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Big honkin' theropod of the southern hemisphere

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By ScottHartman   |   
© 2012 - 2020 ScottHartman
I think Giganotosaurus epitomizes what people are expecting in their imaginations when we talk about giant predatory dinosaurs. It's stocky and overbuilt, but not in the "weight-lifting ballerina" style of T. rex. Instead, it's basically just a really big carnosaur with a big head.

Keen observers (or just those with too much time on their hands) will notice that the head is not reconstructed at all like the earlier elongate one that was created for the cast that is mounted at several museums - that's because it was wrong. For this version I drew on skull elements from Mapusaurus and Tyrannotitan to better flesh out the missing bits of its noggin.

Edit 2: As I was working on a size comparison between large theropods, I realized that there was a minor scaling error in the hind legs relative to the rest of the body (or more accurately, I realized I'd used Mapusaurus's proportions instead). This has now been fixed, and Giganotosaurus is somewhat closer to the ground (although still with a relatively tall femur).
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anonymous's avatar
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Megalosaurid's avatar
:star::star::star::star::star: Overall
:star::star::star::star::star: Vision
:star::star::star::star::star: Originality
:star::star::star::star::star: Technique
:star::star::star::star::star: Impact

By far the best reconstruction of Giganotosaurus I have ever seen, its bizarre, its beautyful, its original, and most important of all, more accurate.
I does not only correct the errors in the mount but just includes many new things like longer legs and more accurate posture, deffinitely worth to watch and original.
Best Giganotosaurus drawing ever!!!
The ribs are restored correctly and the skull is shorter, also correcting things like the curvature of the spine and improving a slighly more slender neck than the mounts or at least that is the impression.
Deffinitely a Masterpiece of Paleontology and reconstructions.
ScottHartman's avatar
ScottHartmanProfessional Digital Artist
Caprisaurus's avatar
:star::star::star::star::star-half: Overall
:star::star::star::star::star-half: Vision
:star::star::star::star::star: Originality
:star::star::star::star::star: Technique
:star::star::star::star::star: Impact

1 critique already, but 2 critiques are better than 1. As for the deviation, i've only got 1 word:awesome. The head of this magnificent beast is original and certified. This is perfectly positioned and totally fantastic. The bones are correctly thickened and adds the picturesque view to it. This would be a great feature drawing on any place! Instead of putting the feet's pad flattened to the the ground, you lifted to put in more impact. The teeth grows to certain length and width. Good score on the rates and this is how it should be for correct motion. I'm happy this exists.
piche2's avatar
piche2Hobbyist Traditional Artist
:star::star::star::star::star-half: Overall
:star::star::star::star::star-empty: Vision
:star::star::star::star::star-empty: Originality
:star::star::star::star::star: Technique
:star::star::star::star::star: Impact

es el mejor esqueleto de giganotosaurus que e visto en esta pagina web.Me gustaría ver mas ver mas de ti ya eres un excelente artista y que seria mejor que hicieras mas ejemplares de dinosaurios ya que este dinosaurios es el emblema de mi país.Sin duda eres uno de los mejores paleoartistas que visto y me gusta muchos tu ilustraciones, me gustaría que incluyeras el tamaño que tiene haber tenido este sorprendente depredador que existido y que existirá jamas sobre la tierra.
Lo digo y lo vuelvo a decir eres uno de los mejores paleoartistas que visto aquí y suerte
ScottHartman's avatar
ScottHartmanProfessional Digital Artist
Muchas gracias!
piche2's avatar
piche2Hobbyist Traditional Artist
No gracias a vos por dibujar tan bien eres muy bueno ento
tireironsforthegays's avatar
tireironsforthegaysHobbyist Digital Artist
the great devourer
TeamDinosauria21's avatar
TeamDinosauria21Student Digital Artist
Sadly Giga is underrated due to the stronger Carcharodontosaurus
mark0731's avatar
Carcharodontosaurus was hardly stronger than Giganotosaurus, as it was a bit smaller or comparable in size to the holotype of Giganotosaurus, which means it was quite a bit or a bit smaller than the paratype of Giganotosaurus.
christina1969's avatar
Do you know anything about that mixture between Gregory Scott Paul's earlier skeletal and yours:… ?
TimeToGoHero's avatar
TimeToGoHeroHobbyist General Artist
What I don't get is if the skeletal is based on components from both Tyrannotitan and Mapusaurus; then why is the skull so much shorter? As far as phylogenetics have shown most the Carcharodontosaurids had elongated skulls; so for whatever reason for the Giga to give up the elongated skull when the prey items were mostly defensless (unless targeting the Sauropods and Ankylosaurs) then I can not see why it would evolve in such a manner.
ScottHartman's avatar
ScottHartmanProfessional Digital Artist
No, the elongated skulls are basically a myth. There's no such thing as a complete Carcharodontosaurus skull, and the one that NGS helped popularized is notoriously elongated beyond what the bones support (the nickname for that reconstruction is Pinnochio). But that was published while a team of sculptors was creating the Giganotosaurus mount, so they copied it assuming that it was a good guide. No carcharodontosaur ever had a skull like that, and Giganotosaurus was no exception.

