Juvenile Tomistoma schlegelii, LACM Herpetology 166483, with me for scale. It wasn’t until I picked up the skull that I realized it was the same specimen I had looked at back when. I was looking at its neck in 2011, and its tail today, for reasons that will be revealed at the dramatically appropriate moment. I was only playing with the skull because it’s cute, an intricate little marvel of natural selection. Photos by Vanessa Graff (2011) and Jessie Atterholt (2018). Many thanks to collections manager Neftali Camacho for his hospitality and assistance both times!


John Yasmer, DO (right) and me getting ready to scan MWC 8239, a caudal vertebra of Diplodocus on loan from Dinosaur Journey, at Hemet Valley Imaging yesterday.

Alignment lasers – it’s always fun watching them flow over the bone as a specimen slides through the tube (for alignment purposes, obviously, not scanning – nobody’s in the room for that).

Lateral scout. I wonder, who will be the first to correctly identify the genus and species of the two stinkin’ mammals trailing the Diplo caudal?

A model we generated at the imaging center. This is just a cell phone photo of a single window on a big monitor. The actual model is much better, but I am in a brief temporal lacuna where I can’t screenshot it.

What am I doing with this thing? All will be revealed soon.

Re-reading an email that Matt sent me back in January, I see this:

One quick point about [an interesting sauropod specimen]. I can envision writing that up as a short descriptive paper, basically to say, “Hey, look at this weird thing we found! Morrison sauropod diversity is still underestimated!” But I honestly doubt that we’ll ever get to it — we have literally years of other, more pressing work in front of us. So maybe we should just do an SV-POW! post about the weirdness of [that specimen], so that the World Will Know.

Although as soon as I write that, I think, “Screw that, I’m going to wait until I’m not busy* and then just take a single week* and rock out a wiper* on it.”

I realize that this way of thinking represents a profound and possibly psychotic break with reality. *Thrice! But it still creeps up on me.

(For anyone not familiar with the the “wiper”, it refers to a short paper of only one or two pages. The etymology is left as an exercise to the reader.)

It’s just amazing how we keep on and on falling for this delusion that we can get a paper out quickly, even when we know perfectly well, going into the project, that it’s not going to work out that way. To pick a recent example, my paper on quantifying the effect of intervertebral cartilage on neutral posture was intended to be literally one page, an addendum to the earlier paper on cartilage: title, one paragraph of intro, diagram, equation, single reference, DONE! Instead, it landed up being 11 pages long with five illustrations and two tables.

I think it’s a reasonable approximation to say that any given project will require about an order of magnitude more work than we expect at the outset.

Even as I write this, the top of my palaeo-work priority list is a paper that I’m working on with Matt and two other colleagues, which he kicked off on 6 May, writing:

I really, really want to kill this off absolutely ASAP. Like, seriously, within a week or two. Is that cool? Is that doable?

To which I idiotically replied:


A month and a bit later, the answers to Matt’s questions are clear. Yes, it’s cool; and no, it’s not doable.

The thing is, I think that’s … kind of OK. The upshot is that we end up writing reasonably substantial papers, which is after all what we’re meant to be trying to do. If the reasonably substantial papers that end up getting written aren’t necessarily the ones we thought they were going to be, well, that’s not a problem. After all, as I’ve noted before, my entire Ph.D dissertation was composed of side-projects, and I never got around to doing the main project. That’s fine.

In 2011, Matt’s tutorial on how to find problems to work on discussed in detail how projects grow and mutate and anastamose. I’m giving up on thinking that this is a bad thing, abandoning the idea that I ought to be in control of my own research program. I’m just going to keep chasing whatever rabbits look good to me at the time, and see what happens.


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.


  • 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.


  • 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).

I recently reread Dubach (1981), “Quantitative analysis of the respiratory system of the house sparrow, budgerigar and violet-eared hummingbird”, and realized that she reported both body masses and volumes in her Table 1. For each of the three species, here are the sample sizes, mean total body masses, and mean total body volumes, along with mean densities I calculated from those values.

  • House sparrow, Passer domesticus, n = 16, mass = 23.56 g, volume = 34.05 mL, density = 0.692 g/mL
  • Budgerigar, Melopsittacus undulatus, n = 19, mass = 38.16 g, volume = 46.08 mL, density = 0.828 g/mL
  • Sparkling violetear,* Colibri coruscans, n = 12, mass = 7.28 g, volume = 9.29 mL, density = 0.784 g/mL

* This is the species examined by Dubach (1981), although not specified in her title; there are four currently-recognized species of violetears. And apparently ‘violetear’ has overtaken ‘violet-eared hummingbird’ as the preferred common name. And as long as we’re technically on a digression,  I’m almost certain those volumes do not include feathers. Every volumetric thing I’ve seen on bird masses assumes plucked birds (read on).

This is pretty darned interesting to me, partly because I’m always interested in how dense animals are, and partly because of how the results compare to other published data on whole-body densities for birds. The other results I am most familiar with are those of Hazlehurst and Rayner (1992) who had this to say:

There are relatively few values for bird density. Welty (1962) cited 0.9 g/mL for a duck, and Alexander (1983) 0.937 g/mL for a domestic goose, but those values may not take account of the air sacs. Paul (1988) noted 0.8 g/mL for unspecified bird(s). To provide more reliable estimates, the density of 25 birds of 12 species was measured by using the volume displacement method. In a dead, plucked bird the air-sac system was reinflated (Saunder and Manton 1979). The average density was 0.73 g/mL, suggesting that the lungs and air sacs occupy some quarter of the body.

That result has cast a long shadow over discussions of sauropod masses, as in this paper and these posts, so it’s nice to see similar results from an independent analysis.  If you’re curious, the weighted mean of the densities calculated from Duchard’s Dubach’s (1981) data is 0.77. I’d love to see the raw data from Hazlehurst and Rayner (1992) to see how much spread they got in their density measurements.  Unfortunately, they did not say which birds they used or give the raw data in the paper (MYDD!), and I have not asked them for it because doing so only just occurred to me as I was writing this post.

There will be more news about hummingbirds here in the hopefully not-too-distant future. Here’s a teaser:


Yes, those are its hyoids wrapped around the back of its head–they go all the way around to just in front of the eyes, as in woodpeckers and other birds that need hyper-long tongue muscles. There are LOADS of other interesting things to talk about here, but it will be faster and more productive if I just go write the paper like I’m supposed to be doing.

Oh, all right, I’ll say a little more. This is a  young adult female Anna’s hummingbird, Calypte anna, who was found by then-fellow-grad-student Chris Clark at a residential address in Berkeley in 2005. She was unable to fly and died of unknown causes just a few minutes after being found. She is now specimen 182041 in the ornithology collection at the Museum of Vertebrate Zoology at Berkeley. Chris Clark and I had her microCTed back in 2005, and that data will finally see the light of day thanks to my current grad student, Chris Michaels, who generated the above model.

This bird’s skull is a hair over an inch long, and she had a body mass of 3.85 grams at the time of her death. For comparison, those little ketchup packets you get at fast-food burger joints each contain 8-9 grams of ketchup, more than twice the mass of this entire bird when it was alive!


  • Dubach, M. 1981. Quantitative analysis of the respiratory system of the house sparrow, budgerigar and violet-eared hummingbird. Respiration Physiology 46(1): 43-60.
  • Hazlehurst, G.A., and Rayner, J.M. 1992. Flight characteristics of Triassic and Jurassic Pterosauria: an appraisal based on wing shape. Paleobiology 18(4): 447-463.