Guest Post: Shark toothed theropods in Asia – introducing Shaochilong

ResearchBlogging.orgSteve BI may be headding off to sunny Inner Mongolia, but have left the keys to the Musings behind in the hands of theropod specialist and memeber of the worldwide Bristol mafia, Steve Brusatte to talk to you about a new paper (and a new taxon) we have described with our colleagues (though I honestly and non-self depreactingly didn’t do much of it apart from coming up with the name). Take it away Steve:

A funny thing happened while I was in Beijing last January. I arranged a two-week visit to examine a bunch of basal coelurosaurs for my PhD thesis, and spent most of my time looking at Dilong and Guanlong. And then my gracious hosts, Dave Hone and Corwin Sullivan, both picked up a nasty flu bug [ed: Corwin got it and gave it to me] that knocked them out of service for an entire week. Suddenly I had the entire office to myself, and freed from Dave’s entertaining interruptions about pterosaurs and football and other passion-inducing topics, I began to work really fast [ed: what’s wrong with pterosaurs eh?]. So fast that I had a couple of free days left over at the end of my second week. Most reasonable people would use those days to act like a tourist and see the sights in Beijing, but I figured I may as well look at other specimens. And that’s how I came across a bizarre theropod called “Chilantaisaurusmaortuensis.


Shaochilong maxilla (top), braincase (middle) and skull roof (bottom) from Brusatte et al., 2009.

This creature, known from a single specimen that includes some very nice skull bones and vertebrae, was originally described by Hu in 1964. Today, over 45 years after it was discovered, “C.maortuensis remains one of the only large-bodied theropods known from a large gap in the Asian predatory dinosaur fossil record. We know a decent amount about Asian large theropods from the Middle-Late Jurassic, as well as the tyranosaurid-dominated faunas of the Late Cretaceous (Campanian-Maastrichtian). However, the intervening 60 million years is largely a dark period in Asian theropod history. For this reason, “C.maortuensis remains a critically important animal.

By now, many intrepid readers have probably noticed that I have placed “Chilantaisaurus” in quotation marks. That is because Hu referred this specimen—clearly a new and distinct species—to the genus Chilantaisaurus, which was based on another, much larger theropod specimen from the same mid Cretaceous rock unit in Inner Mongolia where “C.maortuensis was found. However, there are no overlapping bones between the two specimens, so we really can’t tell if the two species belong to the same genus. This problem has long been noted by many authors, beginning with my colleague Dan Chure when he visited IVPP and examined lots of theropod specimens in the 1990s. Dan had long planned to redescribe “C. maortuensis and name a new genus, but he was sidetracked by other things.

While in Beijing I figured I should try to see the specimen, as some authors had suggested that it was a tyrannosauroid or a basal coelurosaur of some kind. Either way, it would provide important data for my thesis. But when I opened the drawer and saw the maxilla and braincase staring back at me, I quickly realized something was wrong. “C.maortuensis is not a coelurosaur, but possesses a number of unique features seen in Carchrodontosaurus, Giganotosaurus, Mapusaurus, and Acrocanthosaurus—colossal theropods closely related to Allosaurus that are grouped within their own subclade, the Carcharodontosauridae (the shark-toothed reptiles, named for obvious reasons).

Carcharodontosaurids are a relative newcomer to the pantheon of large theropods. The first specimens of this group were found by Ernst Stromer in the early 1900s in Egypt, but most were destroyed during World War II (from the same beds as the legendary Spinosaurus). Only in the 1990s were more complete specimens discovered in Early Cretaceous rocks in Africa and South America. For a while people thought that carcharodontosaurids were restricted to the southern continents, but then it became clear that the humongous North American Acrocanthosaurus was also a member of the group. And over the past decade people have been finding more carcharodontosaurids from across the globe, including Neovenator from the Isle of Wight in England. However, one place that seemed to lack carcharodontosaurids was Asia. Some workers interpreted this absence as significant—perhaps it meant that Asia separated from the rest of Pangaea before carcharodontosaurids evolved.