FWIW, if you dig up the original Giganotosaurus publication (which came out before the pinocchio carcharodontosaur paper) you will see that while the skeletal is pretty horrible, they actually reconstructed the skull surprisingly close to what I've done here.
TimeToGoHero's avatar
TimeToGoHeroHobbyist General Artist
Hmm, I don't know just reconstructing the skull myself last night. The skull of Giganotosaurus seems to be a slight bit elongate, but not to how narrow popular restorations have it. And I never said Carcharodontosaurus had a complete skull; needful to say that Giganotosaurus doesn't have neither, and yes at least one does. The other Giganotosaurians, Mapusaurus and Tyrannotitan seem to have similarily elongate skulls to that of the popularized Giganotosaurus restoration (however they're not nearly as long). Based on the mandibular and the partial remains of the naris both theropods respecitvely, Coria & Currie have at estimated both partials in the 10 cm range (which of course is obvious to do) adding the math, if multiplying in cm to converting into inches; the upper jaws of both theropods are roughly estimated to be around 5ft (1.53-1.55 m) while in Tyrannosaurus skulls (specifically our lovable, Sue) it is 4.8ft (1.47 m) in length; in skeletal metrics, Franoys seems to clarify this using possibly the same principals.

That being said putting all the pieces together to apply in the Giga's skull proportions it likely had a 5ft skull as well; tis the case with phylogenetics when all the members share similarites. I also happened to accquire the original publication (at least I think) and also looked upon the original holotype skull taken by a photographer on Wikipedia. From what I've seen in Currie and Coria's illustration (See Fig. 6, Page 121) the skull isn't as long as like the ones from the museum, but is more pronounced and looks very much similar to illustrations I've been well acquinted with in the Don Lessem book from my childhood, and the Dino Crisis video game (although that Giga was exaggerated); from what I foretold earlier up above (elongated, but not like this which to me seems like good Greg has some help presenting that). As I look into your restoration you've compressed the snout of the poor Allosaur's skull in, making it look almost a fixate cross between T.Rex and Allosaurus (e.g. Indominus Rex) or even close to what Land Before Time's Giganotosaurus design. Moreover based on these variables, where you assume that the elongate skulls of Carcharodontosaurids are a myth, this supposed "myth" wasn't too far off from the truth; granted that if Giganotosaurians grew their skulls 10 (2x)cm longer or in referring to the other specimens like MUCPv-95 and MCF-PVPH-108.145 then it could easily can be discussed over that Giga(s) had longer skulls as their size progressed in age (much like how humans' noses grow bigger as we grow older) and that the only member of the group with the shortest skull is Carcharodontosaurus. It is interesting concept considering Giganotosaurus might have had ontogeny differences; which I like to thank from not only you and Greg's reconstructions, but Coria & the Currie's as well.

Here is the restoration of my Giganotosaurus skull from I've gathered; restoration is made for The Land Before Time, but still it's carefully studied on. I'll be using these for a species identification debate involving the Theropods (Sharpteeth) on The Land Before Time wiki.
LBT Giga resto20170814 141603 by TimeToGoHero   Sharpteeth '17 part 1 20170814 141545 by TimeToGoHero  

On top from this hope you can see from where I've presented this, and I don't mean to sound show boaty and with all do respects Mr. Hartman hope we can come to an agreement.
ScottHartman's avatar
ScottHartmanProfessional Digital Artist

I apologize if I'm overly curt in my reply, but I'm smack in the middle of a massive skeletal project on a tight timeline, preparing to lecture a course, and have SVP in another week and a half where I'm presenting research. The problem here is you keep linking to drawings, rather than the real bones. I've played with Carcharodontosaur skull bones and they don't go together that way either. Greg Paul doesn't draw Carcharodontosaurus like that anymore either; in his most recent edition of his Field Guide he's fixed it to look more like how I restore Giganotosaurus (not saying he copied me, it was independently). 

