October 20, 2015
I just gave an answer to this question on Quora, and it occurred to me that I ought to also give it a permanent home here. So here it is.
This is a great example of a question that you’d think would have a simple, clear answer, but doesn’t. In fact, as a palaeontologist specialising in dinosaur gigantism, I have an abiding fear of being asked this question in a pub quiz, and not being able to produce the name that’s written on the quizmaster’s answer sheet.
First, what do we mean by “biggest”? Diplodocus was longer than Apatosaurus, but Apatosaurus was heavier. Giraffatitan was taller than either. Let’s simplify and assume we want to know the heaviest dinosaur.
Second, estimating the masses of extinct animals is incredibly hard even when we have a pretty complete skeleton. For example, the gigantic mounted brachiosaur skeleton in Berlin (which used to be called “Brachiosaurus” brancai but is now recognised as the separate genus Giraffatitan) has been subject to at least 14 estimates in the published scientific literature,. They vary from 13,618 kg to 78,258 kg — a factor of 5.75 for the same individual. That’s like looking at a human skeleton and not knowing whether its from Kate Moss or Arnold Schwazenegger. (There are reasons for this and I urge you to read the linked article.)
Third, the big dinosaurs tend to be very poorly represented. Giraffatitan is probably the heaviest dinosaur known from a more or less complete skeleton (though even that is put together from several different individuals) so I could give that as the answer to the hypothetical pub-quiz — though the answer sheet would probably be out of date and call it Brachiosaurus.
Fourth, which individual of a given species do we mean? I said Giraffatitan is known from a more or less complete skeleton. And my best guess is that that individual massed, say, 30,000 kg. But an isolated fibula of the same species is known that’s 12.6% longer than the one in the skeletal mount. That suggest an animal that masses 1.126^3 = 1.43 times as massive as the mounted skeleton — say 43,000 kg. There might be yet bigger Giraffatitan individuals. On the other hand, there isthat Apatosaurus, which is usually thought of as not being so big, might have got even bigger.
Fifth, the very biggest specimens tend to be known from only a handful of bones. A good example here is the titanosaur Argentinosaurus, which is known from several vertebrae and a few limb bones, but not all from the same individual. It’s a good bet that it massed 60-70 tonnes — so maybe about twice as much as Giraffatitan, but much less than the often-cited 100 tonnes. Other, more recently discovered, titanosaurs seem to be in the same size class: Puertasaurus, Futalognkosaurus, Dreadnoughtus and more. They they are hard to compare directly due to the paucity of overlapping material, or at least . (Scientists are working on getting more of this stuff properly described in the literature, which will help.)
But, sixth, the very biggest dinosaurs tend to be apocryphal. There’s Amphicoelias fragillimus,. This may have been 50 m long and massed 80 tonnes; but other published estimates say 58 m and 122 tonnes. We really can’t say from the very poor remains.
So if you get asked this question in a pub quiz, your best bet is to roll a dice, pick an answer, close your eyes and hope. Roll 1 for Giraffatitan, 2 for Brachiosaurus, 3 for Apatosaurus, 4 for Argentinosaurus, 5 for Dreadnoughtus and 6 for Amphicoelias fragillimus. Good luck!
October 19, 2015
I imagine that by now, everyone who reads this blog is familiar with Mark Witton’s painting of a giant azhdarchid pterosaur alongside a big giraffe. Here it is, for those who haven’t seen it:
(This is the fifth and most recent version that Mark has created, taken from 9 things you may not know about giant azhdarchid pterosaurs.)
It’s one of those images that really kicks you in the brain the first time you see it. The idea that an animal the size of a giraffe could fly under its own power seems ludicrous — yet that’s what the evidence tells us.
But wait — what do we mean by “an animal the size of a giraffe”? Yes, the pterosaur in this image is the same height as the giraffe, but how does its weight compare?
