten, David J. (ed.), English Wealden Fossils. The Palaeontological Association (London).

Back when Darren and I did the Xenoposeidon description, we were young and foolish, and only illustrated the holotype vertebra NHM R2095 in four aspects: left and right lateral, anterior and posterior.  No dorsal or ventral views.

Also, because the figure was intended for Palaeontology, which prints only in greyscale, I stupidly prepared the figure in greyscale, rather than preparing it in colour and then flattening it down at the last moment.  (Happily I’d learned that lesson by the time we did our neck-posture paper: although it was destined for Acta Palaeontologia Polonica, which also prints in greyscale, and though the PDF uses greyscale figures, the online full-resolution figures are in colour.)

As if that wasn’t dumb enough, I also composited the four featured views such that the two lateral views were adjacent, and above the anterior and posterior views — so it wasn’t easy to match up features on the sides and front/back between the views.  Since then, I have landed on a better way of presenting multi-view figures, as in my much-admire’d turkey cervical and pig skull images.

So, putting it all together, here is how we should have illustrated illustrated Xenoposeidon back in 2007 (click through for high resolution):

(Top row: dorsal view, with anterior facing left; middle row, from left to right: anterior, left lateral, posterior, right lateral; bottom row, ventral view, with anterior facing left.  As always with images of NHM-owned material, this is copyright the NHM.)

Of course, if we’d published in PLoS ONE, then this high-resolution (4775 x 4095), full colour image could have been the published one rather than an afterthought on a blog somewhere.  But we didn’t: back then, we weren’t so aware of the opportunities available to us now that we live in the Shiny Digital Future.

In other news, the boys and I all registered Xbox Live accounts a few days ago.  I chose the name “Xenoposeidon”, only to find to my amazement that someone else had already registered it.  But “Brontomerus” was free, so I used that instead.

UPDATE (from Matt): I also bring good news … and bad news.

The good news is that the entire dinosaur issue of Anatomical Record is open access after all. So this post is mainly of historical interest now, and you should get on over to the page for this issue and download all the free dinosaurian goodness.

The bad news is that the representatives from Wiley never told anyone any of this when inquiries were made two weeks ago–if they had, this particular teacup could have stayed storm-free–and that they apparently still want institutions to pay $575 for a single Open Access issue of the journal. Whether those moves are predatory or just clueless, they are not earning Wiley any friends.

—————-

I bring good news … and bad news.

Good news! Tom Holtz reported in a message to the Dinosaur Mailing List that there is new issue of The Anatomical Record out that is concerned entirely with dinosaurs!  The online table of contents shows that there’s lots of good stuff.

Bad news! It’s not open access.

Good news! You can buy access to the articles.

Bad news! The price of the articles is NOT STATED.  That’s right, folks: you have to register with Wiley InterScience before they will EVEN TELL YOU THE PRICE!  Way to go, Wiley!  THAT’s the way to make sure important research is widely disseminated!

Good news! B tH wrote to ask the publisher for a price, and got a reply, which he shared in another Dinosaur Mailing List message:

Bad news! This is the reply (which I can’t format better, thanks to totally unnecessary limitations in WordPress):

Date: Mon, 31 Aug 2009 12:48:21 -0700 (PDT)

From: B tH <soylentgreenistrex@yahoo.com>

To: dinosaur@usc.edu

Subject: re: special all-dino issue

I wrote to ask them how much ordering this singl issue was – they wanted to know if I was ordering for an institution or myself. This is the price they quoted me to buy and read it at night with a flashlight under the blankey – and I am totally serious:

$575.00 US

That’s right, five HUNDRED and seventy-five buckeroos.   I assured them they were quite mad, and have to face the fact I won’t get to see it.   Waaah.

Good news! B tH realised that Wiley had quoted him the institutional rate and wrote to clarify.  The exchange is documented in yet another Dinosaur Mailing List message.

Bad news! This is the exchange:

Sent: Monday, August 31, 2009 6:07 PM

To: cs-journals@wiley.com

Subject: RE: wanting to purchase an issue of the magazine [pfCase:1078353,

pfTicket:10108736]

Um, I think you’ve made an error.

