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

It’s no secret – at least, not if you’re a regular SV-POW! reader – that the Lower Cretaceous Wealden Supergroup of southern England includes more than its fair share of enigmatic sauropod remains (see Mystery sauropod dorsals of the Wealden part 1, part 2, part 3). Poor taxonomic decisions, a dearth of adequate descriptive literature, and (perhaps) the vague concept that sauropod diversity in the Lower Cretaceous of Europe must be low have combined to prevent adequate appraisal. Recent comments on Wealden sauropods have been provided by Naish et al. (2004), Naish & Martill (2007), Taylor & Naish (2007) and Mannion (2008).

Chondrosteosaurus_R46870_resized

One of the most interesting Wealden sauropods – and I mean ‘interesting’ in an entirely subjective, historiographical sense – is Chondrosteosaurus gigas. This taxon has a rather confusing history that I don’t want to repeat here. The type series consists of two cervical vertebrae: BMNH R46869 and BMNH R46870 (and it is BMNH R46870, despite the occasional use in the literature of ’46780′). We’ve looked at BMNH R46869 before. This time round I want to briefly talk about BMNH R46870. Anyone familiar with the literature on Wealden sauropods will know that this specimen was sectioned and polished. However, to date, only half of BMNH R46870 has been published (Owen 1876, plate V; Naish & Martill 2001, text-fig. 8.4), on both occasions as a mirror-image of the actual specimen. Previously unreported is that both halves of the specimen were polished, and both are in the Natural History Museum’s collection today. And here they are, shown together for the first time ever. I screwed up on the lighting, so sorry for the poor image quality [images © Natural History Museum, London].

A little bit of science has been done on this specimen. Chondrosteosaurus has had a mildly controversial history: it’s been suggested at times to be a camarasaur, but its camellate interior show that it’s a titanosauriform. Because the exact ratio of bone to air can be measured, the specimen lends itself particularly well to an Air Space Proportion analysis of the sort invented by Matt. Indeed, Matt did some ASP work on the figured half of BMNH R46870 in his thesis, finding an ASP of 0.70 (Wedel, Phd thesis, 2007). The average ASP of sampled neosauropod vertebrae is 0.61, and an ASP of 0.70 for the mid-centrum (as opposed to the condyle or cotyle) is most similar to the values present in camarasaurs and brachiosaurs. Mid-centrum ASP values of titanosaurs seem to be lower (Wedel, Phd thesis, 2007).

Chondrosteosaurus_R46870_closeup_resized

Anyway, more on Wealden sauropods – hopefully, a lot more – in the future.

References

  • Mannion, P. 2008. A rebbachisaurid sauropod from the Lower Cretaceous of the Isle of Wight, England. Cretaceous Research 30, 521-526.
  • Naish, D. & Martill, D. M. 2001. Saurischian dinosaurs 1: Sauropods. In Martill, D. M. & Naish, D. (eds) Dinosaurs of the Isle of Wight. The Palaeontological Association (London), pp. 185-241.
  • Naish, D. & Martill, D. M. 2007. Dinosaurs of Great Britain and the role of the Geological Society of London in their discovery: basal Dinosauria and Saurischia. Journal of the Geological Society, London, 164, 493-510.
  • Naish, D., Martill, D. M., Cooper, D. & Stevens, K. A. 2004. Europe’s largest dinosaur? A giant brachiosaurid cervical vertebra from the Wessex Formation (Early Cretaceous) of southern England. Cretaceous Research 25, 787-795.
  • Owen, R. 1876. Monograph on the fossil Reptilia of the Wealden and Purbeck Formations. Supplement 7. Crocodilia (Poikilopleuron). Dinosauria (Chondrosteosaurus). Palaeontographical Society Monographs, 30, 1-7.
  • Taylor, M. P. & Naish, D. 2007. An unusual new neosauropod dinosaur from the Lower Cretaceous Hastings Beds Group of East Sussex, England. Palaeontology 50, 1547-1564.

