November 12, 2009


Scaled restoration of the giant titanosaur Puertasaurus by Nima Sassani, from the Art Evolved Sauropod Gallery

Get on over to Art Evolved and scope out the sauroponderous Sauropod Gallery. It’s brobdingnaginormous. I don’t want to seem biased, but there’s a lot of hot brachiosaurian action on display. I’m happy to say that the other clades are not ignored–diplodocids, dicraeosaurids, titanosaurs, mamenchisaurids, basal eusauropods, and even a basal sauropodomorph are all in the mix.

Normally my brachiosaurcentricity would lead me to steal one of the numerous brachiosaur images–perhaps the awesome parade of brachiosaurs that includes both Sauroposeidon and the Archbishop (!!)–BUT my laziness led me to choose another piece by the same artist, Nima Sassani. That would be the Puertasaurus reconstruction shown at top, which includes vertebrae and thus fulfills our titular mandate. That means I can stop writing now and get back to gawking. Go do likewise.

…oh, and don’t forget to stop by Dracovenator and congratulate Adam Yates on his new critter, Aardonyx. You’ll be hearing more about Aardonyx here at SV-POW! in the hopefully not-too-distant future. I can say no more for now…

58 Responses to “Sauropod-art-O-rama!”

  1. Should Puertasaurus have a big thumb claw? Or even any unguals at all?

  2. davidmaas Says:

    Hi! I’m the author of the Supersaurus (titled Diplodocus) at the gallery. Would love to have my work picked apart by knowledge people, and we’ve already had very interesting exchanges in that thread’s comments section (ie. body volume, position of body over legs etc.).
    So – as a word of encouragement, we welcome scathing crits as much as praise. Personally (and as time allows), I’d even welcome requests.

  3. Nima Says:

    Thanks a bazillion for posting my Puertasaurus, Matt! It’s downright awesomeness being ArtEvolved’s first visual ambassador to SV-POW ;)

    I’ve been working on this guy practically since I first saw the pictures of the dorsal on SV-POW (though I’m a bit shocked that Peter Bond of ArtEvolved resized the image so small… no worries, I’ll post a link to the full-sized version soon enough!)

    As for Tom Holtz’s question (BTW I’m also a Baltimorean…) The big thumb claw is part personal interpretation, and part based off of other species – notably Janenschia and Diamantinosaurus, which are at nearly opposite ends of the “derivedness” spectrum in titanosaurs. Short answer: they probably DID have a thumb claw, unless proven otherwise. Long answer: hold on – this’ll be a doozie…

    Basically my theory regarding titanosaur claws is that they DID retain the thumb claw in most cases, but modified it heavily. Many titanosaur hands are missing thumb claws and in some cases even the other phalanges. But I view this mostly as absence of evidence rather than evidence of absence. The fact that Diamantinosaurus includes complete thumb claw and phalanges, indicates these guys probably did not simply lose them – and it was a highly derived lithostrotian.

    The prevailing theory today is that since Saltasaurs have never been found with unguals or phalanges, that ALL titanosaurs, with few exceptions, lacked them. This simply doesn’t make sense to me. Saltasaurus are only the final most derived clade in a long tree of diverse titanosaur families, but sadly I see trends where artists restore EVERY titanosaur (even the really huge basal ones) like a scaled-up saltasaurus – short neck and all. This may be partially due to the popularity of Ken Carpenter’s saltasaur-silhouhette skeletals for Argentinosaurus, Paralititan, and “Antarctosaurus” giganteus in his paper on “Megasauropods” (which reduces their length estimates far too much, IMO), but there are similar errors by other authors as well. It’s a great paper in every other respect, but I question the accuracy of that particular depiction – it makes me wonder, what if artists assumed every stegosaur looked just like Stegosaurus as well? Very problematic, and not too plausible. All saltasaurs are titanosaurs, but not vice versa, and there are vast differences between saltasaurs and basal giant forms. Puertasaurus is less basal, but it also is not a saltasaur. So some more details need to be sorted out……..


    The classification of titanosaurs needs revision in this respect. This will help with the whole thumb claw issue. In most cladograms I’ve seen, every species more derived than Andesaurus seems to be thrown haphazardly into “lithostrotia” when in fact TRUE “lithostrotia” should include only Saltasaurs and Nemegtosaurs, and perhaps Ampelosaurus and a few other heavily armored types with long low skulls.

    In fact there appear to be THREE main superfamilies of titanosaurs – Andesauroidea, Lognkosauria, and Lithostrotia (based on the structure of the vertebrae, for which I am as ever grateful to the SV-POWsketeers for making such stunning photos accessible to the public).

    Andesauroidea includes Andesauridae, Argentinosauridae (which may include Paralititan), probably Huanghetitan, and PERHAPS Antarctosauridae and Argyrosaurus as well…

    Lognkosauroidea includes Malawisaurus and its relatives, as well as Lognkosauridae (Futalognkosaurus and Mendozasaurus) as well as PUERTASAURUS as a highly derived Maastrichtian descendant. Argyrosaurus may go here too, but until vertebrae are found, it’s anybody’s guess.

    Finally there is lithostrotia, which MAY include Antarctosauridae and Argyrosaurus, but is far more certain on things like Ampelosaurus, Rapetosaurus, Rinconsaurus, Saltasaurus, and perhaps even the seemingly un-armored Alamosaurus. These guys have low diplodocid-like skulls, usually heavy osteoderm armor, and tend to be on the small side. They are usually short-necked, though Alamosaurus and Rapetosaurus both break this rule into smithereeens.

    Finally back to thumb claws… They were definitely present in the more basal groups, so it’s likely that Andesauroidea had them. In Janenschia it’s fairly unspecialized. Then in Lognkosauroidea and Argyrosaurus, I predict that the thumb claw was modified into a bulky digging implement, thick rather than particularly sharp. The thumb metacarpal became flanged outwards to support the stresses on the massive claw. Puertasaurus, I think, likely enhanced this condition.

    Finally, lithostrotia…. well we have Diamantinosaurus, which retained a small thumb claw and seems to have reverted it to the old brachiosaur-like configuration. The first metacarpal is no longer flared out like crazy Argyrosaurus. Perhaps this happened to saltasaurs too. Perhaps they didn’t lost their claws and phalanges after all, but simply made them looser and anchored with more cartilage than earlier forms. The structure of the hands of Diamantinosaurus seems to show looser and more “cushioned” phalange attachments than Janenschia and certainly looser than the phalange attachments of Brachiosaurus/Giraffatitan brancai.

    Also, considering how massive and wide even small titanosaurs were relative to their length, more cushioned phalange joints were probably a very good thing. The thumb claw may have simply become a fixed spur in the most derived saltasaurs.

    Of course feel free to disagree, but I don’t see any solid evidence that any titanosaur lacked thumb claws, foot claws, or phalanges. And I don’t think it’s likely that any but the most derived titanosaurs could have lost them entirely. I know Darren posted a long time ago on sauropod hands on his blog, where he endorsed the claw-loss theory in saltasaurs. What’s everyone else’s take on this?

    I’m going to pose links to the full sized images, so by all means critique my art. I’d really appreciate any professional input here.

  4. Nima Says:

    Update: Peter Bond corrected the image sizing on the ArtEvolved Sauropod Gallery.

    Nevertheless I will still post the Puerta on my blog and leave a link here to the full size image. Actually I want to tweak it a bit more.

  5. Peter Says:

    Hey SV-POW readers,

    Nima Sassani’s FULL-SIZED images are now up on the ART Evolved Sauropod Gallery. Just click on each image to enlarge them. (If you could replace the Puertasaurus above with the high-res version, that would be appreciated, Matt. Thanks!)

    Just to let your readers know, ART Evolved is open to everyone! Join us if you are into recreating extinct life – our next Gallery is in January!

  6. Lognkosauridae and Lognkosauroidea can’t exist due to the absence of a Lognkosaurus. Excellent artwork though.

  7. Nima Says:

    Thanks! I hope this is the first of many titanosaur reconstructions to come.

    Though I wasn’t aware that every clade no matter how large or general needs a namesake genus… (or does it?) With titanosaurs it’s tricky because you have a superfamily (I used to say “titanosauridae” until it became clear that there was far too much taxonomical diversity for that)… interestingly, Lithostrotia doesn’t have a namesake genus either (nor do things like Coelurosauria, Carnosauria, Somphospondylii, tetanurae, maniraptora, marginocephalia – okay, well I admit THAT’s a stretch….) Whatever the rules are, they seem to have been broken a number of times…

    I’m not the ICZN, so I’m only using “Lognkosauria” and its derivatives (lol did I actually say Lognkosauroidea?!?!?! NOOOOOOOOO!!!!!) for the sake of simplicity. Whenever the titanosaur taxonomic maelstrom gets sorted out to any appreciable degree, I suppose then the ICZN will decide on a name for the middle of the three main titanosaur clans… I just hope they don’t pick “Futalognkosauroidea”…. that’s ten mouthfuls right there.

    Come to think of it, Futalognkosaurus is pretty high on my list of titanosaurs to restore. But I just might wait for Calvo et al. to publish their upcoming paper with ALL the measurements for the entire pre-caudal vertebra series first, so I can do the best restoration possible, with a lot less guesswork than went into this Puertasaurus (which is, after all, known from only four bones, two of which remain unpublished).

  8. The rule involves family-level taxa, so anything that ends with -idae, -oidea or -inae. Taxa with other endings can be named without an eponymous genus, such as Lognkosauria, Lithostrotia and the others you mentioned.