Five-Hundred and Seventy-Five dollars for an issue of a magazine?  ??

==============

From: <cs-journals@wiley.com>

Dear __________

The Anatomical Record, Volume 292, Issue 9

Thank you for your email.

As we do not have Individual rates for this title, hence the Institutional single issue rate was quoted instead.

Please provide us with a billing and shipping address if you require a proforma invoice for this order and I will happy to assist you.

Kind Regards,

Jacqueline Choong

Customer Services Advisor

Journal Customer Services for John Wiley & Sons

Good news! The revolution is coming, and things like this can only bring it on.  And Wiley’s InterScience department are a bunch of mindless jerks who will be first up against the wall when the revolution comes.

Yes, Wiley’s behaviour here is totally absurd and absolutely unethical.  No, Wiley didn’t themselves write the articles that they want to charge FIVE HUNDRED AND SEVENTY-FIVE FREAKIN’ DOLLARS for.  Neither did they pay the authors to do so.  Do you know how it comes to be that Wiley are the owners of these articles, and thus in a position to extort for access?  Happily, the reason is right here in the Instructions to Authors:

MISCELLANEOUS

[…]

Upon acceptance of an article for publication, the author will be asked to sign a Copyright Transfer Agreement transferring rights to the publisher, who reserves copyright.

Yes, it’s as simple as that.  Like all of us do most times we submit a manuscript, the authors just signed away the ownership of their work.  Just like that.  Work that was funded, if at all, by public funds, just handed over to a grossly exploitative for-profit commercial enterprise that — quite clearly, from the exchanges above — has no interest whatsoever in the advancement or dissemination of science.

Folks, we have got to stop doing this.  I can (just) stomach handing copyright of my work over to professional societies such as the Society of Vertebrate Paleontology (required for the Journal of Vertebrate Paleontology) or the Palaeontological Association (required for Palaeontology) [although frankly there is absolutely no good reason for these journals to make that requirement].  But I will NOT give my work to these parasitic commercial publishers, and I strongly urge you not to, either.  We should all of us be supporting open-access journals where possible; and failing that, at least those published by non-profit organisations.  I am not going to be propping up Elsevier, Wiley and the rest with any of my stuff.

Deep in our heart, we all — Wiley included — know that non-open academic publishing is dead, even if the corpse is still blundering around trying to eat our brains.  This sort of extortion (I mean the FIVE HUNDRED AND SEVENTY-FIVE FREAKIN’ DOLLARS kind) is death throes.  It’s probably going to get messier before the stakes are finally driven through the hearts of the bloodsuckers.  But take heart: morning is coming, and they will all turn to dust.

And finally …

More Good news! I give you NHM 46869, the holotype of Chondrosteosaurus gigas Owen 1876, a badly eroded cervical centrum from some kind of sauropod, in right lateral view:

NHM 46869, holotype of Chondrosteosaurus gigas, a cervical centrum, in right lateral view.

NHM 46869, holotype of Chondrosteosaurus gigas, a cervical centrum, in right lateral view.

This is the mate of NHM 46870, a specimen that we have already given way too much coverage, and which has sometimes been considered the cotype along with 46869.  Unlike its mate, it has not been sliced down the middle, and is — for what it’s worth — “complete” (i.e. not actually complete at all).