We have sometimes neglected tails on SV-POW!, in favour of the more obviously charismatic charms of presacral vertebrae, but every now and then you come across a caudal vertebra so bizarre that it just cries out to be blogged.

One such is this specimen, which may or may not be BMNH R 2144:

Sauropod caudal with co-ossified chevrons, lateral view

Sauropod caudal with co-ossified chevrons, right lateral view

Sauropod caudal with co-ossified chevrons, right posterolateral view

Sauropod caudal with co-ossified chevrons, right posterolateral view

The reason I’m not sure whether this is BMNH R2144 is that I noticed this at the very last minute while visiting the NHM collections to see a different specimen, and just had time to take a couple of quick photos before kicking-out time.  The label on the side of the vertebra has the unexplained number 2144 written on it, so I am guessing this is the specimen number, but I wouldn’t stake my life on it.

(By the way, both these photographs are copyright the NHM.)

The interesting thing about this vertebra is of course that that the chevrons are co-ossified with the centrum — an extremely rare condition in sauropods, in fact unique as far as I know.  As we’ve shown here and here, among other places, the chevrons are usually separate bones from the vertebrae.

This vertebra caught my eye not only because it’s, well, weird, but also because I’d seen it a couple of times in published figures.  It’s in Mantell’s (1850) description of Pelorosaurus, where it appears as figure 11 in plate XXIII, and is considered to belong to Pelorosaurus; and also in Owen (1859: plate V: figs. 3-4).  Owen seems pretty confused about the identity of this element, and in this paper alone assigns it to Streptospondylus (p. 22), Iguanodon(!) (p. 25) and implicitly Cetiosaurus (p. 34).  So what is it?  Well, its provenance is vague in the extreme, so given that it’s not associated with any more diagnostic material, about the best we can say with any honesty is that it’s Sauropoda incertae sedis.

Let’s take a look at those old figures:

Mantell (1850: plate XXIII, fig. 11)

Mantell (1850: plate XXIII, fig. 11)

Owen (1850: plate V, figs 3-4)

Owen (1850: plate V, figs 3-4)

If you’re like me, your first thought was that Owen’s figures are simply mirror images of Mantell’s.  I checked this out by Photoshopping the two sets of figures, flipping them horizontally, scaling and rotating as necessary, and found to my mild surprise that Owen’s figures are in fact redrawn, despite the startling resemblance they bear to Mantell’s.  As it happens, the same is true with the Owen 1859 plate that is the humerus of Pelorosaurus figured by Mantell 1850, and in that case Owen’s figure is rather better than Mantell’s, so let’s give a bit of credit to Owen here.  Most embarrassing for Mantell (not that he cares, having been dead for 157 years) is that Owen’s flipped images seem to be correct (at least, as best I can judge from the photographs I took) — looks like Mantell or his illustrator badgered this up.

So what is going on with these co-ossified chevrons?  As is so often the case, we just don’t know.  Some possibilities: this might be a pathology of an individual, caused either by injury or infection; it might be a natural ontogenetic character in very old individuals; or it might by a taxonomically significant character of a taxon we’ve not yet found — or one that we have found, but don’t yet recognise as being the same thing.  It’s perfectly possible that this is a chevron of Xenoposeidon, for example, but until someone finds a nice complete specimen we’ll never know.

Not much is known about skeleton fusion in sauropods, and most of what’s in the literature is anecdote.  That is set to change, I am pleased to say, as Matt is putting together a paper with his colleague Elizabeth Rega that will survey and interpret the various fusions known in sauropod vertebrae.  I’m looking forward to seeing what they have to say about this vertebra.