    One time it’s been broken lately has been Kellner’s Archaeopterodactyloidea. That’s an odd case, since by ICZN rules a superfamily including Pterodactylus must be called Pterodactyloidea (so Unwin’s Ctenochasmatoidea also fails), but that’s already in use for a larger clade. The way out seems to be to use a non-oidea ending for the clade in question.

  9. Mike Taylor Says:

    On zoological “family-group” names (those ending in -idae, -inae and -oidea): it is true that the ICZN mandates that families (and their coordinate subfamilies and superfamilies) must have a type genus; but of course that rank-based code says nothing about clades, which are governed (if at all) by the Draft Phylocode. Does the Phylocode have special rules about names ending in -idae etc.? That would be surprising (though I admit it’s been a while since I read it.)

  10. Mike Taylor Says:

    Matt, good choice: I too would have been torn between whether to feature the brachiosaur parade or the Puertasaurus tri-view, and I think that I too would have plumped for the latter. Nima, that is great to work: it’s good to see a titanosaur fattened up to this degree — they are too often restored looking like brachiosaurs, and even if it turns out that your work here is exaggerating the weird proportion, it will still have been a much-needed corrective.

    By the way, that broad torso is potentially indicative of an amphibious lifestyle. Just sayin’, is all.

    On the other hand, Nima, your thoughts on titanosaur classification are pretty bold in light of the lack of evidence that you’ve presented. Maybe you have more up your sleeve, but at this stage you seem, in the memorable words of a DML poster (David Marjanovic?) to be “doing cladistics with one character”. It’s certainly true that titanosaur phylogeny is up in the air right now, and also that the blithe use of the term “titanosaur” obscures a huge amount of generic diversity (100 genera or more) and of morphological disparity. It’s also true that some recentish contributions on the subject of titanoaur phylogeny have made the situation less, rather than more, clear. But on the positive side, good people are working on this problem. To name but two, Jeff Wilson has a long-term titanosaur thing going on, and Paul Upchurch’s in-progress mammoth sauropodomorph analysis is well represented for titanosaurs. Give it another five years and I think things will start to be clearer. (And perhaps the new, clearer ideas will not closely resemble what we think we know now.)

  11. Nima Says:

    Thanks for the input Mike! Great stuff. I figured, titanosaurs are generally very plump, (with a few exceptions), and based on the known fossil material Puertasaurus should be just about the plumpest one yet known, so it was the best choice of genus to illustrate this way (though based on the new photos from Juan Porfiri, Futalognkosaurus would certainly have come close).

    Based on what I’ve seen, the various lineages of titanosaurs seem to run the gamut of girth, from deep-bellied creatures like Isisaurus, which essentially look like highly specialized brachiosaurs, to super wide Lognkosauria like Puertasaurus and potentially even more squat-bodied creatures in Saltasauridae. I’m restoring Argentinosaurus now, and I figure that being a bit more basal, its torso probably has a more or less circular cross section… but even the most educated guesswork is still guesswork. At least Futalognkosaurus still has the ribs, and hopefully Calvo et al. will provide a decent multi-view skeletal…

    My theory of titanosaur classification may look radical, but there are other supporting facts “up my sleeve” that I didn’t mention (otherwise that comment would have been way too long even for my taste!)

    1. Andesauroidea – caudals are platycoelus, a fairly primitive trait… now we just need some Argentinosaurus caudals, lol! But the dorsals seem fairly similar (or course, I’d love to be corrected on this point by actually seeing the monograph or some other published material on Andesaurus instead of hunting down the odd photo all over the internet!) Also in some ways they still resemble robust brachiosaurs, at least in vertebral and pelvic structure, such as in Argentinosaurus. (I put Janenschia and Chubutisaurus in an even more basal group, Eotitanosauria). Skulls probably tend towards the boxy side of things.

    2. Lognkosauria: Now things get really interesting. Common traits? Tall, “shark-fin” neural spines on the cervicals. Also, cervical ribs are wide-gauge and splayed out – like honkin’ wide “Apatosaurus-splayed-out”. Indicative of a very deep and wide neck, perhaps heavily muscled – these guys may have been tearing through some very tough food. Not sure if Isisaurus or perhaps Argyrosaurus has any relation to this group, but they might. Solid vertebrae, not a whole lot of pneumatization besides under the zygapophses. Skulls are still usually boxy, as in Malawisaurus. Very wide rib cages and massively flared ilia. (i.e. “Puerta” and “Futa”). Also, every creature in this group seems to have a very small neural canal surrounded by solid bone (and thus a freakishly tiny spinal cord!) This next part is pure speculation, but based on the relation between spinal cord circumference and motor ability, Lognkosaurs may have been some of the slowest and most ponderous sauropods… making Brachiosaurus look like an olympic hurdle-jumper. Julius Csotonyi’s excellent Futalognkosaurus painting illustrates this point well.

    3. Lithostrotia: Many different forms here, but all have a similar body plan, hugely enlarged pubes, longer, more diplodocid-like skull, wide rib cage, armor plating is common (but probably not universal to the clade). Many species are small and short-necked (though at least a few smaller ones retain longer necks – such as Rapetosaurus, to which the one existing short-necked skeletal does anything but justice).

    Of course this scheme could be totally off, and that wouldn’t surprise me (and with Wilson and Upchurch’s research, even the conventional cladistics of titanosaurs could be shaken up once again). Perhaps there’s a need for some true metataxa to sort out this mess as well. I’d be very impressed to see a cladogram which sorts them out cleanly… It’s in a sense what I’ve been trying to do, just with a lot less data on my hands (though you guys have certainly helped with all the pictures you post up here!)

  12. Matt Wedel Says:

    Nima Sassani’s FULL-SIZED images are now up on the ART Evolved Sauropod Gallery. Just click on each image to enlarge them. (If you could replace the Puertasaurus above with the high-res version, that would be appreciated, Matt. Thanks!)

    Eeez done. Thanks for putting up the big versions–they’re much more appropriate for the subject matter!

  13. DD Says:

    meanwhile on the lighter side of things…

    presenting Nanofuertasaurus, “little lighter lizard”
    (kin to fire breathing dragons, note the neck posture!)

  14. William Miller Says:

    That thing is *immense*. (And so wide; it must weigh a *lot*.) Even Giganotosaurus only comes up barely above its knee.

    When an animal’s *vertebra* is as big as a man, that says something… Wow.

  15. Mike Taylor Says:

    Someone with some spare time on their hands (i.e. not me) should do a GDI on Nima’s Puertasaurus multiview and get a volume estimate (which can be converted into a mass estimate so long as you have a density figure that you’re reasonably happy with). Anyone up for the challenge?

  16. Very cool work Nima! One thing though, on your brachiosaur parade–didn’t a recent SV-POW explain that the size of Breviparopus was way overestimated, because the authors included the combined length of manus and pes into the ‘total print size’? Or is yours based on the downsized version(!)?

  17. Nima Says:

    Breviparopus? Hey, I was wondering when someone would notice that…. Well the short answer is, my Breviparopus is anything but downsized. I ended up with a much bigger Breviparopus than Matt Wedel did because I used different references that gave entirely different measurements.

    In the “Trackmakers” post, Matt was referring to some figures in Dutuit and Ouazzou’s original 1980 paper about a combined manus and pes set that was 115 cm long but only 50cm wide. That truly is impossibly small for a “record sized” sauropod, but I respectfully disagree with Matt’s conclusion that it was a diplodocus-sized animal because I don’t think Dutuit and Ouazzou got the figures correct to begin with.

    So I don’t base my Breviparopus on Dutuit and Ouazzou (1980) at all. Instead, I used Ishigaki’s scale figure of the prints published nine years later, which actually has a very clear scale bar that indicates that the pes prints ALONE are a meter long, and almost as wide. Here’s the URL for the figure:

    I also used Greg Paul’s classic 1988 paper on Giraffatitan, which contained pes width measurements for many sauropods (which, as Matt explained, are far more reliable than pes length measurements) – including Breviparopus.

    In Paul’s paper, the Breviparopus prints are recorded as 900mm wide – or just under a meter, which is EXACTLY the same size they are in Ishigaki’s scale figure. Neither of them supports Dutuit and Ouazzou’s claim of a little 50cm print, and I’m sure that the 50cm must have been an error considering how they publicized the thing as a record-breaker. So I’ll take the combined word of Paul and Ishigaki over Dutuit and Ouazzou’s – and here’s the main reason why:

    In Ishigaki’s figure, it’s actually the narrowest of the manus prints (third manus from the left) that is 50cm wide or so, not any of the pes prints. It may be that Dutuit and Ouazzou simply got the manus and pes measurements mixed up. Basically this guy had narrow hands and huge feet.

    Here’s another source that also lists only the manus as 50cm wide but omits the width of the pes! (though the pes is shown in diagrams elswhere in the text as about a meter wide based on Ishigaki):

    It appears so far that Dutuit and Ouazzou messed up (assuming they said the pes is 50cm wide instead of the manus; I don’t have their paper so I’m going entirely on what Matt said about it earlier). The Pes print in fact huge.

    If you assume Paul and Ishigaki are correct, you get a creature with feet bigger than either HMN SII (750mm wide) or the biggest Paluxy prints (~800mm).

    And THAT would result in a Brachiosaur that could easily be a hundred-footer. If you give it Sauroposeidon or Wealden Brachiosaur neck proportions, it could be even longer. And that is pretty much what I did.

