Some further thoughts on Patagotitan
August 10, 2017
A bunch of stuff, loosely organized by theme.
Media
First up, I need to thank Brian Switek, who invited me to comment on Patagotitan for his piece at Smithsonian. I think he did a great job on that, arguably the best of any of the first-day major media outlet pieces. And it didn’t go unnoticed – his article was referenced at both the Washington Post and NPR (and possibly other outlets, those are the two I know of right now). I don’t think my quotes got around because they’re particularly eloquent, BTW, but rather because reporters tend to like point-counterpoint, and I was apparently the most visible counterpoint. They probably would have done the same if I’d been talking complete nonsense (which, to be fair, some people may think I was).
Paleobiology vs Records
The most commonly reproduced quote of mine is this one, originally from Brian’s piece:
I think it would be more accurate to say that Argentinosaurus, Puertasaurus and Patagotitan are so similar in size that it is impossible for now to say which one was the largest.
That may seem at odds with the, “Well, actually…[pushes glasses up nose]…Argentinosaurus was still biggest” tack I’ve taken both in my post yesterday and on Facebook. So let me elaborate a little.
There is a minor, boring point, which is that when I gave Brian that quote, I’d seen the Patagotitan paper, but not the Electronic Supplementary Materials (ESM), so I knew that Patagotitan was about the same size as the other two (and had known for a while), but I hadn’t had a chance to actually run the numbers.
The much more interesting point is that the size differences between Argentinosaurus, Puertasaurus, and Patagotitan are astonishingly small. The difference between a 2.5m femur and a 2.4m one is negligible, ditto for vertebrae with centra 59cm and 60cm in diameter. OMNH 1331, the biggest centrum bit from the giant Oklahoma apatosaur, had an intact max diameter of 49cm, making it 26% larger in linear terms than the next-largest apatosaur. The centra of these giant South American titanosaurs are more than 20% bigger yet than OMNH 1331, just in linear terms. That’s crazy.
It’s also crazy that these three in particular – Argentinosaurus, Puertasaurus, and Patagotitan – are so similar in size. Dinosaur developmental programs were ‘messy’ compared to those of mammals, both in having weird timings for things like onset of reproduction, and in varying a lot among closely related taxa. Furthermore, sauropod population dynamics should have been highly skewed toward juveniles and subadults. So is the near-equality in size among Argentinosaurus, Puertasaurus, and Patagotitan just a coincidence, or does it mean that something weird was going on? There’s really no third option. I mean, even if some kind of internal (biomechanical or physiological) or external (ecological, food or predation) constraint forced those three to the same adult body size, it’s weird then that we’re finding only or at least mostly near-max-size adults. (If the available specimens of these three aren’t near-max-size, then any hypothesis that they’re forced to the same size by constraints is out the window, and we’re back to coincidence.)
BUT
With all that said, the title of “world’s largest dinosaur” is not handed out for effort expended, number of specimens collected, skeletal completeness, ontogenetic speculation, or anything other than “the dinosaur with the largest measured elements”. And that is currently Argentinosaurus. So although for any kind of paleobiological consideration we can currently consider Argentinosaurus, Puertasaurus, and Patagotitan to all be about the same size – and Alamosaurus, Paralititan, Notocolossus, and probably others I’ve forgotten should be in this conversation – anyone wanting to dethrone Argentinosaurus needs to actually show up with bigger elements.
So, if you’re interested in paleobiology, it’s fascinating and frankly kind of unnerving that so many of these giant titanosaurs were within a hand-span of each other in terms of size. Patagotitan is one more on the pile – and, as I said yesterday, exciting because it’s so complete.
But if you want to know who holds the crown, it’s still Argentinosaurus.
Humeri
In a comment on the last post, Andrea Cau made an excellent point that I am just going to copy here entire:
Even Paralititan stromeri humerus is apparently larger than Patagotitan humerus (169 cm vs 167.5 cm). I know humerus length alone is bad proxy of body size, but at least this shows that even in that bone Patagotitan is just another big titanosaur among a well known gang of titans, not a supersized one.