Mark says “The giraffe is a big bull Masai individual, standing a healthy 5.6 m tall, close to the maximum known Masai giraffe height.” He doesn’t give a mass, but Wikipedia, citing Owen-Smith (1988), says “Fully grown giraffes stand 5–6 m (16–20 ft) tall, with males taller than females. The average weight is 1,192 kg (2,628 lb) for an adult male and 828 kg (1,825 lb) for an adult female with maximum weights of 1,930 kg (4,250 lb) and 1,180 kg (2,600 lb) having been recorded for males and females, respectively.” So it seems reasonable to use a mass intermediate between those of an average and maximum-sized male, (1192+1930)/2 = 1561 kg.
So much for the giraffe. What does the azhdarchid weigh? The literature is studded with figures that vary wildly, from the 544 kg that Henderson (2010) found for Quetzalcoatlus, right down to the widely cited 70 kg that Chatterjee and Templin (2004) found for the same individual — and even the astonishing 50 kg that seems to be favoured by Unwin (2005:192). In the middle is the 259 kg of Witton (2008).
It occurred to me that I could visualise these mass estimates by shrinking the giraffe in Mark’s image down to the various proposed masses, and seeing how credible it looks to imagine these reduced-sized giraffes weighting the same as the azhdarchid. The maths is simple. For each proposed azhdarchid mass, we figure out what it is as a proportion of the giraffe’s 1561 kg; then the cube root of that mass proportion gives us the linear proportion.
- 544 kg = 0.389 giraffe masses = 0.704 giraffe lengths
- 259 kg = 0.166 giraffe masses = 0.549 giraffe lengths
- 70 kg =0.0448 giraffe masses = 0.355 giraffe lengths
Let’s see how that looks.
On the left, we have Mark’s artwork, with the giraffe massing 1561 kg. On the right, we have three smaller (isometrically scaled) giraffes of masses corresponding to giant azhdarchid mass estimates in the literature. If Don Henderson (2010) is right, then the pterosaur weighs the same as the 544 kg giraffe, which to me looks pretty feasible if it’s very pneumatic. If Witton (2008) is right, then it weighs the same as the 259 kg giraffe, which I find hard to swallow. And if Chatterjee and Templin (2004) are right, then the giant pterosaur weighs the same as the teeny tiny 70 kg giraffe, which I find frankly ludicrous. (For that matter, 70 kg is in the same size-class as Georgia, the human scale-bar: the idea that she and the pterosaur weigh the same is just silly.)
What is the value of such eyeball comparisons? I’m not sure, beyond a basic reality check. Running this exercise has certainly made me sceptical about even the 250 kg mass range which now seems to be fairly widely accepted among pterosaur workers. Remember, if that mass is correct then the pterosaur and the 259 kg giraffe in the picture above weight the same. Can you buy that?
Or can we find extant analogues? Are there birds and mammals with the same mass that are in the same size relation as these images show?
- Chatterjee, Sankar, and R. J. Templin. 2004. Posture, locomotion, and paleoecology of pterosaurs. Geological Society of America, Special Paper 376. 68 pages.
- Henderson, Donald M. 2010. Pterosaur body mass estimates from three-dimensional mathematical slicing. Journal of Vertebrate Paleontology 30(3):768-785.
- Witton, Mark P. 2008. A new approach to determining pterosaur body mass and its implications for pterosaur flight. Zitteliana 28:143-159.
October 15, 2015
Following on from his recent, and extensively discussed, offer to host SVPCA 2017, and a plan for the future, Richard Butler is now circulating his update, soliciting volunteers for the committee that virtually everyone agreed was a good idea.
Dear SVPCA/SPPC friends and colleagues,
We have identified you as a member of the SVPCA/SPPC community through having attended the meeting within the last five years. Many of you will doubtless be aware of the vibrant, lengthy, and occasionally fiery debate currently taking place about the future of the meeting on Mike Taylor and Matt Wedel’s SV-POW blog.
The least controversial proposal for change in the meeting has been my suggestion that we establish a ‘steering group’ to (i) try and solve some of the short-term and long-term logistical challenges (bank accounts, abstract submission and online registration issues etc.); (ii) provide support to meeting organisers and develop a comprehensive set of useful information; (iii) help identify and encourage future hosts to come forward; (iv) think about and discuss the future of the meeting, including discussing how best to make sure the meeting appeals to the entire community, from students to amateurs and from professors to preparators.