    Of course I’d certainly welcome a debate on this from anyone here if there’s anything I missed or got wrong…

  18. Matt Wedel Says:

    Nope, I think you’ve got me dead to rights, at least on the size of the Breviparopus prints. I’ll update the other post with a pointer to this comment thread.

    However, the 750mm width for HM SII only accounts for the width of the foot bones. Throw on some soft tissue and it does not seem unreasonable that SII might have made tracks 900mm wide. At least, it seems that the 900mm track might have belonged to an animal 10% bigger than SII, but probably not 20% bigger. XV2 is 13% bigger. Not saying your recon is wrong, just pointing out that there is no evidence that Breviparopus was any larger than the biggest known individuals of Giraffatitan. And the general point that Breviparopus doesn’t hold a candle to the Broome and Plagne tracks is still valid.

  19. Nima Says:

    It’s true that Breviparopus was a good bit smaller than the spectacular gigapods that made the Plagne and Broome tracks, but a couple of things complicate the matter further…

    1. From the Ishigaki diagram it looks like there’s a bit of mud backflow for Breviparopus, as if the mud around the prints was very wet, and the edges fell inward, so the prints may be deceptively smaller than they should be. This is also noticeable with the foot claw prints, they’re classically shrunken from collapsed edges, as with many prints (though this is usually more common in theropod prints).

    IMO, a 68cm (or more liberally, 73cm wide) Giraffatitan foot won’t easily translate into a nearly meter-wide footprint with mud backflow and collapsed edges, even with the fleshy padding… but that’s just my preference in restoring these awesome animals.

    2. Another thing I noticed is that brachiosaurs have unusually small feet relative to their body size and mass (especially when compared to diplodocids and particularly Apatosaurines). So I think conceivably 90cm wide prints (which may have been a meter wide originally) could well belong to a 100+ foot brachiosaur in the 50 ton range (if it is indeed a brachiosaur). If it is some other sauropod, it may have been even longer due to tail length.

    HMN XV2 looks to be an 85-foot to 90-foot animal, so overall a 100-foot Breviparopus is not all that unrealistic, it may be in the Sauroposeidon range apparently…

    Another interesting thing I found is that the Breviparopus prints were originally dated to the Early Cretaceous and they certainly look like derived neosauropod prints (but too narrow-gauge to be titanosaurian). But now at least one paper dates them to the middle jurassic, which I find a bit odd… I wasn’t aware that Middle Jurassic sauropods had these sorts of narrow-gauge prints…

    The creatures that made the Broome and Plagne tracks were at least 50% longer though. Something at least the size of Puertasaurus (or at least my interpretation of Puertasaurus) would have to have made those.

  20. Mike Taylor Says:

    All known Middle-Jurassic sauropods are outside Titanosauria, which is the main group for which wide-gauge tracks have been proposed — so it’s not surprising that the MJ tracks you mention are narrow-gauge, like most others. There are some exceptions, though: for example, Day et al. (2004) reported MJ tracks of which some were wide-gauge, and could I guess have been made by brachiosaurs (yay!). All in all, I am not sure the wide/narrow dichotomy, and its association with non-titanosauriform/titanosauriform taxa, is as cut and dried as it’s sometimes been assumed to be.

    Day, Julia J., David B. Norman, Andrew S. Gale, Paul Upchurch and H. Philip Powell. 2004. A Middle Jurassic dinosaur trackway site from Oxfordshire, UK. Palaeontology 47:319-348. doi:10.1111/j.0031-0239.2004.00366.x

  21. Nima Says:

    Thanks for clarifying the MJ issue! :)

    The confusing part is that I’ve read both opinions on the (partially caved-in) Breviparopus prints; that they were from MJ AND that they were from EK. (from different sources of course).

    I was always under the impression that this was a very large Sauroposeidon-like creature that lived at roughly the same time as Sauroposeidon and Argentinosaurus (perhaps Charcharodontosaurus may have been its greatest enemy, just like Argentinosaurus had Giganotosaurus in patagonia). Yes, pure speculation… I don’t claim that it is valid but it’s certainly tantalizing and

  22. Nima Says:

    …and it seems that it’s at least plausible given what’s known about these mysterious prints. (my bad, posted that last one before it was done ;))

  23. […] doing the detective work. Rather than steal his thunder, I’ll point you to his explanation here. Point A above is still valid: Breviparopus was dinky compared to the Broome and Plagne […]

  24. I took up Mike Taylor on his suggestion of a GDI estimate for Puertasaurus using Nima’s multi-view reconstruction…

    The break-down in results are as follows:

    -Neck and head: 12,825 liters (~12.8 m^3)

    -Torso: 134,617 liters (~134.6 m^3)

    -Tail: 9,945 liters (~9.9 m^3)

    -Forelimb (each): 3,418 liters (~3.4 m^3); four separate limbs=13,673 liters (~13.6 m^3) (I had to use the forelimbs as a rough estimate of the hindlimbs because the hindlimbs are hidden by the forelimbs so I cannot measure them accurately; thus four times the estimate for the forelimbs to get a total for all the limbs)

    Total volume of head, neck, torso, tail, and four limbs: 171,059 liters (~171 m^3)

    Thus is one assumes a density of 500 kg/m^3 for the neck, we get a mass estimate of 6,400 kg (or 6.4 tonnes) for the neck. And if ones assumes a density of 800 kg/m^3 for the torso, we get a mass estimate of 107,680 kg for the torso (or ~107.6 tonnes). If one assumes a density of 900 kg/m^3 for the tail, we get a mass estimate of 8,910 kg for the tail (or about 8.9 tonnes). Finally, if one assumes a density of 1,000 kg/m^3 for the limbs (apparently the limbs are not pneumatic at all, so would be close to the density of water), we get a total mass estimate for the four limbs of about 13,670 kg or ~13.6 tonnes.

    Thus one get an approximate mass estimate for Puertasaurus of approximately 136.6 tonnes (136,660 kg). If I recall correctly, this is larger Matt Wedel’s mass estimate for Amphicoelias fragillimus of 122-123 tonnes.

    The caveat to this of course is that I think Nima made his Puertasaurus too wide from side to side on the torso. True, Puertasaurus was wide, but the widest point on the torso according to my calculation is over 6 m! I find that a little difficult to believe when the D2 itself is only 1.68 m wide. This means it is 3.57 times the width of the vertebrae.

    I am skeptical of this because of the fact that of the only multi-view titanosaur skeletal that I know of (Greg Paul’s Opisthocoelicaudia in The Complete Dinosaur), the width of the torso at the widest point is only 2.75 times the width of the vertebrae.I suspect that if it were correctly restored with a less wide torso, the true weight would be closer to 100-110 tonnes.

    Still, a very large animal it is.

  25. Actually, quick update: I “slimmed” the Puertasaurus‘s torso in my drawing to be much closer to the observed torso-width-to-vertebral-width ratio in Greg Paul’s Opisthocoelicaudia skeletal. It is still a slightly larger ratio, but much closer to that observed in Paul’s titanosaur skeletal.

    This changes the torso’s volume from 134,617 liters to 102,920 liters (or from 134.6 m^3 to 102.9 m^3). Assuming a density of 800 kg/m^3 the torso would mass about 82,320 kg, or ~82.3 tonnes.

    This changes the total weight from 136,660 kg to 111,300 kg (from about 136.6 tonnes to 111.3 tonnes). So, it is slightly smaller than Amphicoelias fragillimusmaybe.

  26. Matt Wedel Says:

    Zach, you magnificent bastard, I don’t know whether to kiss you or kill you! These two comments just pre-empted most of a very long post I had in the works. Good work! :-)

    My conclusions were very similar to yours, especially as regards torso width. Compare the Puertasaurus to this cross section of Diplodocus. I know that at least some titanosaurs were quite a bit wider-of-beam than diplodocids, but try drawing the ribs on the front view of the Puertasaurus and the results don’t look like any known sauropod.

    Also–and I don’t mean to harsh on Nima here, because it is a really attractive life restoration and this discussion wouldn’t be possible if he hadn’t put in so much detail–the scale is off. The 4m scale bar in the picture is 139 pixels tall, giving 34.75 pixels/meter. The total length of the cervical is 52 pixels or 1.49 meters, and the total width of the dorsal is 74 pixels or 2.11 meters. The actual measurements are 1.18 meters for the cervical length and 1.69 meters for the dorsal width. Both of those measurements are almost exactly 80% of the measurements depicted in the life restoration. So the internal scaling is consistent, the darn scale bar is just too small. So you can multiply the mass estimate you got by 0.8*0.8*0.8=0.512, or in other words, cut ’em in half. That gets Puertasaurus back down into the realm of mortal sauropods. OTOH, it is sobering that an animal just 20% bigger would mass what you estimated.

    Now I gotta find another example to use in my graphic double integration tutorial… ;-)

  27. Good thing I didn’t use the scale-bar, then (I noticed it was off)…I took your advice from your MMYD post (about the Futalognkosaurus paper and not trusting scalebars!), and I scaled everything off from the cervical that was drawn in the picture, which is 1.18 m between the prezygs and post-zygs.

    I don’t think you have to worry about my pre-empting comments for your post on GDI, as I doubt very many people will see this comment, seeing that it is on an old post.

    BTW, I also did a new and hopefully more accurate GDI on Giraffatitan. Suffice to say, I get a volume of 44,534 liters. Which means, even using a really low density for the neck (say, 400 kg/m^3 and 800 kg/M^3 for the rest of the body), you still get a 30-tonne animal.

  28. William Miller Says:

    Waitaminute … then … if that 110-ton thing is not using the scale bar – then Puerta really was over the 100 ton mark, in ranges previously reserved only for semi-legendary monsters like Amphicoelias and the Broome trackmaker?