Now we know that carcharodontosaurids lived in Asia as well, thanks to our reinterpretation of “C.maortuensis. Just the other day our paper on this animal was published, in which we name it a new genus (Shaochilong, meaning “shark teeth dragon” in Chinese, in obvious reference to the carcharodontosaurids), reinterpret it as the first unequivocal Asian carcharodontosaurid known from the fossil record, and discuss its implications for Asian theropod evolution. In a nutshell, Shaochilong is further evidence that carcharodontosaurids were a truly global group, and that the large-bodied theropod faunas of Early-mid Cretaceous Asia were cosmopolitan in nature, not endemic as had been the case during the Middle-Late Jurassic.

Carcharodontosaur phylogeny showing their relationships, ages and continent of origin. In the upper left you can see the appearance of the large bodied tyrannosaurids and how late they appear compared to the carcharodontosaurs. From Brusatte et al., 2009.

Carcharodontosaur phylogeny showing their relationships, ages and continent of origin. In the upper left you can see the appearance of the large bodied tyrannosaurids and how late they appear compared to the carcharodontosaurs. From Brusatte et al., 2009.

Not only that, but the 60 million year gap in the Asian large-theropod record had been a thorny issue. We knew that large basal tetanurans lived in Asia during the Middle-Late Jurassic and that colossal tyrannosaurids dominated ecosystems during the last 10 million years of the Cretaceous. But what large predators lived in Asia during the intervening gap? Were these predators late holdovers from the basal tetanuran lineages, or did large tyrannosaurids actually originate much earlier than observed in the fossil record? Shaochilong is only a single data point, but it suggests that basal tetanurans such as carcharodontosaurids still held sway in Asia deep into the Cretaceous, and that tyrannosaurids experienced a relatively rapid ascent into the top predator role much later in the Cretaceous.

But now the really fun part begins. We need to test these ideas with other fossils, so the race is on to discover more large theropods from the once impenetrable middle Cretaceous gap of Asia. What new discovery lurks within the next outcrop…or within the next museum drawer?

*I would like to acknowledge the help of my co-authors on the project: Roger Benson, Dan Chure, Xu Xing, Corwin Sullivan, and some paleo blogger called Dave Hone [who he? ed]. I also thank Dave, Corwin, and Xu for their hospitality in China, and my travel companions (Mark Norell, my PhD advisor and source of funding, and Gabe Bever and Mick Ellison, my two favorite valentines).
Brusatte, S., Benson, R., Chure, D., Xu, X., Sullivan, C., & Hone, D. (2009). The first definitive carcharodontosaurid (Dinosauria: Theropoda) from Asia and the delayed ascent of tyrannosaurids Naturwissenschaften DOI: 10.1007/s00114-009-0565-2

51 Responses to “Guest Post: Shark toothed theropods in Asia – introducing Shaochilong”

  1. 1 Andrea Cau 04/06/2009 at 1:39 pm


    Finally, the “C.” maortuensis has been re-studied!
    Now, it’s time someone could re-study Labocania…

  2. 2 David Hone 04/06/2009 at 1:47 pm

    There’s a hell of a lot of taxa that need working on and I’m doing a couple with various people as I have mentioned before, though taxonomy is not my main interest, it’s important to keep these things ticking over. As I’ve said before, we badly need more taxonomists, in biology as a whole, not just theropod research.

  3. 3 Andrea Cau 04/06/2009 at 2:11 pm

    I agree.
    I’ve cited Labocania because it has been suggested to be a close relative of “C.” maortuensis. If confirmed, it would imply a Latest Cretaceous (mid-sized?) carcharodontosaurid in North America.

  4. 4 David Hone 04/06/2009 at 2:58 pm

    Would that be a big surprise given the presence of Acrocanthosaurus? I know that’s a big taxon, but we do have carchs in the middle Cretaceous of North America already.

  5. 5 Andrea Cau 04/06/2009 at 3:42 pm

    “Big surprise”? No, I find it interesting, not more “surprising” than any other new study.
    The presence of more than one lineage of large-sized predatory theropods in Latest Cretaceous of Laurasia may indicate that the Asiamerican ecosystems were not so different (in term of theropod disparity) than the other terrestrial Mesozoic (both Laurasian and Gondwanan) communities, where we usually find two or more sympatric lineages of big theropods (see for example, the Morrison Fm., the Wealdien Supergroup, the Kem Kem Beds…).

  6. 6 Manabu Sakamoto 04/06/2009 at 4:28 pm

    I had an opportunity to look at “C.” maortuensis when I visited the IVPP a few years ago, but I was not shown the maxilla…I didn’t even know there was one.