As there has been no opposition and plenty of support for a steering group, I propose we move forward with establishing this. No alternative to my proposal for the composition of this group has been put forward: I proposed a group of seven including past, current and future organisers (Gareth Dyke, Peter Falkingham, me as proposed host for a 2017 Birmingham meeting), and four elected members representing the student, early career academic (up to 10 years post-PhD), senior academic, and non-professional communities. This would not draw any museum/university distinction when it comes to students and academics. Although I have not heard formally from the GCG, I understand there is interest in one of their members being co-opted onto the steering group to represent SPPC. Elected steering group members could serve three-year terms, to match the terms served by meeting hosts.
At this stage we need volunteers: people willing to stand for election to this group and help secure and shape the future of the meeting! If you are a member of the SVPCA/SPPC community and are interested in serving, then please email me and Richard Forrest and let us know which of the four elected positions you wish to stand for. We would like volunteers by October 23rd. After this date, we will set-up online elections to allow the SVPCA/SPPC community to vote for each of the positions.
I look forward to hearing from you and to working together to shape the future of the meeting.
Copyright: promoting the Progress of Science and useful Arts by preventing access to 105-year-old quarry maps
October 11, 2015
In my recent preprint on the incompleteness and distortion of sauropod neck specimens, I discuss three well-known sauropod specimens in detail, and show that they are not as well known as we think they are. One of them is the Giraffatitan brancai lectotype MB.R.2181 (more widely known by its older designation HMN SII), the specimen that provides the bulk of the mighty mounted skeleton in Berlin.
That photo is from this post, which is why it’s disfigured by red arrows pointing at its epipophyses. But the vertebra in question — the eighth cervical of MB.R.2181 — is a very old friend: in fact, it was the subject of the first ever SV-POW! post, back in 2007.
In the reprint, to help make the point that this specimen was found extremely disarticulated, I reproduce Heinrich (1999:figure 16), which is Wolf-Dieter Heinrich’s redrawing of Janensch’s original sketch map of Quarry S, made in 1909 or 1910. Here it is again:
For the preprint, as for this blog-post (and indeed the previous one), I just went right ahead and included it. But the formal version of the paper (assuming it passes peer-review) will by very explicitly under a CC By licence, so the right thing to do is get formal permission to include it under those terms. So I’ve been trying to get that permission.
What a stupid, stupid waste of time.
Heinrich’s paper appeared in the somewhat cumbersomely titled Mitteilungen aus dem Museum fur Naturkunde in Berlin, Geowissenschaftliche Reihe, published as a subscription journal by Wiley. Happily, that journal is now open access, published by Pensoft as The Fossil Record. So I wrote to the Fossil Record editors to request permission. They wrote back, saying:
We are not the right persons for your question. The Wiley Company holds the copyright and should therefore be asked. Unfortunately, I do not know who is the correct person.
Thank you for your enquiry.
We are currently experiencing a large volume of email traffic and will deal with your request within the next 15 working days.
We are pleased to advise that permission for the majority of our journal content, and for an increasing number of book publications, may be cleared more quickly by using the RightsLink service via Wiley’s websites http://onlinelibrary.wiley.com and www.wiley.com.
Within the next fifteen working days? That is, in the next three weeks? How can it possibly take that long? Are they engraving their response on a corundum block?
So, OK, let’s follow the automated suggestion and try RightsLink. I went to the Wiley Online Library, and searched for journals whose names contain “naturkunde”. Only one comes up, and it’s not the right one. So Wiley doesn’t admit the existence of the journal.
Well, there’s lots to enjoy here, isn’t there? First, and most important, it doesn’t actually work: “Permission to reproduce this content cannot be granted via the RightsLink service.” Then there’s that cute little registered-trademark symbol “®” on the name RightsLink, because it’s important to remind me not to accidentally set up my own rights-management service with the same name. In the same vein, there’s the “Copyright © 2015 Copyright Clearance Center, Inc. All Rights Reserved” notice at the bottom — copyright not on the content that I want to reuse, but on the RightsLink popup itself. (Which I guess means I am in violation for including the screenshot above.) Oh, and there’s the misrendering of “Museum für Naturkunde” as “Museum fÃ¼r Naturkunde”.