    WOW. Titanosaurs were well named…

  29. I do have to say this: at least in the “length” dimensions, Nima’s reconstruction is nearly spot-on.

    I estimated Puertasaurus dimensions using the now very completely measured material of Malawisaurus. Scaling of Malawisaurus to Puertasaurus on the base of the D2 and the C9, I get a neck length of 10.86 m, a torso length (i.e., the dorsals and sacrum)of 8.87 m, and a tail length of 11.98 m. This combines for a total length of 31.71 meters (~104 ft).

    Judging from the cervical 9 in Nima’s drawing, his torso is 8.85 m, the tail is 12.53 m, and the neck is 10.95 m. So his tail and neck lengths are slightly exaggerated, but his torso length is right on.

    Taking the exaggerations into account, the neck “should” be about 6.35 tonnes and the tail “should” be 8.55 tonnes. All in all, a total weight of for Puertasaurus of 110.82 tonnes seems reasonable.

    We can further corroborate this mass estimate another way: In an earlier post by Matt Wedel(see here), the length for Alamosaurus was given to be 15.8 m, with a mass of 15 tons (is that short tons or metric tons??).

    Since Puertasaurus is 31.7 m, it is 31.7/15.8=2.006…about two times longer than Alamosaurus. Since volume is the cube of length, we have 2^3*15=8*15=120 tons. Assuming these are short tons, 1 metric tonne is about 90.72% of a short ton, so 120*.9072=108.86 metric tonnes, which is reasonably close to my estimate of 110.82 tonnes using Malawisaurus as a model.

  30. don lessem Says:

    Hey Matt, et al:
    I’m in need of size estimates and bones found which I can’t find for about 50 dinos from the entire roster for a kids’ dictionary – anyone willing to give them? Here’s but a few (if they’re valid taxa at all).
    Pitekunsaurus – pronounced, meaning ,size, date
    Platyceratops – size and fossils
    Pookyangosaurus meaning and size
    Pycnomemosaurus – name, meaning, size
    Qingxiusaurus – all
    Qinlingosaurus – size
    Ruyangosaurus – size, date, fossils
    Shenzousaurus size
    Shidaisaurus size
    Sinosonasus fossils
    Shidaisaurus – size
    Similacaudipteryx fossils
    Tastavinasaurus –
    Tavierosaurus size
    Tehuelchesaurus – bones
    Thecodontosaurus – what fossils
    Zapalasaurus – size?

  31. Nima Says:

    Thanks guys for all your comments and calculations! It’s been a while since I looked at this post and Zach, I’m truly AMAZED by your dedication to the numbers!

    It’s true, I did overscale slightly with the bones, but this was for two reasons: one, to compensate for crushing, erosion, and potential surface damage (the fossils look pretty worn down in practically every photo I’ve seen), and TWO, I wanted to depict a SLIGHTLY larger Puertasaurus (i.e. what the type specimen might have looked like if it had lived, say, five or ten years longer).

    The biggest danger of incorrect scaling, of course, is that it may be taken as fact. So even though it’s not off by that much, I’ll try to keep the scaling more conservative in my future giant sauropod restorations.

    Right now I’m working on the seven biggest titanosaurs, I already have finished Argyrosaurus and it’s been up on ArtEvolved for a while. Argentinosaurus and Paralititan are also in the works. For those I’m using a straight 1/100 scale ratio on paper to keep the scaling reliable and consistent; i.e. 1cm on paper = 1m in real life. So hopefully I should have them finished soon and I’ll inform everyone here when they’re done.

    The main thing driving me now is that I’m not satisfied with any of the current depictions of these often fragmentary record-breakers, and I want to be the first to draw them accurately given the material and what’s known about their relatives.

  32. Thanks for your comment Nima. Like I said, your Puertasaurus is very accurate in the length dimensions, in fact my comment about the neck “should” be this long and the tail “should” be this long only meant if one was scaling it off Malawisaurus and so should be taken with a grain of salt, it is certainly possible that the neck and tail were slightly longer, like you had drawn it. It is certainly within the realms of possibility, anyway.

    The only thing that bothered me was the width of the torso, which I think is still too wide as restored. you restored it, according to my calculations, 6.04 meters wide at the widest point, whereas I “slimmed” down your reconstruction to 4.42 m wide at the widest point, based off of the proportions in Greg Paul’s Opisthocoelicaudia skeletal. The truth is Opisthocoelicaudia is fairly wide even for a titanosaur. So that is why I would not restore Puertasaurus with a maximum torso width any greater than 4.5 meters in total width at the widest point.

    The good thing, as Matt said, is that the scaling was internally consistent, just that the scale-bar was too small in proportion to the drawing, see his comments for the specifics. I was recently about to do a GDI for Rapetosaurus based off of Mark Hallett’s skeletal (the only one in existence, really, except for near-verbatim copies of it) in the new monograph of Rapetosaurus that was recently out, the only thing is that the internal scaling is inconsistent, terribly. For example, going off the skeletal , you get a femur length of over a meter, whereas the length listed in the table is about 65.7 cm, or .657 m. Now, when you compare the tibia and fibula, you get different lengths also, but they are not even off by the same ratio! Very frustrating. From the table, one gets an approximate length for the cervical series C3-C16 of 320.1 cm, while the length of the dorsal series D3-D10 is 81.5 cm. This means the neck is approximately 3 times the length of the dorsal series. However, in the skeletal, they are nearly the same length! *Sigh*

    So at least your drawing is more accurate than can be said for some peer-reviewed paper’s figures.

    I am definitely looking forward to those titanosaur drawings!

  33. Nima Says:

    Thanks Zach! I’m making sure they come out top-notch.

    I made the neck and tail longer (mostly the neck) because my theory about titanosaurs is that the bigger they got, the longer their necks got proportionally (the cervicals also grew to more or less uniform length, post-axis anyway). This trend towards mega-necked gigantism also occurs in other sauropod groups (Supersaurus and Sauroposeidon come to mind)… Furthermore, there’s a bit of a secret about the neck of Puertasaurus that makes me think it really must have had a very long neck to fully take advantage of one of the neck’s oddest features. In other words, this guy likely DID out-neck Malawisaurus and most smaller titanosaurs in terms of proportions.

    As for the width of the Puertasaurus torso, I intentionally made it that wide because I believe it WAS that wide. Puertasaurus has the proportionally widest dorsal of any titanosaur – or any sauropod period! Both in terms of with:height ratio AND centrum width, AND it’s got the biggest zygapophyses.

    So, long story short, a torso width of 7-8m is not THAT crazy for such a crazy-proportioned animal. Also, Opisthocoelocaudia was a Saltasaurid (or close to it) and even though those guys are fairly squat, they don’t have the crazy-wide proportions of Lognkosauria. So Opisthocoelocaudia may not be the best model for Puertasaurus.

    Basically in my view, Titanosaurs sort out into three main “family clusters”:

    -Basal types (Chubutisauridae, Andesauridae, Argentinosaurus and its sort…)

    -Transitional types (Lognkosauria, Argyrosauridae, Malawisaurus, etc.)

    -Late types (Lithostrotia: Antarctosauridae, Saltasauridae, Nemegtosauridae, Ampelosauridae)

    Puertasaurus is essentially a very late-evolving descendant of a “transitonal” family (Lognkosauria) which actually departed from the “main line” of titanosaur evolution towards lithostrotia, i.e. it’s a major side branch. So scaling it to the proportions of Opisthocoelocaudia, a rather small sauropod from Lithostrotia, is NOT all that accurate IMO. They are from two long-divergent lineages.

    As for Rapetosaurus – yes, Hallett DID make a bunch of very odd scaling errors. He even left out two dorsals, making the torso too short. He also made the neck vertebrae far smaller and shorter than they actually are, and exaggerated the thickness of the limb bones. The mounted juvenile skeleton shows that this guy had a long torso and a VERY long neck. It’s not your typical pocket-size mini-necked Saltasauroid.

    But I REALLY want to see that Opisthocoelocaudia Greg Paul skeletal. I didn’t even know he did titanosaurs! (well I saw his Argentinosaurus silhouette, but that barely counts…) Send it over if you can…it would be a great reference for future titanosaur drawings after I’m done with the biggest ones.

  34. Actually Nima, Hallett did not leave out two dorsals, as Rapetosaurus is only reported as having ten of them. The first dorsals two were co-opted to be cervicals, according to the new monograph, which leaves Rapetosaurus with a total of 17 cervicals! So in actuality, Hallett left out two cervicals in addition to making the neck too short. I am currently working on a skeletal of Rapetosaurus to correct the matter.

    Your assertion that Puertasaurus was more wide-bodied than other titanosaurs is not true:

    *Malawisaurus has a C9 29.5 cm wide and a D2 43.5 cm wide, or basically the C9 is 67.816% of the width of the D2.

    *Whereas in Puertasaurus, the C9 is 140 cm wide, and the D2 is 168 cm wide, meaning the C9 is 83.333% of the width of the D2. This actually means that the dorsal vertebrae was narrower in proportion to its neck than in Malawisaurus.

    Thus a torso width of 6 m+ for Puertasaurus is unwarranted in light of the evidence. A more reasonable torso width of less than 5 m is much more likely. A torso width of between 4-4.5 m at the widest point seems most probable.