Likewise, the Calvo and Coria paper you cite didn't add any new bones to justify their pinnochio Giganotosaurus skull drawing, they just copied Sereno's infamous pinnochio skull sculpture and scaled it up to their dentary. If one were cynical, one could note that this happened in a paper where they were evaluating how big Giganotosaurus was, and by adopting the pinnochio-style skull it made the skull much longer than their earlier (and more accurate) reconstruction.

What has been published of Mapusaurus skulls is not complete enough to show an elongate snout either, it was just assumed based the previous reconstructions. It's bad skull reconstructions all the way down, none of them are known from complete skulls that justify either the long snout or the gigantic post-orbital region, so everyone just kept copying a (very!) wrong reconstruction. The rest of the books/images you linked to are life reconstructions and/or video games - I'm not against those, but they don't count as data.

Once we find a complete Giganotosaurus skull I'm sure I will have to make some updates. But nothing in the current collection of fossils, nor what is known of Mapusaurus and Carcharodontosaurus supports the stretch-job version, it's all the byproduct of people copying the original, incorrect Carcharodontosaurus skull sculpture. The funny thing is that paleontologists (and some paleoartists, though I wasn't clever enough to be one of them at the time) started criticizing that Sereno/NGS skull sculpture right away, but once it was copied by the people sculpting the skull for the Giganotosaurus mount and used in the Currie & Coria size paper it just got accepted (even though there weren't any new Giganoto bones to support it).
arvalis's avatar
arvalisProfessional Digital Artist
You're a real trooper for dealing with this guy. He has been harassing everyone with his terrible ideas and then gets very upset when he is corrected.
ScottHartman's avatar
ScottHartmanProfessional Digital Artist
Actually I was concerned I was being too big of a dick, but I'm really short on time these days.
paleosir's avatar
paleosirHobbyist General Artist
Really you shouldn't even bother wasting your time on him. I've tried to reason with him, to no avail.
ScottHartman's avatar
ScottHartmanProfessional Digital Artist
I try to be reasonable with everyone. It's a character flaw ;)
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TimeToGoHero's avatar
TimeToGoHeroHobbyist General Artist
Yeah well sometimes your reconstructions aren't exactly always right. Your work is a bunch of drawings too; you don't think these other paleoartist take the time to reconstruct these incomplete skulls as best as possible? Most importantly the majority of the illustrations I've presented are well good variables to what I'm trying to conclude.

"Using both Tyrannotitan and Mapusaurus as reference", "putting the pieces together/monomorphology" did that not a ring bell Mr. Hartman? Despite that your rebuttal on the claim as I've stated from earlier today the estimates are quite clear just by estimating the naris bone by the 10cm scale bar; rough estimate at least adds up 60 which I then converted as inches, 60 inches in reference to a foot measure is roughly 5 feet in length. Though the skulls of most all the Carcharos are incomplete it is possibly to narrow them down. It's one reason why phylogenetics are even a thing to begin with you know, it is how we've been able to reconstruct some dinosaurs from their closest relatives (of course you already knew that).

Third I don't recall Coria and Currie ever drawing a Pinocchio Giga, because as the paper I've showed you seems to be the first one recording the Giganotosaurus holotype which they show is having a moderate elongate skull (much like reconstruction of my Giganotosaurus I had showed you). Honestly I find it distasteful that Paleontology can grow to be that corrupt just from a single copy by some lazy bones. But you know at least it isn't a total lost when the holotype is still…
ScottHartman's avatar
ScottHartmanProfessional Digital Artist
First off, I've hidden two comments from two different posters that had no content except insults. Please (everyone) do that somewhere else on DA, but not on my pages.

Second, of course my skeletals aren't always right. But I make it a point to use the original data as often as possible to avoid copying mistakes made by others, and I update them whenever new data comes to light (though sometimes there's a delay when my schedule is too busy). 