  7. 7 David Hone 04/06/2009 at 4:43 pm

    There’s not much of it there, but enough. I should probably add at this point that there is a longer description in the works as well as this paper with further anatomical details and extra photos etc.

  8. 8 Mike Taylor 04/06/2009 at 5:41 pm

    Great stuff, Steve (and co-authors!)

    Keep ’em coming!

  9. 9 Thomas R. Holtz, Jr. 04/06/2009 at 8:41 pm

    Man,that was a fast turn around from review to publication!!!

    And glad that Chure was in on this: he did do the preliminary redescription in his dissertation.

  10. 11 David Marjanović 04/06/2009 at 9:19 pm

    Fascinating. Someone mention it on the DML already!

    the worldwide Bristol mafia

    Which covers everything with bentonite!!!1!

    Would that be a big surprise given the presence of Acrocanthosaurus? I know that’s a big taxon, but we do have carchs in the middle Cretaceous of North America already.

    That’s not why. The major surprise here is that Shaochilong is the first certain carcharodontosaurid that’s younger than the Cenomanian-Turonian boundary mass extinction event. (A few teeth from South America were thought to count, but they could just as well belong to abelisaurids according to, I think, an SVP meeting abstract from last year.) So far, the most parsimonious hypothesis was that the carcharodontosaurids snuffed it at that mass extinction, and that Asiamerica was then free for the tyrannosaurs to take over. Apparently that’s not what happened…

    • 12 David Hone 04/06/2009 at 10:52 pm

      But that’s not what Andrea was referring to. He was talking about having multiple lineages of large predators, which I don’t think is necessarily a big surprise if Acrocanthosaurs is already present and overlapping with some tyannosaurids.

  11. 13 Steve Brusatte 04/06/2009 at 9:44 pm

    Thanks for your comments, everyone! To answer Andrea’s question about Labocania, I haven’t seen the specimen, but looking at the published information it’s hard to gauge much about it. Dan Chure noted some similarities between Shaochilong and Labocania in his thesis (mostly related to the interdental plates), but he wrote his thesis over a decade ago and we’ve learned quite a bit about large theropods since then. I see no clear reason to consider Labocania a carcharodontosaurid (no obvious synapomorphies), but I would want to see the specimen first. It would be cool to have a late-surviving carch in North America (or, even cooler would be if Labocania is an abelisaurid!).

    Cheers, Steve

    • 14 Andrea Cau 04/06/2009 at 11:26 pm

      Thank you, Steve.

      Dave, are there tyrannosaurids overlapping with Acrocanthosaurus? (For clarity, I use Tyrannosauridae = Tyrannosaurinae + Albertosaurinae)
      I know that basal mid-sized tyrannosauroids (Xiongguanlong) were present in the “mid-Cretaceous”, but not “true” Tyrannosaurids. For example, the same calibrated phylogeny present in Brusatte et al. (2009) and in this post shows a big time gap between Acrocanthosaurus and tyrannosaurids.

      • 15 David Hone 05/06/2009 at 12:00 am

        Sorry that should probably have been tyrannosauroids (don’t have access to my research stuff, I’m at home), but Stokesosaurus is in the Late Jurassic I think, so we should have medium to large bodied tyrannosaurs (of some grade) in the Early Cretaceous of North America. I’d need to check references and just don’t have them right now so amy be in error, ut in general I think it’s reasonable to think tey would be present.

      • 16 Thomas R. Holtz, Jr. 05/06/2009 at 12:19 am

        There are various mid-sized tyrannosauroids from the Middle Jurassic on up to the early Late Cretaceous, including Guanlong, Stokesosaurus, Eotyrannus, Xiongguanlong, and bits an pieces here and there. But these are typically dwarfed by spinosauroids and/or allosauroids in their same formations.

        And even the immediate outgroups to Tyrannosauridae (Appalachiosaurus, Alectrosaurus, Dryptosaurus)–while larger than their precursors–are still noticeably smaller than Tyrannosauridae proper. (Sort of like how Zuniceratops is larger than “protoceratopsians” but smaller than members of Ceratopsidae).