All of this gets me precisely nowhere. As far as I can tell, my only recourse now is to wait three weeks for Wiley to get in touch with me, and hope that they turn out to be in favour of science.
It’s Sunday afternoon. I could be watching Ireland play France in the Rugby World Cup. I could be out at Staverton, seeing (and hearing) the world’s last flying Avro Vulcan overfly Gloucester Airport for the last time. I could be watching Return of the Jedi with the boys, in preparation for the forthcoming Episode VII. Instead, here I am, wrestling with copyright.
How absolutely pointless. What a terrible waste of my life.
Is this what we want researchers to be spending their time on?
Update (13 October 2015): a happy outcome (this time)
I was delighted, on logging in this morning, to find I had email from RIGHTS-and-LICENCES@wiley-vch.de with the subject “Permission to reproduce Heinrich (1999:fig. 16) under CC By licence” — a full thirteen working days earlier than expected. They were apologetic and helpful. Here is key part of what they said:
We are of course happy to handle your request directly from our office – please find the requested permission here:We hereby grant permission for the requested use expected that due credit is given to the original source.If material appears within our work with credit to another source, authorisation from that source must be obtained.Credit must include the following components:– Journals: Author(s) Name(s): Title of the Article. Name of the Journal. Publication year. Volume. Page(s). Copyright Wiley-VCH Verlag GmbH & Co. KGaA. Reproduced with permission.
So this is excellent. I would of course have included all those elements in the attribution anyway, with the exception that it might not have occurred to me to state who the copyright holder is. But there is no reason to object to that.
So, two cheers for Wiley on this occasion. I had to waste some time, but at least none of it was due to deliberate obstructiveness, and most importantly they are happy for their figure to be reproduced under CC By.
- Heinrich, Wolf-Dieter. 1999. The taphonomy of dinosaurs from the Upper Jurassic of Tendaguru, Tanzania (East Africa), based on field sketches of the German Tendaguru expedition (1909-1913). Mitteilungen aus dem Museum fur Naturkunde in Berlin, Geowissenschaftliche Reihe 2:25-61.
October 9, 2015
Since I posted my preprint “Almost all known sauropod necks are incomplete and distorted” and asked in the comments for people to let me know if I missed any good necks, the candidates have been absolutely rolling in:
- The Kaatedocus siberi holotype SMA 0004 (thanks to Oliver Demuth for pointing this out)
- The Futalognkosaurus dukei holotype MUCPv-323 (thanks to Matt Lamanna)
- The referred Rapetosaurus specimen FMNH PR 2209 (Matt Lamanna again)
- The as-yet unnnamed DGM ‘Series A’ titanosaur from Peirópolis (Matt Lamanna again)
- The Camarasaurus lewisi holotype BYU 9047 (thanks to John D’Angelo)
- Dicraeosaurus hansemanni — the mounted “m” specimen, probably MB.R.4886 (thanks to Emanuel Tschopp)
- Maybe some Omeisaurus specimens (Emanuel Tschopp again)
I will be investigating the completeness of all of these and mentioning them as appropriate when I submit the revision of this paper. (In retrospect, I should have waited a week after posting the preprint before submitting for formal review; but I was so scared of letting it brew for years, as we’re still doing with the Barosaurus preprint to our shame, that I submitted it immediately.)
So we probably have a larger number of complete or near-complete sauropod necks than the current draft of this paper suggests. But still very few in the scheme of things, and essentially none that aren’t distorted.
So I want to consider why we have such a poor fossil record of sauropod necks. All of the problems with sauropod neck preservation arise from the nature of the animals.
First, sauropods are big. This is a recipe for incompleteness of preservation. (It’s no accident that the most completely preserved specimens are of small individuals such as CM 11338, the cow-sized juvenile Camarasaurus lentus described by Gilmore, 1925). For an organism to be fossilised, the carcass has to be swiftly buried in mud, ash or some other substrate. This can happen relatively easily to small animals, such as the many finely preserved stinkin’ theropods from the Yixian Formation in China, but it’s virtually impossible with a large animal. Except in truly exceptional circumstances, sediments simply don’t get deposited quickly enough to cover a 25 meter, 20 tonne animal before it is broken apart by scavenging, decay and water transport.