Third, I've looked at all the skulls you are bringing up - they didn't just get published yesterday. Tyrannotitan is known from fewer bones than Giganotosaurus. The jugal of Tyrannotitan can be used to fill in for the missing element in Giganotosaurus, but that doesn't do anything to support the old pinocchio skulls, and nothing else in the scrappy Tyrannotitan skull remains supports it either. Mapusaurus actually has a decent back half of the skull, and it does NOT look like the old drawings with a super-sized infratemporal fenestra. It looks like a typical acrocanthosaur-allosaur style back of the skull. The snout of Mapusaurus is not complete, but the maxilla and partial nasals are perfectly compatible with a normal, acrocanthosaur-style snout. Since Carcharodontosaurus skulls are not complete enough to make this case, there is literally zero animals that show this, meaning there's no reason to draw it that way.

Fourth, and this is where it gets a bit wearying to have to continue, but the paper you linked to is:

Calvo, J. O. & Coria, R. A. (1998). New specimen of Giganotosaurus carolinii (Coria & Salgado, 1995), supports it as the largest theropod ever found. Gaia15, 117-122.

You say you "
the paper I've showed you seems to be the first one recording the Giganotosaurus holotype", but that's simply not true. The first time the skull bones were figured was in the original description:

Coria, R. A., & Salgado, L. (1995). A new giant carnivorous dinosaur from the Cretaceous of Patagonia. Nature377(6546), 224.

I don't have time to crop out the image from the pdf and upload it, but it doesn't look anything like the image in the 1998 paper you linked to. And they didn't find any new bones that justified this change (they were just describing the a new chunk of dentary). Literally the only thing that had changed was that Sereno and NGS published the inaccurate skull reconstruction of Carcharodontosaurus that was based on very scrappy remains. Calvo and Coria copied it for their paper, and then artists started copying that. Like I said, it's bad reconstructions all the way down. Moreover, the fact that you don't know the publishing history, but you are making giant, sweeping statements about who is right and who is wrong is a bit tiresome, when I could be using what little time I have to answer questions for people that aren't making up arguments without knowing all the data.

It's fine that you haven't read every paper out there, I don't expect most people on DA to read the technical literature. And having read them is obviously not a guarantee that someone is right. But when you are arguing about how a certain style of reconstruction came to be (and whether it's based on anything other than people copying other people's bad illustrations) you really need to have access to the real fossils (or at least images of them) and you need to have a good knowledge of the order all those illustrations occurred in, and it helps if you were there at the time and talking to the people who did them.

You are right that the holotype is around, and if you look closely at the image you linked you can actually see the problems right there. Moving from left to right, there is no reason for the giant gulf of white plaster in the squamosal. The postorbital is actually really short on the posterior end - all that plaster was invented to try and match the inaccurate Carcharodontosaurus skull. Likewise, look at the giant swaths of plaster where they invented the jugal and quadratojugal. Tyrannotitan jugal does not look like that, and the back half of the Mapusaurus skull bones look nothing like that. It's just sculpted to look like the erroneous Carcharodontosaurus reconstruction. Moving to the snout, notice again the white between lacrimal and the nasals - there's no reason for that. There's also no reason for there to be a gap between the broken ascending process of the maxilla and the floating chunk of nasals. It's all arranged to meet a pre-existing concept set up by the incorrect Carcharodontosaurus skull. Nothing in these fossils supports the spacing here, or the way the plaster had been sculpted to fill in that spacing.

And it's not the way it was originally reconstructed:…
DEDEYE64's avatar
I hear you. You're not the first professional to be pestered by this guy either.…
Franoys's avatar
FranoysStudent Digital Artist
Hi Scott. Looking at Giganotosaurus more in depth ( it is kind of hard due to how little it has been written or described in depth about it) and following the most recent literature there are some things that I noticed differ from your skeletal.

As a first note it seems that the diminutive pectoral girdle portrayed here and in the mounts of the animal is artificial, a product of bad preservation ( the scapulocoracoid is very damaged as can be seen in the images in which it has been ilustrated or photographed). So the low acromion and unexpanded distal end are artificial. The coracoid is broken in two pieces and has a preservational crack, but part of it is attached and fused with the scapula. In your skeletal you seem to portray this crack, perhaps as if it was the separation between the bones.

 This was recently adressed in the Tyrannotitan osteology study by Canale and Novas (2014) in which they say this:

"A detailed comparison between the scapulocoracoid of Tyrannotitan and Giganotosaurus reveals that in the coracoid of the holotype of Giganotosaurus, the dorsal and anterior borders are damaged and only the sector over the glenoid cavity is preserved. The scapula of the type specimen of Giganotosaurus also has the acromial process broken, which was previously interpreted as a low acromial process.