    As far as I know, Puertasaurus has not been included in any cladistic analysis so your classification of it in the Lognkosauria is speculative, although not implausible. It probably is not a saltisaurid, so comparing it with Opisthocoelicaudia may not be the best comparison, but even so, comparing it with Malawisaurus (which is apparently a basal titanosaur, judging from the skull), a torso of a width exceeding 4.5 m is very unlikely IMHO. Thus, a torso width of 7-8 m, as you suggested is pretty crazy considering the evidence. It should be noted that it was the derived saltisaurids that are known for widening their torsos, not the basal or ‘intermediate’ titanosaurs of which Puertasaurus is likely one.

  35. Nima Says:

    All the same, Puertasaurus has insanely wide zygapophyses compared to Malawisaurus, Mendozasaurus, and even Futalognkosaurus (at least based on the pictures I’ve seen)… and these appear to be its closest relatives based on vert structure alone. They are all wide-bodied, and have wide, thick loop-like cervical ribs that to some extent seem to imitate those of Apatosaurus.

    But Puertasaurus may have had the widest body of all IMO. The fact that its dorsal-cervical width ratios are not as extreme as Malawisaurus does not prove anything about the rib cage. It merely shows that Malawisaurus had a proportionally narrower NECK.

    In my restoration I made it clear that Puertasaurus had a wide neck as well as a wide torso. The neck is extremely wide and also very flattened (even when you factor out the crushing) compared to other sauropods. The cross section must have been very “oblate”. Furthermore, you didn’t list the LENGTH to width ratio of any vertebrae. THAT is the real clue to how wide a rib cage can be.

    Puertasaurus’s dorsals were not only the widest on record, but also the MOST widened relative to their length. That in itself indicates a very wide torso, and when you also consider that most of the vert’s width is due to those huge wing-like zygapophyses (which of course anchored the ribs), then the only logical conclusion for having such gigantic buttressed zygapophyses is that this creature had to support an insanely wide rib cage. These are not delicate little structures, they are giant wedges designed to anchor a lot more weight than the more rectangular or rod-shaped zygapophyses of nearly all other sauropods. Even Malawisaurus’ dorsals can’t match it in this regard.

    So an 8m wide rib cage is not too outlandish in that regard, but to be conservative lets say 7m (and to make up for any *possible* scaling discrepancies). Still, in terms of proportions and curvature of the proposed ribs, this seems perfect for the huge wedge-shaped design of the zygapophyses.

    Saltasaurids also had squat torsos. This is simply a case of convergent evolution, their dorsal vertebrae were not all that similar to Puertasaurus. Puertasaurus appears to be a highly derived Lognkosaurian, which appeared much later than other members of the group. Earlier ones like Mendozasaurus likely had narrower torsos, but likely still proportionally wider than the basal titanosaurs like Argentinosaurus (which itself had a pretty hefty barrel-like torso, but proportionally narrower vertebrae than Lognkosauria).

  36. Actually, Nima, it’s the diapophyses that are insanely wide, not the zygapophyses, but I won’t hold that against you. You said, “The fact that its dorsal-cervical width ratios are not as extreme as Malawisaurus [sic] does not prove anything about the rib cage. It merely shows that Malawisaurus [sic] had a proportionally narrower NECK,” OR it means just what I said, that the the torso is not as wide proportionally to the neck as in Malawisaurus. At any rate, it means the same thing, the torso was not all that extra wide in comparison to its other body members.

    You claim that the neck is very flattened, but the fact is you restored it without restoring the full neural spine height as restored in the paper. The neural spine apparently suffered some erosion and/or damage that reduced the neural spine height.

    You also said, “Furthermore, you didn’t list the LENGTH to width ratio of any vertebrae. THAT is the real clue to how wide a rib cage can be.” How so? Is the width of the rib cage somehow related to the length of the dorsal vertebrae? Please find a reference to back that up, as you offer no support for that blatant assertion. You continued, “the only logical conclusion for having such gigantic buttressed zygapophyses is that this creature had to support an insanely wide rib cage.” Oh really? What about the possibility that it may have had thicker dorsal ribs that were not pneumatic? As animals grow larger, they have to thicken their bones in order to compensate for the increased volume and hence weight. As Tyrannosaurs got bigger their femurs got proportionally shorter and thicker NOT proportionally longer, might it be the same thing with Puertasaurus ribs and diapophyses? Might it be that they got thicker and denser, rather than growing out sideways? Just because the diapophyses grew large and winglike does NOT mean that the ribcage got proportionally wider to its vertebrae, and you have given no hard evidence to support that.

    You say that a 7-8 meter wide ribcage is not unreasonable, but it IS considering that the widest part of the ribcage in your drawing above is about 6.04 m-6.19 m depending on if you measure using the aerial view or frontal view. Even this is WAY too wide based on the evidence considered, much less considering a ridiculous width you promote of 7 meters (even 8!!). As Matt Wedel commented above, in order to get a ribcage that wide you have to draw a rib configuration like no other sauropod, titanosaur or not, which is both highly speculative and unparsimonious.

    What we do know is that the vertebrae got wider in proportion to their centrums, but this does not indicate anything about changing proportions in ribcage width. Your reasoning here is a total non sequitur, the premise (i.e., the vertebrae is wider in proportion to the centrum width) and the conclusion (i.e., thus the ribs must have also been proportionally wider) are unrelated.

    You said that here Puertasaurus converged with saltasaurids, but before you said that Opisthocoelicaudia (which is a saltasaurid) was not a valid comparison (one that I had used). Now all the sudden it is okay? Like I said way back at the beginning of this discussion, in Opithocoelicaudia (based of Greg Paul’s skeletal), “the width of the torso at the widest point is only 2.75 times the width of the vertebrae.” Thus, 2.75*168=462 cm, or 4.62 meters. Thus the widest point of the torso should be about 4.62 meters in Puertasaurus. A torso width of 6 meters is 3.57 times the D2 width, 7 m is 4.166 times the width of the D2, and 8 m is 4.76 times the width of the D2. This is way over and above a 2.75 ratio. And yet, no logical qualitative or quantitative argument you have given supports these widths which seem to have come out of thin air (i.e., WHERE, oh where, are you getting these 7 meter or 8 meter figures??? Also, how does one conclude that because the vert width to centrum width ratio is increasing does one conclude this has anything to do with the width of the ribcage???)

  37. Mike Taylor Says:


    (I’m talking to Nima and Zach),

    You’re both making good and interesting points in this discussion, but you seem to be getting increasingly snarky. Please reverse this trend: I’m keen to see where your discussion goes, but I don’t want to see it drag down the high level of discourse we’ve become accustomed to. That’s all, thanks for listening.

  38. Sorry about that Mike (and Nima), I’m mostly the one to blame for the snarky tone. Won’t happen again.

  39. Nima Says:

    1. The length of the vertebrae: My main argument here was that Zach didn’t include the lengths of Malawisaurus vertebrae, thus it’s impossible to calculate their average length-to-width ratio given what he’s posted. I could look them up myself, but I’m just lazy I suppose… I have seen the photos and they look nowhere near as laterally widened as those of Puertasaurus. With Puertasaurus we see an insanely wide but short dorsal. So OVERALL it’s reasonable to assume that the torso itself is insanely wide relative to its length – whereas the torso of Malawisaurus, I suspect, would be more moderately proportioned.

    2. I measured the rib cage at around 7m. I scaled all the measurements with a ruler and in any case, I tried to be conservative with the width. I said this is the first high-fidelity restoration of Puertasaurus, and so far the best. I didn’t say it’s perfect, because it can never be when you only have a couple of the bones! They can give some clues but basically those clues (and what they mean in related animals) are the best you have to go on. For example, in more complete titanosaurs, widened Zygapophyses roughly correspond with wider rib cages. This is the case in all titanosaurs with decent rib material preserved, compare it to Giraffatitan with its narrower, slab-sided torso (which is still pretty wide compared to diplodocids).

    3. My point on Saltasaurs and Opisthocoelocaudia: I admit this got a little confused. Opisthocoelocaudia is not “everySaltasaur”. It has a more barrel-shaped rib cage than some. Saltasaurs are actually quite a diverse group and it’s hard to tell whether some species belong in there at all. But all that aside, YES, in terms of wide rib cages, Puertasaurus did converge on Saltasaurs. But that doesn’t contradict the apparent fact that it took the widening further than one certain Saltasaur did. Even the much earlier Futalognkosaurus had very wide dorsals, and what looks to be a big, robust rib cage. To this widening trend seems to be pretty common in titanosaur evolution.

    3. Widening the dorsals simply to support non-pneumatic ribs doesn’t make a whole lot of sense IMO. Comparing it to T. rex limbs is also a bit odd, because T. rex limbs had to withstand the stress of a running bipedal animal, whereas each rib of a giant titanosaur was not directly supporting half of the creature’s weight! I don’t claim to be an expert on titanosaur rib pneumaticity (this seems a great area for research though!) but from what I know, members of a clade didn’t necessarily decrease pneumaticity but they simply altered its form. Titanosaurs actually DO have pneumatic vertebrae, only most of it is internal camellae rather than big scooped out external camerae like in brachiosaurs.

    Rib pneumaticity has for obvious reasons always been internal. So I’m not sure this would have disappeared. And in any case, if this was the real reason for widening the diapophyses, then how does one explain the lack of any significant attachment point for the lover part of the “rib fork” on the centrum? This creature does not appear to have unusually dense ribs. Furthermore, that theory also does not explain why the entire vertebra is so squat and even the neural spine is short and wedge-shaped.