References

  • Brusatte, Stephen L., Roger B. J. Benson, and Stephen Hutt.  2008.  The osteology of Neovenator salerii (Dinosauria: Theropoda) from the Wealden Group (Barremian) of the Isle of Wight.  Monograph of the Palaeontographical Society 162 (631): 1-166.
  • Calvo, Jorge O., Juan D. Porfiri, Claudio Veralli, Fernando Novas and Federico Poblete.  2004.  Phylogenetic status of Megaraptor namunhuaiquii Novas based on a new specimen from Neuquen, Patagonia, Argentina.  Ameghiniana 41 (4): 565-575.
  • Mantell, Gideon Algernon.  1850.  On the Pelorosaurus: an undescribed gigantic terrestrial reptile, whose remains are associated with those of the Iguanodon and other saurians in the strata of Tilgate Forest, in Sussex.  Philosophical Transactions of the Royal Society of London 140: 379-390.
  • Owen, R.  1859a.  Monograph on the fossil Reptilia of the Wealden and Purbeck formations.  Supplement no. II (pages 20-44 and plates V-XII): Crocodilia (Streptospondylus, &c.) [Wealden].  Palaeontographical Society, London.

Update

Thanks to Mickey Mortimer for pointing out that this kind of centrum-chevron fusion is known in the theropod Megaraptor.  Here is the relevant figure from Calvo et al.’s (2004) revision of that genus:

Calvo et al. (2004: fig. 5). Caudal vertebrae of Megaraptor with co-ossified chevron

Calvo et al. (2004: fig. 5). Caudal vertebrae of Megaraptor with co-ossified chevron

The strange thing is this comment in the text (p. 569): “Two articulated caudal vertebrae are preserved (figure 5), slightly laterally compressed.  Their centra and the neural arches are firmly co-ossified, as well as their respective haemal arches [i.e. chevrons].  This fusion, not infrequent among dinosaurs, may be pathological.”  Not infrequent?  Is this going on all over the place and I’ve just never noticed it?  Anyone have any more examples?

Update 2

Here is that pair of fused Neovenator caudals with a co-ossified chevron, which Darren mentions in the comments below.

Brusatte et al. (2008: fig. 16d), fused Neovenator caudals with co-ossified chevron

Brusatte et al. (2008: fig. 16d), fused Neovenator caudals with co-ossified chevron

It’s been a while since we looked at everybody’s favourite partial dorsal vertebra, and there may be those who feel we’ve said all that can be said about it, but there is one feature of Xenoposeidon that we’ve never really highlighted here and which is well worth a look.

For anyone who’s not up to speed, a super-brief resumé: Xeno is an indeterminate neosauropod which Darren and I named in 2007 on the basis of a single element, a superbly preserved partial dorsal vertebra loaded with distinctive features that make it very clearly distinct from any other named taxon.  For anyone who wants more background, the original paper is freely available, as is a page summarising the story for the media, some unnofficial supplementary information, and a whole week’s worth of SV-POW! posts.

Here is the canoncial Xenoposeidon photo: the specimen in left lateral view.  (Don’t worry, this old chestnut is not your Picture of the Week — it’s just setting the scene.)

Xenoposeidon holotype BMNH R1095, left lateral view

Xenoposeidon holotype BMNH R1095, left lateral view. Copyright the NHM.

Up near the top of the preserved part of the vertebra, where the neural arch is broken off, there is a distinctive “V”-shaped pair of laminae, which we identified as accessory infraparapophyseal and infrapostzygapophyseal laminae. The more posterior of the two (on the right as we look at at it here) has a very distinctive wrinkled texture, and that’s what I want to show you today:

Xenoposeidon holotype BMNH R2095, left lateral view. Detail of accessory infrapostzygapophyseal lamina showing wrinkled bone texture

Xenoposeidon holotype BMNH R2095, left lateral view. Detail of accessory infrapostzygapophyseal lamina showing wrinkled bone texture. Copyright the NHM.