That made me want to start a list of the longest sauropod humeri. Here goes – if I missed anyone or put down a figure incorrectly, I’m sure you’ll let me know in the comments.
- Giraffatitan: 213cm
- Brachiosaurus: 203cm
- Ruyangosaurus: 190cm (estimated from 135cm partial)
- Turiasaurus: 179cm
- Notocolossus: 176cm
- Paralititan: 169cm
- Patagotitan: 167.5cm
- Dreadnoughtus: 160cm
- Futlognkosaurus: 156cm
Admittedly the Patagotitan humerus is from a paratype and not from the largest individual, but that is true for some others on the list, including Giraffatitan. And we have no humeri from Argentinosaurus, Puertasaurus, and some other giants.
Dorsal Vertebrae
A couple of further thoughts on how the dorsal vertebrae of Patagotitan compare to those of Argentinosaurus. First, now that I’ve had some time to think about it, I have a hard time seeing how the dorsal polygon method used by Carballido et al. in the Patagotitan paper has any biological meaning. In their example figure, the polygon around the Puertasaurus vertebra is mostly full of bone, and the one around Patagotitan has a lot of empty space. It’s easy to imagine an alternative metric, like “area of the minimum polygon actually filled by bone”, that would lead to a different ‘winner’. But that wouldn’t mean much, either.
Something that probably does have a real and important biomechanical meaning is the surface area of the articular face of the centrum, because that’s the area of bone that has to bear the compressive load, which is directly related to the animal’s mass. The biggest Patagotitan centrum is that of MPEF-PV 3400/5, which is at least a local maximum since has smaller centra both ahead and behind. The posterior face measures 59cm wide by 42.5cm tall. Abstracted as an ellipse, which may not be perfectly accurate, those measurements give a surface area of (pi)(29.5)(21.25)=1970 cm^2. For Argentinosaurus, the largest complete centrum has a posterior face measuring 60cm wide by 47cm tall (Bonaparte and Coria 1993: p. 5), giving an elliptical surface area of (pi)(30)(23.5)=2210 cm^2. (I’d use hi-res images of the centra to measure the actual surface areas if I could, but AFAIK those images either don’t exist or at least have not yet been made public, for either taxon.) So although the Argentinosaurus dorsal seems like it is only a bit bigger in linear terms, it’s 12% larger in surface area, and that might actually be a meaningful difference.
Cervical Vertebrae
One thing I haven’t commented on yet – Patagotitan is the newest member of the “world’s longest vertebrae” club. The longest Patagotitan cervical, MPEF-PV 3400/3, is listed in the ESM as having a centrum length of 120cm, but it’s also listed as incomplete. In the skeletal recon in the paper, the centrum is colored in as present, but the neural spine is missing. So is the centrum complete in terms of length? I don’t think it’s clear right now.
Anyway, here’s the current rundown of the longest cervical centra of sauropods (and therefore, the longest vertebrae among animals):
- BYU 9024, possibly referable to Supersaurus or Barosaurus: 137cm
- Price River 2 titanosauriform: 129cm
- OMNH 53062, Sauroposeidon holotype: 125cm
- KLR1508-77-2, Ruyangosaurus giganteus referred specimen: 124cm
- MPEF-PV 3400/3, Patagotitan holotype: 120cm (+?)
- MPM 10002, Puertasaurus holotype: 118cm
You may be surprised to see the Price River 2 cervical in there. It was reported in an SVP abstract a few years ago (I’ll dig up that ref and update this post), and Mike and I saw it last year on the Sauropocalypse. We measured the centrum at 129cm, making it just a bit longer than the longest centrum of Sauroposeidon, and therefore the second-longest vertebra of anything ever.