The character ‘externally open coracoid foramen’ is also produced by a misinterpretation of the type material of this taxon. The coracoid foramen is present, but in a fragment of the left coracoid that is fused and preserved attached to the scapula .This fragment of the coracoid was interpreted as part of the scapula by previous authors, but the suture scar between the coracoid and scapula is visible in the type material.  The position of the coracoid foramen is almost the same as in Tyrannotitan, located centrally on the lateral surface of coracoid.  The interpretation of the fragment of the coracoid as part of the scapula led previous authors to postulate the autapomorphic character of Giganotosaurus ‘proximal end of the scapula forwardly projected over the coracoid.

In sum, the pectoral girdle of the type material of Giganotosaurus (Coria and Salgado 1995) is incompletely preserved and led to a misinterpretation of its anatomy. We interpret the scapula and coracoid of Giganotosaurus as similar to that of Tyrannotitan (Novas et al. 2005) and Mapusaurus roseae (Coria and Currie 2006), having a wide coracoid, well-developed coracoid foramen, scapula and coracoid fused, and the contact between these two elements is oriented perpendicular to the long axis of scapula. Although the acromion is only partially preserved in the holotype of Giganotosaurus, the available remains suggest its morphology did not differ from that of Tyrannotitan. "

Some bones are clearly based and drawn after the figures in Coria and Currie 2006, but I'm not sure if this is a good idea at least for the ischium; in the very same paper it is said that the ischium in Mapusaurus is curvy, an autapomorphy in respect to Giganotosaurus, in which the ischium is straight. The ischium in your skeletal is curvy and directly based on the ilustration, resulting on giving Giganotosaurus a Mapusaurus' defining character.

"The dorsoposterior margin of the shaft in Mapusaurus is somewhat convex in lateral view, but because the distal end expands ventrally, it gives the shaft the appearance of a relatively strong ventral curvature. In contrast, the ischium of Giganotosaurus is straight in lateral aspect."

Now onto the size of some bones:

The fibula in your skeletal (110+ cm) is a lot bigger than the measurement that we have of it (83.5 cm, taken from Coria and Currie 2006)

"Four complete Mapusaurus' fibulae (MCF-PVPH-108.132, -108.202, -108.196, -108.189) and several partials (-108.51, -108.220) have been recovered and range in length from 640 to 860 mm. Although the largest one (MCF-PVPH-108.202) is 2.5 cm longer than the holotype fibula of Giganotosaurus carolinii, it is more gracile "

The femur is also bigger than the measurement that Carrano made for it's 2012 Phylogeny of Tetanurae papers; apparently he got it as 1.365 meters. This is the quote from the papers:

"We consider Giganotosaurus to have had a skull almost exactly comparable in length to that of Tyrannosaurus. Likewise, our measurements of femur length in the holotype (136.5 cm, left)record a smallersizethan originally reported (143 cm; Coria & Salgado 1995) and therefore an animal of lower overall body mass

It is true that in the original description of Giganotosaurus in 1995 the femur was claimed to be 1.43 m, however FMNH PR 2081 femur was also claimed to be 1.38 m instead of 1.31 or 1.32 m, both Giganotosaurus and Tyrannosaurus femora are oversized by about 6-7 cm and by a similar percentage which leads me to believe that the femora were measured using a different metric from the traditional one, (Perhaps accounting for some kind of curvature?), while in your skeletal the femur is 1.43 m measuring from the greater trochanter to the condyle.

The tibiotarsus you portrayed at about 120 cm long. In the papers about Giganotosaurus braincase (Coria 2003) the tibia was reported as 112 cm, however I suspect that this is the measurement of the whole tibiotarsus, not just the tibia, otherwise it is extremely incoherent with the fibula measurement reported. In Gregory S.Paul's Giganotosaurus' skeletal, the whole tibiotarsus is 112 cm. I tried giving a Giganotosaurus mount photograph the measurements in the papers (femur and fibula) and the whole tibiotarsus was indeed about that size. This is the one I'm least sure about however.

I hope I didn't came out as too nitpicky.

PD: Where did you get the measurements of the axial skeleton? Were they provided by the ROM?
ScottHartman's avatar
ScottHartmanProfessional Digital Artist
Hi Franoys,

Yes, I need to make a couple of updates to the Giga skeletal. That said, I would not use the mount to scale from, as parts of it were sculpted prior to it being fully prepped. Also FWIW, Sue's femur has been reported at different sizes than just the ones you list - this is apparently a common error, and I'm sure you are correct that it's related at least in part to the shape of the bone (and some people not measuring in a single plane).