Now I’ve heard different things said of those wrinkles: I’ve never seen them on any other specimen of anything before, but then I’ve not seen a whole lot of material. Other people have told me that this kind of thing is pretty common, though I don’t recall anyone ever having told me a specific other specimen that has it.  What is completely clear is that no-one seems to know what it is or what it means.  It’s very hard to see how this could be the result of any kind of post-mortem distortion, so this must be what the bone was like when the animal was alive.  What are the possibilities?

  • I wondered whether it could be some kind of pneumatic feature, perhaps the trace left by a diverticulum or set of diverticula; but IIRC Matt doesn’t think that’s likely.
  • Could it be the result of some sort of infection?  Maybe neontologists, or for that matter doctors, see this kind of thing all the time but we poor palaeontologists are ignorant.
  • Er … what else?  Seriously, I am out of ideas here.

So, comments are open.  Enlighten me.  What is the cause of this distinctive texture?

Oh, and finally — for some reason, I don’t think I’ve ever mentioned that I uploaded most of the Xenoposeidon news spots onto Youtube a while back.  So if anyone wants to hear me talking about my favourite subject, and arguing with the Channel 4 News guy, or indeed just wants to see how mainstream media can get a science story completely right when we trouble to give them good information, get over to http://www.youtube.com/user/Mirk101

Happy Xenoposeidon day!  Today, November 15, 2008, is the one-year anniversary of the publication of Xenoposeidon Taylor and Naish 2007.

By happy coincidence, I’ve just been sent a courtesy copy of Kids Only, a new guide-book for the Natural History Museum … and there is Xenoposeidon on page 5, exemplifying dinosaur diversity.  Rock!

nhm-kids-only-p5-xenoposeidon-500px

It’s good to see our baby out there educating people!

For much more of Xeno, see Xenoposeidon week.

Today, we bring you the long-overdue third installment in everyone’s  favourite Mystery Sauropod Dorsals serial, our trawl through the NHM’s collection of mostly isolated elements from the Wealden Supergroup.

Many of these elements are too bashed up to be diagnostic (with the Xenoposeidon holotype R2095 being an honourable exception).  But there are one or two that are much better preserved, and arguably the best of these are the pair of elements BMNH R88/R89, which in some sense belong to “Eucamerotus” (read on).  These are difficult to photograph well, because they are in a glass case in the public gallery, but fortunately Hulke (1880: plate IV) illustrated the more anterior and better preserved of the two:

plate IV)

Like far too many British sauropod specimens, this one is mired in a taxonomic hell-hole. It was described by Hulke as belonging to Ornithopsis, a genus based on a horribly non-diagnostic type specimen, and it is this name that appears on the exhibit label (along with the incorrect specimen numbers R89/90 … oh well, One Out Of Two Ain’t Bad.)

Here is my least bad photo of R88 and R89, in left lateral view, with R88 on the left:

dscn1431-500px

Blows (1995) referred this pair of dorsals, and a bunch of other specimens, to another ancient British name, Eucamerotus — in fact, he nominated them as paratypes — but didn’t give any reason for doing so.  (He also referred to the R88/R89 pair jointly as R90, thus further muddling the specimen numbering.)  Blows’s reassignment to Eucamerotus is puzzling, because while the Eucamerotus type specimen is also pretty undiagnostic, consisting only of a partial neural arch, it does have one obvious apomophy, which is huge robust parapophyses supported on what I like the call The Prezygaparapophyseal Laminae Of Doom.  (Remind me to show you this specimen some time.)  That feature, of course, R88 and R89 completely lack.

So what are they?  I don’t think they can be referred with any confidence whatsoever to either Ornithopsis or Eucamerotus, two questionable genera of which at least the first is invalid.  So perhaps the right thing to do would be to torpedo those names and raise R88/89 as the type specimen of a new taxon?  There’s more work to do before taking such a step, not least an exhaustive trawl through the historical literature, but I think that might eventually prove the way to go.