Aside – I’m probably getting a reputation as a big ole meanie when it comes to debunking “world’s largest dinosaur” claims. If I’m willing to take the lead in kicking my own dinosaur down the ladder, don’t expect me to be kind to yours. I follow where the numbers lead.
Now, here’s an interesting thing – now that Sauroposeidon is coming out as a basal titanosaur, rather than a brachiosaur, it might not have been a skinny freak. The 120cm cervical of Patagotitan makes the 125cm cervical of Sauroposeidon and the 129cm cervical from Price River 2 look even more tantalizing. Maybe it’s super-giant sauropods all the way down.
Sauropod Vertebra Picture of the Week 
August 11, 2017 at 3:31 am
@ Dr. Wedel:
What could have been the primary driver among the factors you have mentioned that led to the rise of those megasauropods? And given that Patagotitan and Argentinosaurus are sister taxa (at least in the article of Carballido et al., 2017), doesn’t that possibly indicate that Patagotitan evolved into Argentinosaurus (given that the former’s remains are found in slightly older rock horizons than the latter).
August 12, 2017 at 3:20 am
Interesting. I’ve never heard of the Prince River 2 Titanosauriform. Do we know where in the cervical series its placed in (ex. Is it a C7 or C9 or C5, etc.)?
August 12, 2017 at 9:21 am
We don’t know, I’m afraid. The PR2 cervical is poorly preserved and in a dreadful state.
August 13, 2017 at 8:25 pm
I thought Sauroposeidon was sexier as a brachiosaur, but I suppose we must not let awesomebro-ism blind us to the science… That’s where these silly “mine is bigger than yours!” arguments come from, anyway.
August 15, 2017 at 6:16 pm
Longest cervicals reffered to Ruyangosaurus are more than 120 cm long, longest – 124 cm.
Osteology of the giant sauropod dinosaur Ruyangosaurus giganteus Lü et al., 2014
August 15, 2017 at 7:05 pm
I can confirm that Shahen is correct, I have a copy of the osteology paper, shared with me by a friend who received it from Phil Mannion.
Ruyangosaurus would be an absolute giant as reconstructed by Lü et al 2014, but they gave it an extremely high cervical count (19 or 20 if my memory serves). Issues have been brought up in my circles that the referred material doesn’t match the holotypes much (femoral and dorsal overlap) but this is as yet speculation, and doesn’t mean that the material wasn’t from a huge giant.
August 15, 2017 at 7:14 pm
In paper is information about some cervicals 120+ cm, sacrum 147 cm, humerus 190 cm!, Largest dorsal with centrum 61 cm, sacral with centrum 68 cm, etc. Mike and Matt please check email
Best Regards
August 15, 2017 at 8:23 pm
Hikaru – good questions. All of the factors that make sauropods ‘work’, at least in the Sander et al. (2010) model, work progressively better at larger size. To me, that’s enough to explain why super-giant sauropods evolved. What I’m more curious about is what factors kept all the other sauropods from evolving to super-giant sizes. Or maybe most sauropods did, and we just haven’t found the giant, fully mature individuals yet.
Vishal – as far as I know, the 129-cm centrum is the only element from a super-giant titanosauriform from that quarry. I’ve seen the rest of the material, and none of it struck me as being in a size class larger than, say, the B. altithorax holotype. Mike, do you remember anything differently?
Shahen and ijreid – thank you for the very good catch on Ruyangosaurus, I will update the post accordingly!
August 16, 2017 at 7:44 pm
There is my small analyze about biggest Dinosaurs. Also i belive that Puertasaurus was bigger than Argentinosaurus. They centra are the same wide, but Puertasaurus diapophyses are huge. Some people said that Argentinosaurus had much longer torso, but in Lognkosauria second dorsal is smaller than rest. For example first dorsal Futalognkosaurus is 43 cm and second only 2/3 that length, in Patagotitan the same. I belive that Puertasaurus 1st dorsal was around 44 cm.
I send also file
https://www.sendspace.com/file/1in7kr
What do you think about this?