      • 17 David Hone 05/06/2009 at 12:39 pm

        Tom I guess it depends on how we define ‘big bodied’ predators (returning to Adrea’s original comment). I’d still be happy to say something like Stokes. was pretty big and even without direct evidence there were likely equally large or more likely larger tyrannosdauroids of some kind running around at the same time as Acrocantho. Depends on your definitions really I think.

  12. 18 David Hone 04/06/2009 at 10:26 pm

    The Bristol Mafia, I would note is very, very extensive. We have guys in Japan, China, Mexico, U.S.A., Germany, France, Ireland, Brazil, Denmark, Canada, South Africa, Finland, Italy and Panama and that’s just off the top of my head. Oh yeah, and Australia (sorry Roger).

  13. 19 J. S. Lopes 04/06/2009 at 10:46 pm

    Maybe Asian carcharodontosaurids came from some Gondwanan microplate split from Eastern Gondwana (probably near Australia-Melanesia) in Latest Jurassic or Early Cretaceous, later colliding into Southeast Asia.
    Or, alternatively from some route Africa-Europe-Southwestern Asia, or Africa-Europe-North America-Asia.

  14. 20 Thomas R. Holtz, Jr. 04/06/2009 at 11:55 pm

    Andrea: oldest Tyrannosauridae proper and immediate outgroups (like Appalachiosaurus) are mid-Campanian. Alectrosaurus may be older, but its position remains unresolved and it is quite likely outside Albertosaurinae + Tyrannosaurinae.

    The big tyrants were very much latecomers on the stage…

  15. 21 DinoHunter 05/06/2009 at 12:26 pm

    I doubt anyone will figure out what Labocania is. I was at the Mexico University with Dan Chure when he went to study it (I was looking at other things, Lambeosaurus? laticadus). It is so poorly preserved I doubt it’ll ever be understood. Dan was really bummed when he saw it. Also, while I had some time I went looking through the collections and found the ‘square’ toothed tyrannosaurid premaxillary teeth that Dan and I described years ago.

  16. 22 DinoHunter 05/06/2009 at 12:29 pm

    If the Burpee Tyrannosaur volume ever gets published my paper on the geologic history of tyrannosaurids would be in it. Its a chronologic history of tyrannosaurids from the Middle Jurassic to the latest Cretaceous. My reviewer accepted the paper with only a few changes (to my surprise). Hopefully the volume will be published.

  17. 23 DinoHunter 05/06/2009 at 12:32 pm

    Brusssate wrote;
    I see no clear reason to consider Labocania a carcharodontosaurid (no obvious synapomorphies), but I would want to see the specimen first. It would be cool to have a late-surviving carch in North America (or, even cooler would be if Labocania is an abelisaurid!).<<

    If it were either one, then it'd have been living with tyrannosaurids in the same formation. Another interesting fact is that during the Campanian Baja was down by the souther tip of Mexico, which makes it the furthest south Tyrannosaurid and possibly the furthest north abelisaurid (if it is an abelisaurid).

  18. 24 DinoHunter 05/06/2009 at 12:34 pm

    Dave, are there tyrannosaurids overlapping with Acrocanthosaurus? (For clarity, I use Tyrannosauridae = Tyrannosaurinae + Albertosaurinae)
    I know that basal mid-sized tyrannosauroids (Xiongguanlong) were present in the “mid-Cretaceous”, but not “true” Tyrannosaurids. For example, the same calibrated phylogeny present in Brusatte et al. (2009) and in this post shows a big time gap between Acrocanthosaurus and tyrannosaurids.<<

    Ah, if only my aftermentioned article was published…


  19. 26 Mickey Mortimer 06/06/2009 at 10:44 am

    Very cool looking paper. It’s great to see you getting so many theropods redescribed. As for Labocania being a carcharodontosaurid, I coincidentally noted such a possibility on my site-

    “Another so far unpublished possibility is a relationship to carcharodontosaurids, as Mapusaurus has a pneumatic quadrate, lateral dentary shelf, proximolateral ischial fossa and triangular obturator process.”

    I never thought Shaochilong (still “Alashansaurus” on my site) could be carcharodontosaurid though, so that is quite interesting. I’ll have to update my site to reflect the new data once I see the paper.

  20. 27 vvvvv 22/06/2009 at 11:10 pm

    How big is shaochilong? Is it comparable to giganotosaurus or is it in the eocarcharia-acrocanthosaurus size range? How heavy was it?