Based on general proportions and overall “gestalt”, these vertebrae appear to be brachiosaurid — but I put them in a cladistic analysis a while back (so far unpublished) and they didn’t clade unambiguously with Brachiosaurus, so we’ll have to see how that develops when I finally get around to adding my thirty-odd new characters of the dorsals.  Don’t hold your breath.  At least, these elements are much more convincingly brachiosaurid than anything else I’ve seen from the Wealden.  So do they consitute Britain’s best brachiosaur?

Well, maybe.  Not if you count The Archbishop, Migeod’s Tendaguru brachiosaur, which I’ve been working on for waaay too long now, but really, really will describe Real Soon Now.  (Amazingly, this specimen has yet to appear on SV-POW!, unless you count my T-shirt in one of the photos of our Oxford Museum visit.)  But since this specimen is from the Tendaguru Formation of Tanzania, it should probably be discounted from the BBB competition.

In fact, Britain’s Best Brachiosaur is probably the “Barnes High Sauropod” from the Isle of Wight. But that’s in private hands and the ownership/availability situation is complex.  For that reason, no-one has yet published on it, and in fact I have never seen any of the material except what’s embedded in a wall-mount at Dinosaur Isle.  I’m not sure what’s happening with this specimen (I don’t think anyone is) but if I ever get a chance to find out, I will!

References

  • Blows, William T.  1995.  The Early Cretaceous brachiosaurid
    dinosaurs Ornithopsis and Eucamerotus from the Isle of
    Wight, England.  Palaeontology 38 (1): 187-197.
  • Hulke, J. W.  1880.  Supplementary Note on the Vertebræ of Ornithopsis, Seeley, = Eucamerotous, Hulke.  Quarterly Journal of the Geological Society 36: 31-35.  doi:10.1144/GSL.JGS.1880.036.01-04.06

In this article I’d like look at something that I’ve just spoken about at a conference: the ‘Dinosaurs – A Historical Perspective’ meeting held in London on May 6th and 7th (my thoughts on the conference can be found here and here). Mike attended too (and, like me, gave a talk), but Matt couldn’t make it. Anyway…

 

 

Today, the idea that sauropods (and non-avian theropods) were pneumatic animals is well established and universally accepted (a minority view – promoted by those who insist that birds cannot be dinosaurs – maintains that non-avian dinosaurs were not pneumatic, but I see no indication that the workers concerned know what they’re talking about). Indeed, sauropod pneumaticity has been discussed a lot here at SV-POW! But have people always regarded sauropods as pneumatic? As someone particularly interested both in pneumaticity and in the dinosaurs of the Lower Cretaceous Wealden Supergroup of southern England, the whole ‘Pneumaticity: the Early Years’ story is of special interest to me. I hope you get something out of it too…

 

Actually, the very first dinosaur (not just sauropod, but dinosaur) identified as exhibiting skeletal pneumaticity is from the Wealden, and it’s a theropod: it’s the large tetanuran Becklespinax altispinax, discovered prior to 1855 in the Hastings Beds Group of East Sussex (the Hastings Beds Group is the oldest unit in the Wealden Supergroup: for more see the Tet Zoo article here). Consisting only of three articulated posterior dorsal vertebrae, Becklespinax exhibits deep fossae on the sides of the neural arches (by now you’ll all be familiar with the names for the different vertebral laminae present in saurischians, but the fossae have names too: in Becklespinax, we’re talking about the infraprezygapophyseal fossa, the infradiapophyseal fossa, and the infrapostzygapophyseal fossa). The key thing is that, in describing these vertebrae, Richard Own (1804-1892) realised that these fossae were probably pneumatic as they are in birds, and in his 1856 article on Megalosaurus (he assumed that the Becklespinax vertebrae belonged to Megalosaurus), he wrote that ‘Three deep depressions, probably receiving parts of the lungs in the living animal, divide these lamelliform butresses from each other’ (Owen 1856, p. 5). His ‘lamelliform butresses’ are what we call laminae. This is the very first reference to pneumaticity in any Mesozoic dinosaur (Britt 1993), so Becklespinax is the first non-avian dinosaur for which pneumaticity was ever suggested.