August 16, 2017 at 9:18 pm
The position of the Puertasaurus dorsal is a guess. It might be a good guess, it might not be. Given how variable vertebral form is even among titanosaurs, I would not hang a lot of importance on it definitely being D2, rather than D1 or D3. I am also not overly moved by the wide transverse processes. Maaaaybe that means that the torso was wider – certainly some people have been happy to assume that – but maybe it just means that the ribs hung down more vertically. Until we have more material, we need to keep these uncertainties at the forefront of our interpretations.
August 31, 2017 at 2:18 am
[…] 2014 but was formally named and published by José Carballido and colleagues just three weeks ago. While not technically the biggest known sauropod, Patagotitan is the only dinosaur in its class known from reasonably complete remains. The […]
March 9, 2018 at 9:01 am
[…] to ~120cm for Sauroposeidon, Puertasaurus, and Patagotitan, and 137cm for Supersaurus (more details here) – but they are massive. According to the table of measurements (yay!) in Tykoski and […]
April 3, 2018 at 5:39 am
[…] Some further thoughts on Patagotitan […]
November 14, 2019 at 9:41 am
Greetings,
As far as I know Argentinosaurus tibia is 155 cm., but there is a larger tibia of Alamosaurus, at least 165 cm long. Some claims this would be 183 cm: http://psdinosaurs.blogspot.com/2018/12/size-calculations-for-herbivorous.html. Here, the author alleges he made personal measurement of the tibia, but I don t know if this is accurate. If so, this could imply Alamosaurus may have a taller body than Argentinosaurus? There is any tibia from Patagotitan for comparision?
Thank you very much
Kind regards
Liviu Urziceanu
November 14, 2019 at 4:36 pm
Hi Liviu,
First, the 155 cm element of Argentinosaurus is not a tibia, but a fibula, and second, the allegedly giant tibia of Alamosaurus, actually is not a tibia (first-hand information).
November 14, 2019 at 5:37 pm
Asier, that is super-interesting info on the alleged giant Alamosaurus tibia. The description was a little brief and unconvincing for what would probably be the world’s largest sauropod known from fossils that still exist, if it was a tibia. Do you know if there is anything published beyond the initial description, or anything in the works?
November 14, 2019 at 6:16 pm
Hi Matt, not yet, the manuscript describing this specimen is on preparation by a pair of colleagues of mine from Mexico. This is what I can say by the moment, the information is confidential.
Nevertheless, next year will be published a book of mine and Molina-Pérez which estimates the size of all known sauropods. We’ve analyzed thousands of specimens.
https://www.amazon.com/Dinosaur-Figures-Theropods-Sauropods-Molina-P%C3%A9rez/dp/0691190690/ref=sr_1_3?__mk_es_US=%C3%85M%C3%85%C5%BD%C3%95%C3%91&keywords=larramendi&qid=1573755253&sr=8-3
November 14, 2019 at 7:05 pm
Hi Asier, thanks for the fast reply. I’m very glad to hear that the specimen from Mexico is getting a more complete description. I will be looking forward to that paper. And also looking forward to your book! Best of luck with the launch.
November 15, 2019 at 12:20 pm
Hi, Assier, Thank you very much for that very interesting info. Waiting for new data when they will be public and available.
November 16, 2019 at 4:04 am
Since the other giant femur referred to Argentinosaurus was found in isolation, and the fibula is from another specimen of Argentinosaurus, it makes me wonder what a 155 cm fibula calculates to on femur size.
Also, another stat that moves me when talking about giant sauropods is i’m pretty sure Puertasaurus is easily the owner of the widest dorsal and cervical vertebra that I’m aware of.
November 17, 2019 at 4:07 pm
The sauropods usually have a femur 1,5 to 1,65 longer than the fibula.
Opisthocoelicaudia has a femur of 139,5 cm
and a fibula of 83 cm ,has ratio of 1,68.