References

  • Owen, Richard.  1876.  Monograph of the fossil Reptilia of the Wealden and Purbeck formations.  Supplement 7.  Crocodilia (Poikilopleuron), Dinosauria (Chondrosteosaurus),  Palaeontographical Society of London [Monographs], 29:15-93.
This is the reply:
Date: Mon, 31 Aug 2009 12:48:21 -0700 (PDT)
From: B tH <soylentgreenistrex@yahoo.com>
To: dinosaur@usc.edu
Subject: re: special all-dino issue
I wrote to ask them how much ordering this singl issue was – they wanted to know if I was ordering for an institution or myself. This is the price they quoted me to buy and read it at night with a flashlight under the blankey – and I am totally serious:
$575.00 US
That’s right, five HUNDRED and seventy-five buckeroos.   I assured them they were quite mad, and have to face the fact I won’t get to see it.   Waaah.
Good news!  B tH realised that Wiley had quoted him the institutional rate and wrote to clarify.  The exchange is documented in yet another Dinosaur Mailing List message.
Bad news!  This is the exchange:
Sent: Monday, August 31, 2009 6:07 PM
To: cs-journals@wiley.com
Subject: RE: wanting to purchase an issue of the magazine [pfCase:1078353,
pfTicket:10108736]
Um, I think you’ve made an error.
Five-Hundred and Seventy-Five dollars for an issue of a magazine?  ??
==============
==============
From: <cs-journals@wiley.com>
Dear __________
The Anatomical Record, Volume 292, Issue 9
Thank you for your email.
As we do not have Individual rates for this title, hence the Institutional single issue rate was quoted instead.
Please provide us with a billing and shipping address if you require a proforma invoice for this order and I will happy to assist you.
Kind Regards,
Jacqueline Choong
Customer Services Advisor
Journal Customer Services for John Wiley & Sons
Good news!  The revolution is coming, and things like this can only bring it on.  And Wiley’s InterScience department are a bunch of mindless jerks who will be first up against the wall when the revolution comes.
Yes, Wiley’s behaviour here is totally absurd and absolutely unethical.  No, Wiley didn’t themselves write the articles that they want to charge FIVE HUNDRED AND SEVENTY-FIVE FREAKIN’ DOLLARS for.  Neither did they pay the authors to do so.  Do you know how it comes to be that Wiley are the owners of these articles, and thus in a position to extort for access?  Happily, the reason is right here in the Instructions to Authors:
MISCELLANEOUS
[…]
Upon acceptance of an article for publication, the author will be asked to sign a Copyright Transfer Agreement transferring rights to the publisher, who reserves copyright.
###

Condrosteo_scan

By now you’ll recognize this as NHM 46870, a minor celebrity in the world of pneumatic sauropod vertebrae. Darren has covered the history of the specimen before, and in the last post he showed photographs of both this chunk and its other half. He also briefly discussed the Air Space Proportion (ASP) of the specimen, and I’ll expand on that now.

People have mentioned the weight-saving properties of sauropod vertebrae from the very earliest discoveries of sauropods. But as far as I know, no one tried to quantify just how light they might have been until 2003.

That fall I was starting my third year of PhD work at Berkeley, and I was trying to think of everything that could possibly be investigated about pneumaticity in sauropod vertebrae. I came up with a list of four things:

  • external traces of pneumaticity (foramina, fossae, tracks, laminae)
  • form and complexity of internal spaces (camerae, camellae, branching patterns)
  • ratio of bone to air space within a pneumatic element
  • distribution of postcranial skeletal pneumaticity (PSP) in the body

That list of four things formed the outline for my first dissertation chapter (Wedel 2005), and for my dissertation itself. In fact, all of my papers that have anything to do with pneumaticity can be classified into one or more of those four bins:

That list is not exhaustive. It’s every aspect of PSP that I was able to think of back in 2003, but there are lots more. For example, I’ve only ever dealt with the internal complexity of sauropod vertebrae in a qualitative fashion, but the interconnections among either chambers or bony septa could be quantified, as Andy Farke has done for the frontal sinuses of hartebeests (Farke 2007). External traces on vertebrae and the distribution of PSP in the body can also be quantified, and were shortly after I drew up the list–see Naish et al. (2004) for a simple, straightforward approach to quantifying the extent of external pneumatic fossae, and O’Connor (2004, 2009) for a quantitative approach to the extent of pneumaticity in the postcranial skeletons of birds. There are undoubtedly still more parameters waiting to be thought of and measured. All of these papers are first steps, at least as applied to pneumaticity, and our work here is really just beginning.