 

The next milestone in pneumaticity came from Harry Seeley (1839-1909) in his description of the Wealden sauropod Ornithopsis hulkei (Seeley 1870). O. hulkei was named for two dorsal vertebrae: BMNH R2239 from East Sussex and BMNH R28632 from the Isle of Wight, but the former was later removed from O. hulkei (then becoming the type for Bothriospondylus elongatus), leaving BMNH R28632 alone associated with this name and as the lectotype* [the specimen is shown below, from Seeley 1870]. The strong opisthocoely, large lateral foramina and camellate internal anatomy show that BMNH R28632 is from a titanosauriform (it can’t really be identified more precisely than that and whether it’s diagnostic is arguable: see Naish & Martill 2007. For previous SV-POW! comments on the specimen go here). It’s often noted that Seeley suggested that these vertebrae belonged to an animal ‘of the Pterodactyle kind’. However, he did not think that these vertebrae belonged to a giant pterosaur (as, oops, Naish & Martill (2001) said): rather, he thought that O. hulkei represented something entirely new, the first member of a ‘new order of animals which will bridge over something of the interval between birds and Pterodactyles, and probably manifest some affinity with the Dinosaurs’ (Seeley 1870, p. 280).

 

* When a species is erected for more than one specimen, the specimens are all called syntypes. When one syntype from a series is later elected to serve as the type specimen for the species, it is called the lectotype.

 

 

 

Seeley – who has been described as ‘the most defiant’ of Victorian palaeontologists, of exhibiting ‘anarchic tendencies’, and of being considered ‘strikingly individualistic’, even in his own day (Desmond 1982) – gets a lot of flack these days, in particular for his rampant taxonomic splitting and naming of new dinosaur and pterosaur species, and also for his unusual views on how vertebrate groups were related to one another. But I often think that his conclusions on lifestyles and comments on palaeobiology are really not unreasonable in view of what we think today, and in fact often seem quite far-sighted.

 

Here’s where we come back to pneumaticity: Seeley (1870) was very impressed with the enormous lateral foramina present in O. hulkei (these were the main feature which led him to regard O. hulkei as allied to pterosaurs and birds), and he wrote: ‘Seeing that in living animals these foramina exist for the prolongation of the peculiarly avian respiratory system into the bones, and that no other function is known for them, we are compelled to infer for this animal bird-like heart and lungs and brain’ (Seeley 1870, p. 280). In describing the worn condyle of BMNH R28632, Seeley noted the presence within the bone of ‘enormous honeycomb-like cells of irregular polygonal form … divided by exceedingly thin and compact films of bone’ (Seeley 1890, p. 281). Elsewhere in the paper, he referred to the internal cavities as ‘air-cells’, and he also wrote of the Ornithopsis vertebrae (both BMNH R2239 and BMNH R28632) as ‘being constructed after the lightest and airiest plan’. He never explicitly stated it, but I infer from these statements that Seeley imagined the internal cavities of the centrum to be pneumatic: he was describing what today we call the camellae (that is, the numerous small pneumatic cavities that occupy the centrum in mamenchisaurs and titanosauriforms). So, all in all, I say well done Seeley for correcting inferring vertebral pneumaticity in O. hulkei.

 

Like most Victorian palaeontologists, Seeley did not get on particularly well with the most prolific, most respected and most politically powerful Victorian palaeontologist, Richard Owen [although it’s not really accurate to regard Owen just as a palaeontologist: sure, he did palaeontology, but his contribution to mainstream zoology and anatomy was massive and equally significant, if not more so]. In 1876, Owen described another Wealden sauropod and, like the O. hulkei lectotype (BMNH R28632), it was from the Isle of Wight’s Wessex Formation: it’s based on two cervical vertebrae (BMNH R46869 and BMNH R46870: one is shown below). Today, it is obvious that these vertebrae are from sauropods, and their enormous lateral foramina and camellate internal anatomy show that they’re from titanosauriforms (and not from camarasaurs as has been suggested in the past: see Naish & Martill 2001, 2007). However, Owen couldn’t be this confident that identified the material as ‘Dinosauria (?)’.