    • 28 David Hone 23/06/2009 at 7:43 am

      I’ll leave Steve to do the real deatila, but we are talking soemthing Allosaurus sized (say 8-10 m ish) not Giganotosaurus sized.

  21. 29 vvvvv 26/06/2009 at 8:03 pm

    How large and heavy was Shaochilong? I want to know whether it coexisted with basal tyrannosauroids. Also, I want to know=
    how long did it take a titanosaur to grow up into an adult? How big was Andesaurus delgadoi? What were the other Maastrichtian titanosaurs living with Puertasaurus, Epachthosaurus and saltasaurus and others?

    Also, I want to know how many times a titanosaur’s rib equals its body length (I’m trying to calculate the length of Huanghetitan ruyangensis). COuld you help me?

    • 30 David Hone 26/06/2009 at 8:45 pm

      You are asking a lot of difficult and techncial questions. I suggest you put this up on te Dinosaur Mailing List or on Ask A Biologist or somewhere like that. The Shaochilong papaer covers some of this information and others are available online that will answer soem of your questions if you look.

  22. 31 vvvvv 26/06/2009 at 8:09 pm

    I also want to know what kind of environment the South American cenomanian-turonian dinosaurs (Giganotosaurus, Argentinosaurus, Limaysaurus, Buitreraptor…blah blah blah) lived in. What were the plants in the area? What kind of environment did Shaochilong and its contemporaries live in anyway?

  23. 32 vvvvv 06/08/2009 at 1:20 pm

    What exactly sets Chilantaisaurus apart from Shaochilong? I read in the Wikipedia Spanish Chilantaisaurus page that it was a member of the spinosauroidea, and was 11 meters lnog, and 4 meters high. Also, in the Wikipedia English page, there is something else, that though it was difficult to rule out the possiblity that they were the same, the two had ‘an enormous size difference’. So who’s correct, and why has Chilantaisaurus been so…obscure in classification(if what I’m putting here in my words is correct…)?

  24. 33 vvvvv 08/08/2009 at 4:42 pm


    • 34 David Hone 08/08/2009 at 5:33 pm

      VVVVV, please do not act like this. I don’t really know the answer to your question and as you may have noticed I have been extraordinarily busy this last week. I did ask Steve to post an answer up here since this is his speciality though he himself is away on his honeymoon right now so is probably not checking his e-mail or will consider this a priority. If you really need an answer i suggest you ask someone for the original description, or put your question to other people like on AAB, or the DML or Vert Pal mailing list.

      Posting demanding comments like that will not get the answer any faster and as you might have guessed will not endear you to people who might otherwise help.

  25. 35 vvvvv 17/08/2009 at 11:48 am

    Oh, I’m sorry if I was a little too demanding, but I’m just..well…I’M A 13-YEAR-OLD KID! I just…you know, am a dinosaur enthusiast, and love to be a paleo-artist, or whatever you call it, and I’m curious to know more, that’s all. I’m sorry , though.

    • 36 David Hone 17/08/2009 at 12:05 pm

      That’s Ok. It is good to ask questions, but there are lots of ways of getting an answer. I am sorry Steve has not been around to help out, I think he is still away – I have tried again since I last posted on here. I’ll keep trying but in the meantime you can try the other sources i suggested or even Wikipedia – it’s not great, but it’s not bad. Also remember that not all of your questions can be answered, are at least not very well – the fossil record is far from complete and much research is badly out of date – especially for obscure dinosaurs which is why it has taken us decades to get around to this one for starters. Do keep asking, but be patient or try and do the work yourself – it’ll be great in the long term if you learn hwo to obtain this kind of information.

  26. 37 Steve Brusatte 20/08/2009 at 12:40 am

    Yes, to chime in. I have indeed been away for a few weeks, but it’s now back to work! To answer what is an excellent question: we are not 100% certain that Shaochilong and Chilantaisaurus are different animals, but they most likely are. Each genus is only known from one incomplete specimen, and there are no overlapping bones between the two specimens. But, one important difference is that the size of Chilantaisaurus (judging from the preserved limb bones and vertebrae) is absolutely huge, about the same size as Tyrannosaurus. Shaochilong is a much smaller animal. In fact, it is the smallest known carcharodontosaurid and even smaller than Allosaurus and Sinraptor. Although we cannot compare individual bones of the two animals, we can be pretty sure that they were different because of the major size disparity between them.