    It’s one of the beautiful melodies of nature that an animal’s whole torso seems to be constructed the same way as its dorsal vertebrae: with brachiosaurs, the dorsals are tall, with long neural spines and diapophyses with pronounced laminae and flared ends, light but reinforced to handle a vast load – and the rib cage is just as you would expect – grand, large, and most of all deep, just as the vertebrae are tall. With diplodocids, the vertebrae are light, modestly proportioned, and smoothly transition in shape from one to the next, and their diapophyses are not very wide, flaring of vertebral extremities is far less than in brachiosaurs, which indicates a more lightly built creature per meter of length. And fittingly, they also have narrower rib cages and smaller torsos proportional to total length than brachiosaurs. Even bulkier apatosaurs still fall within this spectrum.

    With titanosaurs, the pattern changes yet again. Dorsals get squatter and wider, as do rib cages. Including that of Opisthocoelocaudia, though it’s not the most extreme example. Rapetosaurus is far more squat. It’s as if the whole design philosophy of these animals changes. As the entire torso gets wider and squatter, so does the dorsal column, including its neural arches and neural spines. Laminae connect processes and there is more fusion and less separation of them. The dorsal column becomes wide and buttressed to support ever wider rib cages. This trend gradually increases from the most basal early titanosaurs, to the most derived, including Maastrictian forms like Puertasaurus, and also the Saltasaurs. Both are derived in their own ways.

    The end of the Cretaceous basically sees sauropod rib cages getting very wide and flat-topped like those of ankylosaurs. Titanosaurs seem to have adopted the “built like a tank” design, not least because many of them were armored.

  40. Nima Says:

    whoops, last point was 4, not 3.

  41. In Malawisaurus the D2 centrum is 130 mm long, 159 mm wide, 110 mm tall, total vertebral width is 435 mm+, and the total vertebra height is 308 mm. The C9’s centrum is 415 mm long, 120 mm wide, 85 mmm tall, the total vertebral width is 295 mmm and the total height is 200 mm.

    *D2 centrum width to total vertebral width ratio: 159/435=0.365517 (or the vertebra is 2.735 times the centrum width)
    *C9 centrum width to total vertebral width ratio: 120/295=0.40678 (or the vertebra is about 2.458 times the centrum width)

    In Puertasaurus the D2 is 168 cm wide, about 41.5 cm long, the centrum at its widest point is about 56.62 cm (scaling from the photograph based on the known total width) or 58.51 cm (depending if you measure based on the condyle or the cotyle, the centrum height is about 39.64 cm or 41.52 cm (again depending on whether you measure off the cotyle or condyle). The total height of the vertebrae appears to be 107.59 cm. The C9 on the other hand is 118 cm long between the prezygs and postzygs (or 105.57 cm for the restored centrum length), and is about 130.42 cm between the diapophyses, the centrum is 41.4 cm wide, and 18.63 cm tall.

    *D2 centrum width to total vertebral width ratio:
    56.62/168=0.337024 or 58.51/168=.34827 (or the vertebra is either 2.96 times or 2.87 times the cnetrum width)
    *C9 centrum width to total vertebral width ratio:
    41.4/130.42=0.317436 (or the vertebrae is 3.15 times the centrum width)

    So indeed, the total vertebral width in proportion to the centrum for the D2 in Puertasaurus is wider than in Malawisaurus (either 2.96 or 2.87 compared to 2.735). So, if Malawisaurus had a centrum 56.62 cm wide or 58.1 cm wide, going off the ratio, we’d expect the total vertebral width going off of its own proportions of 56.2*2.735=153.7 cm or 58.1*2.735=158.9 cm. Going off the Puertasaurus proportions we get 56.2*2.96=166.35 cm or 58.1*2.96=171.976 cm (with the true width of 168 cm falling right inbetween).

    This means that using the Malawisaurus proportions, the vertebrae width is at minimum predicted to be 153.7 cm, so 153.7/168=0.91488. So basically in Puertasaurus the ratio is about 9.5% wider than in Malawisaurus (1.095*153.7=168.3).

    Before I quoted that based of Greg Paul’s skeletal, the maximum torso width is 2.75 times the vertebra’s width. This means, then, that in Puertasaurus we can expect this ratio to be exaggerated by about 9.5%, so 2.75*1.095=3.01125. So, 168 cm*3.01125=505.89 cm, or about 5.05 meters wide. This is about 1 meter less than depicted in your drawing, and 2-3 meters less than you say. At any rate, going off all the evidence we’ve seen, your torso is still too wide.

    As for the measurement of the torso in your drawing above, I printed it off, and the C9 is exactly 8mm between the prezygs and postzygs, which are known to measure 118 cm in between. That means that for every millimeter on the paper, it represents 14.75 cm in real life. I measured the maximum width on the torso in your drawing to be either 41 cm or 42 cm depending on whether one goes of the aerial or frontal view. 41*14.75=604.75 cm, 42*14.75=619.5 cm, or a max. width of between 6.04 meters and 6.19 meters.

    As for the non-pneumatic ribs comment, I never said the ribs definitely were not pneumatic, what I did do was raise was the possibility that they were not pneumatic (hence my question mark). I also raised the possibility that they more dense or proportionally thicker. In any case, the point was to show that one does not have to automatically assume a ginormous-ly wide ribcage to account for the fact that it had wide diapophyses, as there may in fact be other possible explanations. As for the T. rex bone comparison, my point was not to say that the ribs of Puertasaurus are analogous to T. rex legs, but to show biomechanics can change dramatically (or even jsut significantly) at such a large sizes. Thus, the reason for large winglike transverse processes may have to do more with Puertasaurus being extremely large than have anything to do with the relative width of the ribcage.

    As for the ‘beautiful melodies of nature that an animal’s whole torso seems to be constructed the same way as its dorsal vertebrae’, that’s fine, but does not change the mathematics I show above, namely that the width of the torso at the maximum point on Puertasaurus was around 5 meters, not 6, not 7, and definitely not 8 meters.

    You have shown no hard quantitative evidence to support the width(s) you advocate for Puertasaurus, and thus I submit to you that the torso width you have portrayed for Puertasaurus is very, very unlikely. Note that this is not a personal criticism of you or your artwork, which is top notch, but of this particular instance in which the scientific support for the torso width portrayed in your reconstruction is found wanting. This greatly affects mass estimates, which would not be possible to do without your highly detailed drawing for which I am grateful (and for which otherwise this conversation would not be possible). Using your recon as-is, one gets a total mass of over 136 tonnes, while using a ‘slimmed’ version, one gets a mass of about 110-111 tonnes, which is much closer to the predicted mass of 108 tonnes using Alamosaurus as a model. Thus it appears your recon is too wide.

  42. Nima Says:

    I also prefer a mass estimate of 110 tons. Perhaps my “recon” may be a bit wide, but it’s good change IMO from titanosaurs that are drawn too narrow. Perhaps the density may have been less with creatures so bulky, to keep the weight down, but it’s hard to know. All the same, we can not all be mathematical geniuses. Given that the vast majority of the skeleton is not known, however, I am reluctant to stick fervently to any mathematical model for Puertasaurus. In addition I scaled the vertebrae a bit larger than the official published measurements to account for erosion and crushing (both vertebrae appear to be substantially eroded and smoothed down) so the actual width including ossified cartilage may have been 10-20% greater. Then, we see a rib cage width of 7m is certainly not outside the realm of possibility.

    In addition, it may be that so much change had occurred between the time of Malawisaurus and the time of Puertasaurus, that their proportions were nothing alike! Once again, it’s hard to tell. So until more remains of Puertasaurus are found, I’m not all that eager to revise this restoration. But if more remains turn up and refute my theory then by all means I will be glad to revise it. To me, bones are usually far more reliable than numbers alone (not that I’m dismissing YOUR numbers, but it would be nice to see what shape this creature really was instead of just going off numbers). Just take a look at all the best early mathematical estimates for “Ultrasauros” when it was discovered – same with Seismosaurus, Supersaurus, etc… and they were all far off. On the other extreme, I think Ken Carpenter’s “dwarfed” estimates for the dimensions of Argentinosaurs, Paralititan, and A. giganteus are far too small. And we are definitely in the early stages of even beginning to understand Puertasaurus. So a purely mathematical approach depends on your interpretation of the actual size/shape of the animal, and causes a lot of room for error.

    That’s unfortunately the territory you have to step into with most of these enigmatic super-titanosaurs. So little of them is known that making a good restoration also takes some talent for guesswork. Obviously the whole thing is not guesswork, the described material is drawn and even restored to some extent and the reconstruction was measured and cleaned up several times, and compared with other known titanosaurs, which is far more than can be said for previous restorations of this beast.

    There’s a good reason why most artists won’t even touch these giants.

    Which reminds me Zach, didn’t you once do a quick rear-view sketch of Puertasaurus on DeviantArt? I’m interested to see if you’d make a more detailed restoration. If you view mine as flawed (albeit in areas we don’t even have hard fossil evidence for) then by all means make a better one and post the link. There are other aspects of this picture I’m not totally satisfied with since completing it, but rather than just arbitrarily change things up, I’d like to have some alternate images to better inform any debate.

    *BTW I know you like my art so I’m not taking any of this personally, and thanks for all the compliments. I myself view Puertasaurus as one of the widest if not THE widest titanosaur, so for me any good restoration of it should be unusually wide, even by the standards of earlier titanosaurs.