Also, it took me an embarrassingly long time to “discover” ASPs.  I’d had CT slices of sauropod vertebrae since January, 1998, and it took me almost six years to realize that I could use them to quantify the amount of air inside the bones. I later discovered that Currey and Alexander (1985) and Casinos and Cubo (2000) had done related but not identical work on quantifying the wall thickness of tubular bones, and I was able to translate their results into ASPs (and MSPs for marrow-filled bones).

Condrosteo_ASP

The procedure is pretty simple, as Mike has shown here before. Open up the image of interest in Photoshop (or GIMP if you’re all open-sourcey, like we are), make the bone one color, the air space a second color, and the background a third color. Count pixels, plug ’em into a simple formula, and you’ve got the ASP. I always colored the bone black, the air space white, and the background gray, so

ASP = (white pixels)/(black + white pixels)

For the image above, that’s 460442/657417 = 0.70.

Two quick technical points. First, most images are not just black, white, and one value of gray. Because of anti-aliasing, each black/white boundary is microscopically blurred by a fuzz of pixels of intermediate value. I could have used some kind of leveling threshold thing to bin those intermediate pixels into the bone/air/background columns, but I wanted to keep the process as fast and non-subjective as possible, so I didn’t. My spreadsheet has columns for black, white, gray, and everything else. The everything else typically runs 1-3%, which is not enough to make a difference at the coarse level of analysis I’m currently stuck with.

Second, I prefer transverse sections to longitudinal, because most of the internal chambers are longitudinally oriented. That means that longitudinal sections, whether sagittal or horizontal, are likely to cut through a chamber wall on its long axis, which makes the walls look unnaturally thick. For example, in the image above the median septum looks 5-10 times thicker than the outer walls of the bone, which would be a first–usually the outer walls are thicker than the internal septa, as you can see here. I don’t think the median septum really is that thick; I strongly suspect that a very thin plate of bone just happened to lie in the plane of the cut. It takes some work to get used to thinking about how a 2D slice can misrepresent 3D reality. When I first started CT scanning I was blown away by how thick the bone is below the pre- and postzygapophyses. I was thinking, “Wow, those centrozygapophyseal laminae must have been way more mechanically important than anyone thinks!” It took me a LONG time to figure out that if you take a transverse slice through a vertical plate of bone, it is going to look solid all the way up, even if that plate of bone is very thin.

Even apart from those considerations, there is still a list of caveats here as long as your arm. You may not get to choose your slice. That’s almost always true of broken or historically sectioned material, like NHM 46870. It’s even true in some cases for CT scans, because some areas don’t turn out very clearly, because of mineral inclusions, beam-hardening artifacts, or just poor preservation.

The slice you get, chosen or not, may not be representative of the ASP of the vertebra it’s from. Even if it is, other elements in the same animal may have different ASPs. Then there’s variation: intraspecific, ontogenetic, etc. So you have to treat the results with caution.

Still, there are some regularities in the data. From my own work, the mean of all ASP measurements for all sauropods is about 0.60. That was true when I had only crunched my first six images, late on the evening of October 9, 2003. It was true of the 22 measurements I had for Wedel (2005), and now that I have over a hundred measurements, it’s still true. More data is not shifting that number at all. And Woodward (2005) and Schwartz and Fritsch (2006) got very similar numbers, using different specimens.

This is cool for several reasons. It’s always nice when results are replicated–it decreases the likelihood that they’re a fluke, and in this case it suggests that although the limitations listed above are certainly real, they are not deal-killers for answering broad questions (we are at this point seeing the forest more clearly than the trees, though).

More importantly, the mean 0.60 ASP for all sauropod vertebrae is very similar to the numbers that you get from the data of Currey and Alexander (1985) and Cubo and Casinos (2000): 0.64 and 0.59, respectively. So sauropod vertebrae were about as lightly built as the pneumatic long bones of birds, on average.