The Barosaurus with a femur of 144 cm and a fibula of 112 cm has a ratio of 1,285.
For a fibula of 155 cm, we can estimate the size of femur between 199 cm and 260 cm.
The difficuty of knowing whether all the bones attributed to Argentinosaurus belong to one or more.
July 20, 2020 at 1:37 am
Bump.
I’m not sure if this if this is the right place for this, but if it is, I happen to be the bearer of news of a humerus that may be the new longest complete titanosaur humerus.
This humerus, 178 cm long, was found in Thailand and described in an abstract.
“THE GIGANTIC TITANOSAURIFORM SAUROPOD FROM THE EARLY CRETACEOUS KHOK KRUAT FORMATION IN THE NORTHEASTERN OF THAILAND: A PRELIMINARY REPORT KHANSUBHA, Sasidhorn, Department of Mineral Resources, Bangkok, Thailand; POTHICHAIYA, Cherdchan, Department of Mineral Resources, Bangkok, Thailand; RUGBUMRUNG, Mana, Department of Mineral Resources, Bangkok, Thailand; MEESOOK, Assanee, Department of Mineral Resources, Bangkok, Thailand
Here is a preliminary report on the most recent find of an immense sauropod from the Early Cretaceous Khok Kruat Formation at Ban Pha Nang Sua, Nong Bua Rawe District, Chaiyaphum Province, Northeastern Thailand. In early March 2016, a local villager found an exposed of a sauropod sacrum at the bank of pond in the village. Later, geologists from the Department of Mineral Resources have investigated and started a systematic excavation during April-June 2016, discovered several post-cranial specimen from an individual sauropod comprising an axial skeleton (dorsal vertebra and sacral vertebrae) and an appendicular skeleton (a complete well-preserved right humerus, 1.78 m in length, a large dorsal ribs, a partial small rib, two pieces of partial right femur, pelvic girdle and many bone fragments), found associated with several isolated teeth which belong to Allosaurid and Spinosaurid theropods. Associated faunas include Hybodont shark Heteroptychodus sp. and crocodilian teeth. According to a preparatory study on humeral morphological features suggested that this new find possibly belong to a new taxon of giant titanosauriform sauropod dinosaur from the Khok Kruat Formation of Northeastern Thailand Lithostratigraphically, the sequence of the Khok Kruat Formation at the Pha Nang Sua dinosaur site, can be divided into nine units i.e., Units 1-9, having approximately 3.96 m thick. Dinosaur bones and fragments are found in the lower part of the sequence consisting mainly of reddish brown sandstones and siltstones with conspicuous climbing ripple lamination of Units 1-3. The top of this part consists of a thin layer (Unit 4) of calcrete paleosol. Sandstones from Units 5-9 display various directions of cross bedding and the bones are absent. This may indicate that the dinosaur bones are found in the crevasse splay layers and the channel and channelized sandstone are represented by the sandstones of the upper part of the sequence. The rocks are interpreted as having been deposited by the meandering rivers for Khok Kruat Formation in semi-arid to arid paleoclimate. More details on anatomical and phylogenetic studies and further excavations are essential to complete and compare with other vicinity sauropods. The measured section and faunal assemblage from Ban Pha Nang Sua dinosaur site is not only useful for lithostratigraphic and faunal correlations to other vertebrate sites. This new dinosaur locality will also shade light and figure out more understanding on sauropod evolution, distribution and paleoenvironment in this region.”
As for how big it might be, Phuwiangosaurus is probably a good animal to use as a base.
Greg Paul (2016) estimates Phuwiangosaurus at 19 meters and 17 tonnes, and according to Klein and Sander (2009) on its histology, the largest humerus from Phuwiangosaurus is 110 cm.
Assuming geometric similarity, this animal would end up at 72 tonnes directly scaled up.
Likewise, in terms of length, it’s about 30.5 meters scaled up as is, up to 34 meters with neck allometry applied. I hope this animal is published soon!