At some point I need to fix it and go Mapusaurus and Tyrannotitan, but I'm not sure if that'll be done this year or not, as I have just about every free moment for skeletals spoke for until 2018 it looks like :(
Franoys's avatar
FranoysStudent Digital Artist
Dear Scott, I hope I'm not being annoying with this (neither too nitpicky) but I dug up a bit more on this regard because of my own work and I figured I could share it with you since you plan to update your Giganotosaurus skeletal and also perhaps work on Mapusaurus in the future:

As a first point, it does seem that the largest fibula in Mapusaurus being 86 cm is false, as there is a fibula (MCF PVPH-108.202) ilustrated in Coria and Currie 2006 (Figure 30) that exceeds this measurements, giving us a more normally sized and proportional fibula-tibiotaursus discrepancy, as it seems to be about 100 cm going by the figure. Furthermore, the tibia ilustrated there ( MCF-PVPH-108.68) matches perfectly with the size that the tibia is expected to be compared to the fibula as it is 104.7 cm m long when measuring it with the scalebar provided in figure 29. In Acrocanthosaurus holotype, the tibia is about 85 cm is and the fibula is  80.1 cm , therefore the ratios (1.047 and 1.061) match rather convincingly, and so do those of other allosauroids. Based on that, and if Giganotosaurus fibula would be only 2.5cm shorter the the largest Mapusaurus fibula; we would have a cm fibula, which is indeed very coherent with a reported size of 112 cm for the whole tibiotarsus.

This is corroborated by the Dataset (S1) of Benson et al 2014 (in which they included forelimb and hindlimb measurements for hundreds of dinsaurian taxa, you can access the paper here :… and the supplementary materials here:….) According to it, the femur of Giganotosaurus would be 132 cm long and the tibia 99 cm. The tibial measurement of 99 cm is coherent with a 112 cm tibiotarsus. 

About Giganotosaurus femur, if we couple this measurement from Benson 2014 (132 cm) with Carrano's 2012 measurement (136.5 cm) and the medial view of it included in this paper with a scalebar (figure 15); (which yields barely over 130 cm too, I got 131) I think it is safe to say it is unlikely it exceeded 140 cm.

About the metatarsals; going by Acrocanthosaurus measurements, NCSM 14345 and OMNH 10147 have both tibiae expected to be about 96 cm long based on Stoval&Langston 1950 and Carpenter & Currie 2000, and femora that could have been well over 120 cm when complete; and the metatarsal lll of OMNH 10147 is 44.5 cm and that of NCSM 14345 is estimated to be 43.9 cm when complete, therefore I doubt Giganotosaurus metatarsals would have been longer than 50 cm; only Tyrannosaurids have such long metatarsals between predatory dinosaurs (as in longer than 60 cm). Carpenter and Currie 2000 dedicates one paragaph to discuss how even juvenile individuals of Tyrannosaurid have longer metatarsals than gigantic Allosauroids like Acrocanthosaurus.

PD: I'm sorry for what you had to go through just under this comment.


Calvo J.O (1999) Dinosaurs and other vertebrates of the Lake Ezequiel Ramos Mexía Area, Neuquén-Patagonia, Argentina. Proceedings of the second Gondwanan dinosaur symposium. pp 13-45. Natural science museum monographs, No.15, Tokio 1999.

Coria RA, Currie PJ. 2002. The braincase of Giganotosaurus carolinii (Dinosauria: Theropoda) from the Upper Cretaceous of Argentina. J Vert Paleontol. 22(4):802 – 811.

Coria RA, Currie PJ. 2006. A new carcharodontosaurid (Dinosauria, Theropoda) from the Upper Cretaceous of Argentina. Geodiversitas. 28(1):71 – 118. 

Carrano MT, Benson RBJ, Sampson SD. 2012. The phylogeny of Tetanurae. J Syst Palaeontol. 10(2):211 – 300.

Benson RBJ, Campione NE, Carrano MT, Mannion PD, Sullivan C, Upchurch P, et al. (2014) Rates of Dinosaur Body Mass Evolution Indicate 170 Million Years of Sustained Ecological Innovation on the Avian Stem Lineage. PLoS Biol12(5): e1001853.…


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