 

 

 

The big deal for our story is that these vertebrae have massive lateral fossae housing large lateral foramina, and again Owen correctly interpreted them as pneumatic, writing: ‘The whole of the side of the centrum is occupied by a deep oblong depression which, probably, lodged a saccular process of the lung’ (Owen 1876, p. 6). So far so good. But Owen had one of these two Wessex Formation specimens (BMNH R46870) sectioned, revealing its incredible (and beautiful) interior [it’s the image shown at the very top]. The internal anatomy of this specimen illustrates camellate texture so well that it’s almost becoming a poster-child in talks and articles on sauropod vertebrae. But while, as we just saw, Seeley had implied that camellae were pneumatic, Owen interpreted them quite differently. He wrote ‘I deem it much more probable that the large cancelli obvious at every fractured surface of this vertebra were occupied in the living reptile by unossified cartilage, or chondrine, than by air from the lungs, and consequently have no grounds for inferring that the whale-like Saurian, of which the present vertebrae equals in length the largest one of any Cetacean recent or fossil, had the power of flight, or belonged to either Pterosauria or Aves’ (Owen 1876, p. 6). To reflect the presence of ‘chondrine-filled’ spaces in the vertebrae of this animal, Owen coined for it the new name Chondrosteosaurus gigas, meaning ‘giant cartilage and bone lizard’.

 

Quite why Owen was happy with pneumatic lateral fossae, but not with pneumaticity within the body of the centrum itself, seems odd, especially when Owen was very familiar with avian anatomy (he specifically referred to the internal anatomy of avian vertebrae in, for example, his 1859 article on pterosaur vertebrae [Owen 1859]). As you can see in Tutorial 3, the internal anatomy of bird and sauropod centra are so similar that it is difficult not to conclude that what applies for one applies for the other. However, it is clear from Owen’s quote given above that, when interpreting C. gigas, he was not just doing an objective description, but also had an axe to grind: he was specifically refuting Seeley’s statements on O. hulkei, hence the rejection of the idea that C. gigas might have been capable of flight, or that it might be allied to pterosaurs or birds. For whatever reason, Owen was also making note of the fact that he disagreed with Seeley’s idea of a pneumatic vertebral interior: the name Chondrosteosaurus itself almost seems like a snub to Seeley.

 

Despite this one-upmanship, ultimately, both Seeley and Owen come out of this early phase in research pretty well, as both workers still win citations for having made key early statements on saurischian pneumaticity (e.g., Wedel 2003, O’Connor 2006). Edward Cope (1840-1897), who liked Seeley and said nice things about him, was to note during the 1870s that sauropod vertebrae were probably pneumatic, and Othniel Marsh (1831-1899) did likewise. What happened in the field of dinosaur pneumaticity after this? Well, that’s a story for another time: for the time being I will direct you to Wedel (2003).

 

References

 