Naturally, there are some deviations from average. Although I didn’t have enough data to show it in 2005, brachiosaurids tend to have higher ASPs than non-brachiosaurids. And Early Cretaceous brachiosaurids from the US and England are especially pneumatic–the mean for all of them, including Sauroposeidon, ‘Angloposeidon’, some shards of excellence from the Isle of Wight, and assorted odds and ends, is something like 0.75-0.80, higher even than Brachiosaurus. So there’s probably a combined phylogenetic/functional story in there about the highly pneumatic, hyper-long-necked brachiosaurids of the Early Cretaceous of Laurasia. Another paper waiting to be written.

Chondrosteosaurus broken face

Here’s another shard of excellence, referred to Chondrosteosaurus, NHM R96. As Mike had discussed here before, there’s no good reason to believe that it actually is Chondrosteosaurus, and the internal structure looks considerably more subdivided than in NHM 46870. This is an anterior view, and normally you’d be seeing a nice hemispherical condyle, but all of the cortical bone is gone and the internal structure is revealed. The little black traces are bone and the brownish stuff is rock matrix filling the pneumatic cavities.

Chondrosteosaurus broken face ASP

A few years ago, Mike asked me to look at that photo and guess the ASP, and then run the numbers and see how close I got. I guessed about 78%, then did the calculation, and lo and behold, the answer was 78%. So I’m pretty good at guessing ASPs.

Except I’m not, because as any of you armed with photo software can tell, that picture has 24520 black pixels and 128152 white ones, so the ASP is actually 128152/(128152+24520) = 0.84. The moral of the story is check your homework, kids! Especially if you seem to be an unnaturally good estimator.

ASP-ESP aside, I think ASP is cool and has some interesting potential at the intersection of phylogeny and biomechanics. But the method is severely limited by sample size, which is severely limited by how much of a pain in the butt preparing the images is. In most cases you can’t just play with levels or curves to get a black and white image that faithfully represents the morphology, or use the magic wand, or any of the other myriad shortcuts that modern imaging programs offer. Believe me, I’ve tried. Hard. But inevitably you get some matrix with the bone, or some bone with the matrix, and you end up spending an impossible amount of time fixing those problems (note that this is not a problem if you use perfect bones from extant animals, which is sadly not an option for sauropod workers). So almost all of my ASP images were traced by hand, which is really time-consuming. I could pile up a lot more data if I just sat around for a few weeks processing images, but every time I’ve gotten a few free weeks there has been something more important demanding my attention, and that may always be the case. Fortunately I’m not the only one doing this stuff now, and hopefully in the next few years we’ll get beyond these first few tottering steps.

Side Note: Does NHM 46870 represent a juvenile, or a dwarf?

This came up amongst the SV-POW!sketeers and we decided it should be addressed here. Darren noted that the vert at top is pretty darned small, ~23 cm for the preserved part and probably only a foot and a half long when it was complete, which is big for an animal but small for a sauropod and dinky for a brachiosaurid (if that’s what it is). Mike made the counter-observation that the internal structure is pretty complex, citing Wedel (2003b:fig. 12) and surrounding text, and suggested that it might be an adult of a small or even dwarfed taxon. And I responded:

I’m not at all certain that it is dwarfed. It matters a lot whether the complex internal structure is polycamerate or camellate. I was agnostic for a long time about how different those two conditions are, but there is an important difference that is relevant in this case: the two internal structures develop differently. Polycamerate verts really do get progressively more complex through development, as illustrated–there are at least two great series that show this, that I need to publish one of these days. But I think camellate vertebrae may be natively complex right from the get-go; i.e., instead of a big simple diverticulum pushing in from the side and making a big camera first, a bunch of smaller diverticula may remodel the small marrow spaces into small air spaces with no prior big cavities. At least, that’s how birds seem to do it. This needs more testing from sauropods–a good ontogenetic sequence from Brachiosaurus would be clutch here–but it’s my working hypothesis. In which case NHM 46870 may be a juvenile of a camellate taxon, rather than an adult of a polycamerate taxon.

The whole camerate-vs-camellate problem deserves a post of its own, and this post is already too long, so we’ll save that for another day.

References

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