  43. “Perhaps my “recon” may be a bit wide, but it’s good change IMO from titanosaurs that are drawn too narrow.” Right, Hallet’s multiview restoration restores Rapetosaurus way too narrow in cross section, more narrow than observed in even some diplodocids(!)–one just has to look at the museum mount to see the difference–so I understand the intent to break down these ‘stereotypes’, at the same time I think we should be more ‘conservative’ in our reconstructions, otherwise some of the ill-advised public may treat it as fact, rather than speculation. For me, a lower-bound estimate of mass or length is always better than a more speculative over-estimate that may turn out to be with a significant error. This goes back to the estimates of Seismosaurus being over 150 ft long and such, which while at the time probably seemed reasonable, but more evidence shows that this was way too long. In a similar vein, I’d rather find in the future with better fossils that my estimate for Puertasaurus (or really, any large sauropod or dinosaur) was an underestimate, rather than an overestimate.

    You said you restored your D2 wider to account for crushing and other fossilization-related distortion. This may be where our estimates differ, then. Would you mind saying how wide you restored it in your reconstruction above?

    You said (in part), “To me, bones are usually far more reliable than numbers alone (not that I’m dismissing YOUR numbers, but it would be nice to see what shape this creature really was instead of just going off numbers)…So a purely mathematical approach depends on your interpretation of the actual size/shape of the animal, and causes a lot of room for error.” Right, but all of my measurements are based off the bones in the descriptions. Without math, we could come to no reasonable mass or length estimate at all. Furthermore, I do do some paleoart, so I am not discounting the shape of the bones or anything else or the visual ‘intuition’ aspect to this. I realize that as a mathematician all mathematical models are subject to errors, the goal however is to minimize those errors and have an estimate as close to the reality as we can get and this is what I am striving for in my calculations. I think from what I related above, it is obvious that there are guesses and assumptions made in my calculations. No one is saying that Puertasaurus *had* to be a scaled up version of Malawisaurus, but it is the best model we have until the paper of Futalognkosaurus‘s measurements is published (which apparently has winglike transverse processes similar to Puertasaurus).

    Until you give me the width you restored for the D2 above (and even the C9, because scaling off the C9 your D2 does in fact appear to be just slightly over 168 cm) which may effect my estimates based on your drawing, all I can say is that the restoration above appears to depart significantly from even the most liberal model based off of known titanosaurs (both visually and mathematically) as regards torso width. This effects mass estimates which were the primary reason for this discussion. I would send you the Greg Paul skeletal of Opisthocoelicaudia, but I am not sure if that would be copyright infringement. Maybe if you can rent “The Complete Dinosaur” edited by James O. Farlow and M.K. Brett-Surman from a library in your area (or even buy it at a used-books shop like I did for cheap), you can find the skeletal on p. 281.

    Yes it is true I did do a quick sketch of Puertasaurus a while back and is still on deviantArt, but it was not done with the amount of detail in your above drawing so any estimate off of it is at best grossly inaccurate. I did do a lateral view skeletal of Puertasaurus (and Rapetosaurus) that is currently up on deviantArt, but it is not multi-view. I may go for a life reconstruction in the not-too-distant future, but no promises.

    I am currently working on a lateral-view skeletal of Malawisaurus also that should be up soon on deviantArt, I may try to do a multi-view in the future, but no promises (only two of the dorsal ribs are actually figured).

  44. Nima Says:

    I doubt sending a scan of the thing is copyright infringement, since the SV-POWsketeers have already posted some of Paul’s published skeletals here for educational purposes and obviously gained no money from doing so. As long as it serves the interest of science alone I’m pretty sure there’s no infringement – you’re not selling copies of the thing or passing it off as your own work. I have yet to come across “The Complete Dinosaur” in my area, so I don’t know where to get a glance at this thing without buying the whole book.

    I’ll check out what you have at DA, it seems you beat me to reworking Rapetosaurus! And yes, the mounted skeleton gives a ton of clues to how wrong Hallett’s skeletal is.

    As for my “restored” Puertasaurus D2, it’s ~1.9m wide. You’ll notice the wing-like processes are a bit longer and sharper than the published photos in Novas at. al.

  45. Peter Says:

    You didnt account for the skull when estimating a 31.7m length, which would bring the length close to 33m. Your estimate is still well below the 40m estimated length give by most sites. Although if a 31.7m individual weighed 110 tonnes then a 40m individual would weigh around the 190 tonne mark which is perhaps unrealistic.

  46. @Peter: I’m assuming this comment was directed towards me, since I gave the length estimate of 31.7 meters. You are correct that I left out the skull in the length, which means if my previous estimate of length was accurate, then it would be around 33 meters long.

    On the other hand, as I have pointed out above and in other places, Nima’s restoration is overly-speculative. He restored the width of the 2nd dorsal significantly wider than it is preserved as he claims there is evidence of significant erosion. However, the paper in which Puertasaurus is described does not suggest this, so Nima’s restoration above way over estimates the dimensions of Puertasaurus.

    Since Puertasaurus seems to be a Lognkosaurian, if we compare it to Futalognkosaurus, which had second dorsal with a functional length (that is, excluding the length of the cranial articular surface) of approximately 28.6 cm long, and the functional length of the second dorsal of Puertasaurus is about 30.6 cm (scaling of the figures in the original paper). This means that, scaling of this one dorsal mind you, Puertasaurus was only 30.6/28.6= 1.0699 times the length of Futalognkosaurus and therefore only 1.0699^3=1.2247 times as massive.

    A 2008 abstract reduced the estimated length of Futalognkosaurus to 26 meters (this is assuming a rather long tail,see here), which might imply that Puertasaurus was about 31.84 meters long.

    If Futalognkosaurus massed about 30 tonnes, then Puertasaurus might have massed only 36 tonnes. However, if we note that the 2nd dorsal was 168 cm in Puertasaurus and the anterior dorsals are described as being 100 cm wide in Futalognkosaurus this might imply that Puertasaurus was 1.68 times wider. So if it was 1.0699 times longer, 1.68 times wider and assuming it was about 1.26 times taller (based on own estimation of the height of the anterior dorsals in Futalognkosaurus at about 84 cm tall) this translates to 1.68*1.0699*1.26=2.26 times as massive as Futalognkosaurus which might suggest a mass as high as 68 tonnes. I should point out that going off of diapophysial width is probably misleading since there is no correlation between diapophysial width and total width of an animals torso. In fact, in Malawisaurus the anterior dorsals are progressively wider than their mid-to-posterior dorsals, in spite of the fact that the narrowest part of the body cavity is at the anterior portion and the widest part is in the mid-to-posterior portion. A better benchmark for comparison would be centrum width, however the widths of the centra in Futalognkosaurus have not been given. My guess is that the centrum width ration is probably similar to the height ration which would suggest a mass of 1.0699*1.0699*1.26=1.44 times that of Futalognkosaurus which would result in a mass estimate of about 43.2 tonnes for Puertasaurus.

    Taking the speculation further, a hypothetical, “world-record-sized” 40-meter long individual would then be (40/31.84)^3=1.98 times as massive, or about 135 tonnes if a 68-tonne “normal-sized” individual is assumed or about 85.5 tonnes assuming a 43.2 tonne “normal-sized” individual.

  47. Peter Adlam Says:

    Futaloognkosaurus was estimated at between 32-34m in length and considering the whole torso and neck were found (only the tail had to be “filled in”) a new 26m estimate is a bit shocking i mean how can they overestimate the tail by 6-8m, it seems slightly amateurish.

  48. Considering that almost every new “giant” sauropod in recent years has tended to be over-estimated in size (both in length and mass) it should not be surprising that this happened in the case of Futalognkosaurus.

    For instance, “Seismosaurus hallorum” was originally estimated to be 39-52 meters long by its original describer, and now is only thought to be between 28-32 meters long, which is a reduction of over 20 meters! So a reduction of 6-8 meters is comparatively small.

    I’m not sure it was the tail alone that caused the overestimate in Futalognkosaurus. Sadly, we have to *wait-for-the-paper* for a more complete description and estimation of its size to be sure. However, if the length estimate was based on the skeletal in the original paper describing Futalognkosaurus, and since the scale as indicated be the scale-bar was off in the skeletal restoration, this probably is what contributed to a overestimate of its size.

    I think sometimes people don’t realize just how hard it can be to get the scale-bars right and consistent between images, especially if you don’t have the technology available to make it easier. This is on top of the fact of how difficult it can be to create an accurate skeletal restoration of which a length estimate should be based. In the original paper, it appeared that the dorsal vertebral column had not yet been prepared and possibly the sacrum was not prepared either when the original paper came out. Due to this uncertainty, plus the difficulties of doing accurate skeletal reconstructions (even when all the bones are prepared and nicely photographed–trust me) and matching up corresponding scale-bars, it should not be too surprising that they over-estimated the length.

    Thus, I think it may be a bit overly harsh to describe it as “amateurish” especially since they aren’t the only scientists to have overestimated a dinosaur’s size. Remember that it is the nature of science to be self-correcting, and it is unlikely that any study on a particular topic is going to get everything right–in fact the probability is probably close to nil. It’s also interesting to note that the follow-up paper to the original description did not repeat that 32-34 m length estimate, and they have published an abstract that reduced the length to 26 m. So it seems that they caught their error and have corrected/are correcting it, which is good and what we want to happen in the scientific process.

  49. Peter Adlam Says:

    An estimate of 28m for Diplodocus halli (hallorum)? A bit ridiculous when Diplodocus Carnegi is 26m, only 0.93 smaller in linear dimensions. They are practically the same size,despite the reduced estimates many sites say it was still by far the largest diplodocus ever found (this goes against a 28m length, the most common revised estimates put the creature at between 33m-37m which is no way near the 52m length but big enough to be noticeable larger than D.carnegi. I can’t believe the corresponding bones of carnegi our 0.93 times the size of D.halli when even your site says it was Considerably larger than any previously discovered Diplodocus.