  • Britt, B. 1993. Pneumatic Postcranial Bones in Dinosaurs and Other Archosaurs. Unpublished PhD thesis, University of Calgary (Alberta).
  • Desmond, A. J. 1984. Archetypes and Ancestors: Palaeontology in Victorian London 1850-1875. The University of Chicago Press, Chicago. 
  • Naish, D. & Martill, D. M. 2001. Saurischian dinosaurs 1: Sauropods. In Martill, D. M. & Naish, D. (eds) Dinosaurs of the Isle of Wight. The Palaeontological Association (London), pp. 185-241.
  • Naish, D. & Martill, D. M. 2007. Dinosaurs of Great Britain and the role of the Geological Society of London in their discovery: basal Dinosauria and Saurischia. Journal of the Geological Society, London 164, 493-510.
  • O’Connor, P. M. 2006. Postcranial pneumaticity: an evaluation of soft-tissue influences on the postcranial skeleton and the reconstruction of pulmonary anatomy in archosaurs. Journal of Morphology 267, 1199-1226.
  • Owen, R. 1856. Monograph on the fossil Reptilia of the Wealden and Purbeck Formations. Part III. Dinosauria (Megalosaurus). Palaeontographical Society Monographs 9, 1-26.
  • Owen, R. 1859. On the vertebral characters of the Order Pterosauria, as exexmplified in the genera Pterodactylus (Cuvier) and Dimorphodon (Owen). Philosophical Transactions of the Royal Society of London 149, 161-169.
  • Owen, R. 1876. Monograph on the fossil Reptilia of the Wealden and Purbeck Formations. Supplement 7. Crocodilia (Poikilopleuron). Dinosauria (Chondrosteosaurus). Palaeontographical Society Monographs 30, 1-7.
  • Seeley, H. G. 1870. On Ornithopsis, a gigantic animal of the pterodactyle kind from the Wealden. Annals and Magazine of Natural History, Series 4, 5 279-283.

Invading the postzyg

March 30, 2008

7306-fossa-medial-side-postzyg-resized.jpg

Again, another exclusive peek at an interesting specimen: the MIWG.7306 vertebra, aka ‘Angloposeidon’ (Naish et al. 2004). Apologies if, by now, you’re bored of my show-casing of this specimen, but – not only is it the only sauropod vertebra of which I personally have multiple unpublished images – it is also a really nice demonstration of the fact that, even in just a single vertebra, there are multiple interesting, bizarre, and sometimes under-studied or even un-studied details.

What we’re looking at here is the medial (‘inside’) surface of the left postzygapophysis, with the centrum down below the bottom of the image, and the cotyle off to the left (the opposite side of what’s shown here). The image below should help with orientation. The focus of interest is the unusual matrix-filled space in the middle of the image: just what is it? Because it has sharp, clean edges, I am pretty convinced that it’s natural, and I assume it’s a pneumatic foramen. Similar structures are present on the medial side of the right postzygapophysis, and are different in position and shape (Naish et al. 2003, p. 790). We think, based on several lines of evidence, that the space between the postzygapophyses (limited anteriorly by the neural spine) was occupied by an air-sac (Schwarz & Fritsch (2006) called this the interspinal diverticulum), so is this evidence that diverticula from the interspinal air-sac invaded the bodies of the postzygapophyses on their medial sides? If so, was this just a one-off in MIWG.7306, or was it widespread in brachiosaurs, in macronarians, in neosauropods, or even in sauropods as a whole? I admit that I haven’t yet taken the time to check properly, but the big problem is that this part of the vertebra – the medial surface of the postzygapophysis – is rarely figured. Based on what has been published, I have yet to see a similar structure, even in Brachiosaurus (which is very well figured, as sauropods go).

miwg7306-prezyg-fossa-composite-resized.jpg

I’m sure that someone is now going to make me look very, very silly. But, whatever. I can’t pretend to know everything. Note that, again, this is a world first. Yes, all of this stuff should be published… and in time in will, in time.

References

  • Naish, D., Martill, D. M., Cooper, D. & Stevens, K. A. 2004. Europe’s largest dinosaur? A giant brachiosaurid cervical vertebra from the Wessex Formation (Early Cretaceous) of southern England. Cretaceous Research 25, 787-795.
  • Schwarz, D. & Fritsch, G. 2006. Pneumatic structures in the cervical vertebrae of the Late Jurassic Tendaguru sauropods Brachiosaurus brancai and Dicraeosaurus. Eclogae geol. Helv. 99, 65-78.
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