  27. 38 vvvvv 23/08/2009 at 12:36 pm

    Well, that’s nice, thanks. So, to what family does Chilantaisaurus fall into?

    Also, how can I measure a Huanghetitan ruyangensis by checking the rib to body ration of the dinosaur?

    • 39 David Hone 23/08/2009 at 3:52 pm

      Well no-one is really sure what Chiliantisaurus actually is at the moment. It’s a tetanuran, but beyond that we don’t know much as the material is so fragmentary.

      As for Huanghetitan, you can’t really tell how big it was from the rib. First of all there is no obvious correlation between ribs and body size (as there is for say the femur) so it would not give you an accurate answer. Secondly the animal as a whole is very fragmentary – there is not very much of it at all and the big rib itself it not complete (I’ve actually seen this specimen and there’s not much of it).

  28. 40 vvvvv 25/08/2009 at 4:29 pm

    Oh, that’s pitiful. You see, it’s just that I’m interested in the dinosaurs nobody really takes much interest of, instead of always the T-rex and Triceratops fare. Well, if I’m ever to reconstruct a Chilantaisaurus at present, I’ll just have to have it looking like a really huge Shaochilong.

    Also, I’d like to know a thing or two about Borealosaurus and gobitian, like when they lived; building up a cenomanian and turonian picture of Asia is difficult, isn’t it?

    • 41 David Hone 29/08/2009 at 10:55 am

      Well lots of people are interested in obscure animals, but there are not many palaeontologists and a huge number of dinosaurs (and huge numbers of specimens and lots of things to do beyond just taxonomy and anatomical descriptions). As such we don’t always have time to do what we want, when we want and how we want.

      Sorry, but I don’t know much about those animlas at all. I suggest you ask the SVPOW guys who are much more up on their sauropods. Good luck with your project!

  29. 42 vvvvv 29/08/2009 at 1:40 pm

    Thanks for the details. Who are the SVPOW guys?

  30. 44 Sebastian 02/09/2009 at 9:02 pm

    Hi guys,

    I am new to this blog,however, I find it very interesting.I’ve seen some comments on Labocania anomala and I agree with Steve that it is the North American abelisaurid theropod. I am redescribing some of the taxa from India, which were previously described by von Huene and Matley 1933. In my paper, which is almost done L.anomala is included in Abelisauridae.All the best to all of you who is going to SVP this year in UK have fun there and good time.

    • 45 Tor Bertin 03/09/2009 at 8:34 am


      If you’re allowed to talk about it, which Abelisaurid synapomorphies does Labocania convey? Finding the Abelisauridae in North America would be awesome, if not biogeographically bizarre.

      If you can’t talk about it I understand though–will gladly wait for the paper if needed. 😀

  31. 46 RaptorX Jackie 23/05/2010 at 2:39 pm

    Well, now, these Chilantai and Shaochi are both Cenomanian-Turonian animals, and allosaurids were the top bananas in Asia even at that time. Well, there’s something confusing going over here?
    Haven’t any chronologically contemporaneous tyrannosaurs ever been noted there?

    Okay, for the record-
    Super-sauropods like Borealosaurus, Ruyangosaurus and Huanghetitan
    Ankylos like Gobisaurus and Talarurus
    Ornithomimids and therizinosaurs
    A king-size raptor- Achillobator (the formation it lived in is finally dated, right?)

    But….looking back at T-rex relatives, we find Aptian and Barremian critters-
    Dilong, Xiongguanlong, Sinotyrannus, Raptorex.

    What’s going on here?

  1. 1 A New Look For Asia's Ancient "Shark Tooth Dragon" | Dinosaur Tracking Trackback on 12/06/2009 at 10:05 pm
  2. 2 Guest post: a new tyrannosaur – Alioramus altai. « Dave Hone’s Archosaur Musings Trackback on 06/10/2009 at 5:41 am
  3. 3 The Archosaur Musings 2009 Awards « Dave Hone’s Archosaur Musings Trackback on 31/12/2009 at 9:04 am
  4. 4 Shenzhousaurus « Dave Hone’s Archosaur Musings Trackback on 07/02/2010 at 12:14 pm
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