  50. Well I actually gave the range of 28-32 meters, given by some authors. I actually think 30-32 meters is the more accepted figure. To compare, the D9 in “Seismosaurus hallorum” (I put the name in quotes because it is now generally thought to be a specimen of Diplodocus longus; the species epithet was emended from ‘halli‘ to ‘hallorum‘ by George Olshevsky and other sauropod workers have followed suit)was about 1.2 meters tall. In D. carnegi (whose length is usually cited as around 25 meters) this same dorsal was 0.966 meters. This means D. carnegi was 0.966/1.2=0.805 times that in “S. hallorum“, or “S. hallorum” was about 1.2/0.966=1.24 times as large in linear dimensions as D. carngegi which is 1.24*25=31 meters–within the range I cited above.

    For comparison, Lucas et al. (2006) estimated the length at 33 meters and Lovelace et al. (2007) estimated the length at 30 meters based off of Diplodocus longus.

    I’m not sure where you get the up to 37 meter length estimates, as the most recent data all suggests a length of 30-32 meters as most probable which is still 5-7 meters shorter than the upper end of the estimate you cite. This is still close to the 6-8 meters that the original Futalognkosaurus length estimate was off by, so it shows that length overestimates of 5-8 meters are not that uncommon.


    Lucas, S.G., Spielman, J.A., Rinehart, L.A., Heckert, A.B., Herne, M.C., Hunt, A.P., Foster, J.R., and Sullivan, R.M. (2006). “Taxonomic status of Seismosaurus hallorum, a Late Jurassic sauropod dinosaur from New Mexico”. In Foster, J.R., and Lucas, S.G.. Paleontology and Geology of the Upper Morrison Formation. New Mexico Museum of Natural History and Science (bulletin 36). pp. 149–161. ISSN 1524-4156.

    Lovelace, D. M., Hartman, S. A. & Wahl, W. R. 2007. Morphology of a specimen of Supersaurus (Dinosauria, Sauropoda) from the Morrison Formation of Wyoming, and a re-evaluation of diplodocid phylogeny. Arquivos do Museu Nacional, Rio de Janeiro 65, 527-544

  51. Peter Adlam Says:

    Still 31m is still longer than was thought for diplodocus,but it seems to be shorter than Supersaurus,how do the corresponding bones between diplodocus hallorum ( i thought Seismosaurus had been completely ditched) and Supersaurus compare? I suspect Supersaurus was bigger given that the neck alone was at least 13.5m.

  52. Peter Adlam Says:

    To go back to the Peurtasaurus drawing the neck looks considerably oversized for a tintanosaurid and if that is a 6ft man than the creature looks in the region of 45-50m unless that man is Verne Troyer because like you said its a 33m sauropod.

  53. The posterior most or second-to-last dorsal in Supersaurus is 133 cm tall, and the second-to-last dorsal in “Seismosaurus hallorum” is about 120 cm, so Supersaurus probably was larger. I think the most recent estimate of its length is 33-34 meters long. You’re right that Seismosaurus has been ditched as a genus name (hence why I refer to it in quotes most of the time). It’s generally now thought to be an individual of Diplodocus longus.

    *My* estimate for Puertasaurus based on his drawing and the reported dimensions of the bones as described is 33 meters long, *Nima’s* reconstruction (above) is quite a bit larger than my estimate based on the scale bar and because he restored the vertebrae larger than described. For instance, using GIMP’s measuring tool, the 4 meter scale-bar is 139 pixels, and the distance between the pre- and post-zygapophyses of the 9th cervical is 52.5 pixels. This means the cervical is about 151 cm between the pre- and post-zygapophyses. The actual length as measured and given in the description of Puertasaurus is actually 118 cm. So Nima increased the length by about 28% than it should be. So reconstructed above, the total length of Puertasaurus is about 43.8 meters long from tail tip to the tip of the skull, measured along its vertebral axis.

    So if my total mass estimate for Puertasaurus of 136.6 tonnes based strictly on Nima’s drawing is correct, then if we reduced it to 33 meters long, this is about 75% of the length Nima constructed, which implies a mass of about 42% of the original mass estimate, which is about 58 tonnes as he reconstructed it. The guy appears to be 6 ft and 1/2 inch tall based off of Nima’s scale-bar. Nima says the guy is Eugene Sandow, who was actually 5 ft 9 and 1/4 inches tall. So the scale bar is about 4.5% too large from what it is supposed to be, meaning, as reconstructed, the Puertasaurus is about 41.88 meters long. Even taking that into account, the cervical vertebrae is still ~22.5% too long.

  54. Peter Adlam Says:

    I am quite surprised that it turns out that “Seismosaurus” was a Diplodocus longus, so what is the difference between a Diplodocus carnegi and longus? what is the average size of the previously discovered D.Carnegi’s and D.Longus’s? before “Seismosaurus” i thought D.carnegi was perhaps slightly larger but obviously not because at 31m it dwarfs D.carnegi. Is this a freak compared to other specimens of D.longus? Maybe a very old large male (i don’t know which gender was largest i’m just guessing male)? I know i’ve asked a fair few questions but your knowledge is well appreciated, you certainly know your stuff.

  55. As Mike said in a comment on the other post on Amphicoelias fragillimus, the species of Diplodocus are not well known. This problematic aspect was commented on in the Lucas et al. (2006) citation I gave in one of my earlier comments above (that paper is freely available online if you want to check it out). I’m not intimately familiar with the species of Diplodocus, but if you want a visual, see Scott Hartmann’s skeletal reconstructions here which compare D. carnegii, D. longus and “S. hallorum“. At a quick glance, the differences between the two species of Diplodocus appear to be so small that they could well be just intraspecific variation. On the other hand, one’s definition of a species is pretty much arbitrary. In fact, some modern bird species are nearly indistinguishable skeletally and are hardly distinguishable by their outward appearances. Many species of warbler and sparrow, for instance, are very difficult to recognize on outward appearances and are best distinguished by voice and behavior. Obviously, these type of differences are not able to fossilize. Even worse is our knowledge on the gender of most dinosaur species, especially sauropods, so even gender specific variation is not quantifiable at this time.

    As Lucas et al. (2006) (in part) noted, “There are four named species of Diplodocus…These species are routinely listed as valid…although convincing arguments for their validity are difficult to construct. Thus, D. longus and D. carnegii are not demonstrably distinct. D. lacustris is based on fragmentary premaxillary, maxillary and dental material, elements that do not demonstrably vary among Diplodocus species, so it is best considered a nomen dubium (nomen vanum)…D. hayi, however, can be distinguished from D. longus and D. carnegii by characteristics of the skull and caudal vertebrae according to McIntosh…Obviously, a rigorous revision of the species-level taxonomy of Diplodocus is needed, though such a revision faces the obstacle of needing to compare species that are not all known from (or diagnosed on) overlapping anatomical parts. In the absence of such a revision, we consider the holotype of Seismosaurus hallorum to represent a fifth species of Diplodocus, Diplodocus hallorum, distinguished by the minor morphological differences…we have little confidence that these are even valid species-level differences, and suggest it is likely that D. hallorum is a junior subjective synonym of D. longus. However, this suggestion can only be verified by a complete study of species-level variation in Diplodocus that is beyond the scope of this paper. This is a significant problem. Without a better understanding of the real morphological variation present within a genus of sauropod dinosaurs, at least some of the characters and character-states coded in the numerous phylogenetic hypotheses of sauropod relationships now available must be taken with the proverbial grain of salt.”

    As far as I am aware, such a detailed study of the species of Diplodocus has not yet been published. The seismosaur specimen does indeed appear to be significantly larger than the specimen for which D. longus is described. The latter species is known from two skulls and partial skeletons, apparently, so we don’t have a good sample size of which to see how big “normal-sized” individual would be.

  56. Mike Taylor Says:

    There is little to add to what Zach’s written here. It’s an open secret that Diplodocus carnegii was never convincingly diagnosed, as pointed out as long ago as 1932 by Gilmore (“On a newly mounted skeleton of Diplodocus in the United States National Museum”, Proc. U.S. Nati. Mus. 81:1-21). I don’t know the history, but it’s a decent bet that Hatcher felt obliged to name a new species after Carnegie in gratitude for his funding of the work, whether or not the new specimen merited it.

    The real question is, in more than a hundred years, why has no-one revisited the problem of Diplodocus species? It seems like a tantalisingly obvious project, and I can hardly understand why it’s been overlooked for all this time. Someone ought to sort it out.

  57. Odontodactylus Says:

    Multiplying the total body volume calculated by Zach (the slimmed torso version) by a density of 0.7 tonnes per cubic metre – suggested elsewhere on the site for other sauropods – would give a mass of about 96 tonnes. I prefer this estimate to 110 tonnes as I’ve always assumed huge sauropods would be especially pneumatic, but I have nothing to back this up. Also it just sounds more reasonable to me, but that’s personal incredulity for you.

  58. Bryan Riolo Says:

    Much ado about incomplete material. While Zach and Nima have good arguments to make–I side mostly with Nima–there is not enough material on these dinosaurs, as yet, to put forth high probability conclusions, and I say this even if the Futalognkosaurus, Argentinosaurus, and Puertasaurus material was complete. It isn’t. Speculation should be just that–speculation, not theories put out as fiats from on high. The chances that our intrepid scientists have found the largest individuals in each species, genuses, whatever!-are very remote.

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