Secondly, even when complete sauropods are preserved, or at least complete necks, distortion of the preserved cervical vertebrae is almost inevitable because of their uniquely fragile construction. As in modern birds, the cervical vertebrae were lightened by extensive pneumatisation, so that they were more air than bone, with the air-space proportion typically in the region of 60–70% and sometimes reaching as high as 89%. While this construction enabled the vertebrae to withstand great stresses for a given mass of bone, it nevertheless left them prone to crushing, shearing and torsion when removed from their protective layer of soft tissue. For large cervicals in particular, the chance of the shape surviving through taphonomy, fossilisation and subsequent deformation would be tiny.
So I think we’re basically doomed never to have a really good sauropod neck skeleton.
October 7, 2015
Well, I’m a moron again. In the new preprint that I just published, I briefly discussed the six species of sauropod for which complete necks are known — Camarasaurus lentus (but it’s a juvenile), Apatosaurus louisae (but the last three and maybe C5 are badly damaged), Mamenchisaurus hochuanensis (but all the vertebrae are broken and distorted), Shunosaurus lii, Mamenchisaurus youngi and Spinophorosaurus nigerensis.
I did have the wit to say, in the Author Comment:
Although I am submitting this article for formal peer-review at the same time as publishing it as a preprint, I also solicit comments from readers. In particular I am very keen to know if I have missed any complete sauropod necks that have been described in the literature. In the final version of the manuscript, I will acknowledge those who have offered helpful comments.
Happily, several people have taken me up on this (see the comments on the preprint), but one suggestion in particular was a real D’oh! moment for me. Oliver Demuth reminded me about Kaatedocus — a sauropod that we SV-POW!sketeers love so much that it has its own category on our site and we’ve held it up as an example of how to illustrate a sauropod specimen. More than that: we have included several illustrations of its vertebrae in one of our own papers.
Aaanyway … the purpose of this post is just to get all the beautiful Kaatedocus multiview images up in one convenient place. They were freely available as supplementary information to the paper, but now seem to have vanished from the publisher’s web-site. I kept copies, and now present them in the conveniently viewable JPEG format (rather the download-only TIFF format of the originals) and with each image labelled with its position in the column.
Please note, these images are the work of Tschopp and Mateus (2012) — they’re not mine!
C15 (and the rest of the skeleton) is missing, which makes this a very nearly, but not quite, complete sauropod neck.
- Tschopp, Emanuel, and Octávio Mateus. 2012. The skull and neck of a new flagellicaudatan sauropod from the Morrison Formation and its implication for the evolution and ontogeny of diplodocid dinosaurs. Journal of Systematic Palaeontology 11(7):853-888. doi:10.1080/14772019.2012.746589
October 6, 2015
I have a new preprint up at PeerJ (Taylor 2015), and have also submitted it simultaneously for peer review. In a sense, it’s not a paper I am happy about, as its title explains: “Almost all known sauropod necks are incomplete and distorted“.
This paper has been a while coming, and much of the content will be familiar to long-time readers, as quite a bit of it is derived from three SV-POW! posts: How long was the neck of Diplodocus? (2011), Measuring the elongation of vertebrae (2013) and The Field Museum’s photo-archives tumblr, featuring: airbrushing dorsals (2014). It also uses the first half of my 2011 SVPCA talk, Sauropod necks: how much do we really know? (and the second half became the seed that grew into our 2013 neck-cartilage paper.)
So in one sense, publishing this is a bit of a mopping up exercise. But it’s also more than that, because I think it’s important to get all these observations (and the relevant literature review) down all in one place, to help us recognise just how serious the problem is. There are, to a first approximation, no complete sauropod necks in the published literature. And the vertebrae of the necks we do have are crushed to the point where trying to articulate them is close to meaningless.
I’m not happy about this. But I think it’s important to face the reality and be honest with ourselves about how much we can really know about sauropod necks. There’s a lot we can do in a qualitative way, but most quantitative results are going to be swamped in supposition and error.