Friday, September 27, 2013

Q&A III: Nanotyrannus – my turn in the hot seat.

A cast of the Cleveland Skull next to a cast of an adult Tyrannosaurus rex skull in the striking dinosaur gallery of the Royal Ontario Museum.

What follows are my answers to Mickey Mortimer’s extensive questions regarding my post “Nanotyrannus isn’t real, really”. I think they are excellent and require response, especially since it gives me the opportunity to clarify and expand some points regarding my work on the Cleveland skull, the increasingly irrational defenses of the taxon, recently uncovered specimens, and the inaugural article. However, I presently have three projects in the works that have direct bearing on this taxonomic issue. Therefore, I cannot state the conclusions or share data before they are published. You'll have to read the previous post for the appropriate context.
#1: Notably it's only really engaging with the first and worst arguments for Nanotyrannus' validity.
A diagnosis sets out the evidence for the validity of a taxon; for a systematist, it is transparency at its best. In my mind, there is no reason not to accept the diagnosis of Nanotyrannus (or any other taxon) as, prima facie, the best argument for establishing a scientific name. Also, a diagnosis is a hypothesis that can be tested. This is a crucial point, and my hypothesis - the Cleveland skull is a juvenile T. rex - can be falsified if it can be shown that the diagnosis of Nanotyrannus is defensible. What could be easier? Has anyone done this?
The defenders of Nanotyrannus have not even mentioned the diagnosis in their articles.
Perhaps none of them have given the diagnosis any thought, or they assume that it is perfectly fine. Instead, they have just heaped on new characters as if that’ll do the job of rescuing the taxon. In the end, it does not matter if that blind spot is out of agreement, complaisance, avoidance, ignorance, or indifference; we must all agree that the diagnosis is central to the question of the validity of Nanotyrannus, and it has been on borrowed time.
In the case of Nanotyrannus, the diagnosis is important because if the purported type specimen (the Cleveland skull) is not diagnostic, then there is no taxon to which new specimens can be referred. That is why skeletons such as Jane and the ‘dueling tyrannosaur’ are completely irrelevant to this taxonomic issue. Arguably, the diagnosis isn’t low-hanging fruit; in this case, it’s the whole orchard.
#2: Sure Bakker's 25 year old ideas are flawed, but any modern defense of the genus includes a swath of newly recognized characters, and of course Jane.
There are two flaws that I see in Bakker et al. (1988): the article splits tyrannosaurids into many taxa without first assessing ontogeny, including the splitting of Nanotyrannus from Tyrannosaurus rex; I think the last paragraph in their article reveals the philosophical view that motivated the decision. The second flaw was in not conducting a cladistic analysis of the data listed in the caption of their branching diagram. As such, their phylogenetic scheme is a subjective arrangement and not a true test of character congruence.
Aside from those issues, the rest of the article is an excellent and refreshing review of tyrannosaurid cranial osteology. For example, I use their terminology for the basicranium, because no one before or since has done a better job of clarifying the salient landmarks of that region, or in providing a more useful nomenclature for those features.
If by new characters you mean the ones in Larson (2013), then I cannot say anything more than this: I deal with them in my in-progress work on T. rex ontogeny. That article includes a review of all of the literature and ‘evidence’ advanced against the hypothesis in Carr (1999). As you might guess, when the smoke clears, that a juvenile T. rex is still standing.
Again, specimens such as Jane and the ‘dueling tyrannosaurid’ are irrelevant to the issue unless it can be shown that they are different from juvenile T. rex as represented by the Cleveland skull. In that case, we’d have a new taxon or two, but we wouldn’t have Nanotyrannus.
#3. So yes, Bakker was wrong to think the Nanotyrannus type had extensive cranial fusion. But Larson (2013) reports Jane has presacral neurocentral, scapulocoracoid and pelvic fusion. By not addressing these characters (also including glenoid position, subnarial foramen position, quadratojugal pneumaticity, maxillary tooth count, etc.), your analysis loses most of its force.
My analysis is on the way! In the meantime, I’m going to sound like a broken record here, but I do think it is worthwhile repeating: Jane is irrelevant to the Nanotyrannus issue because (a) it is not the type specimen, (b) the diagnosis of Nanotyrannus is indefensible; therefore, (c) no specimen can be referred to a taxon that does not exist in Nature.
If it can be shown that Jane is a different taxon from the Cleveland skull and adult T. rex, then it becomes the holotype of a new taxon. There is no way that Jane can be referred to Nanotyrannus, because the taxon hasn’t a single diagnostic character. Also, I did propose a hypothesis to account for the difference in tooth count (Carr, 1999).
By the way, I am at the helm of the description of Jane, and we do take care of the purported diagnostic characters listed in recently published articles (e.g., Larson, 2013). As such, my hands are tied on discussing those characters specifically until the article is published.
#4. There's also the problem of using similarity to juvenile stages of other species as an argument. In the 4 stage ontogeny of your 1999 paper, Albertosaurus simply never gets to stage 4. It stops at what is stage 3 in Daspletosaurus. Indeed, in 1996, you used this to argue Daspletosaurus are just adult Albertosaurus. You later changed your mind on this, but couldn't Nanotyrannus be a tyrannosaurine that never grows past stage 1? This makes your huge table moot, just as a huge table of stage 3 characters listed as "Yes!" in Albertosaurus would not mean it was a subadult Daspletosaurus.
The goal of Carr (1999) was to establish a baseline of what the primary growth stages of a  tyrannosaurid looks like. In this case, Albertosaurus libratus was the point of reference given the high number of specimens and completeness of the growth series. It turns out that the growth series can be blocked out into four stages (small stage 1, large stage 1, stage 2, stage 3). In this series, the stepwise transformation of the juvenile features into adult can be clearly seen. I then assumed, based on common ancestry, that this pattern is plesiomorphic for Tyrannosauridae.
Working under that parsimonious assumption, I then reviewed the evidence for pygmy tyrannosaurids, such as Nanotyrannus and Maleevosaurus (it is curious that people are a lot less upset about my hypothesis that Maleevosaurus is a juvenile T. bataar). It turns out that both have the same features as juvenile A. libratus; ergo, they must be juveniles. Since that time, every juvenile of other derived tyrannosauroids (e.g., Bistahieversor, Daspletosaurus, T. bataar) that have come to light have the same constellation of juvenile features as are seen in A. libratus, the Cleveland skull, and "Maleevosaurus". If the Cleveland skull had characters seen in adults of A. libratus, Daspletosaurus, T. bataar, or T. rex, then that would be an indication that it might be a pygmy taxon and my hypothesis would have been different, perhaps validating Nanotyrannus.
As the Cleveland skull and juvenile Daspletosaurus (Currie, 2003) show, tyrannosaurines have the same starting point as A. libratus; this is consistent with the hypothesis that the growth pattern is conserved (i.e., plesiomorphic) in Tyrannosauridae (Carr, 1999). The calibration of growth series is a major part of a manuscript that I have in progress, some of this I have presented at SVP meetings over the past few years.
As such, it turns out that tyrannosaurids went through the same ontogenetic changes in the skull. In the table I present (implicitly) the hypothesis that Daspletosaurus is peramorphic relative to A. libratus, in that the growth trends are carried further in the tyrannosaurine. That is why Daspletosaurus is under the heading “Stage 4”. In the same vein, T. rex is peramorphic relative to Daspletosaurus.
The salient point you make is the suggestion that the Cleveland skull is an adult, but it does not grow past stage 1. I think it is reasonable to expect that an adult of a true ‘pygmy’ tyrannosaurid would have all of the adult characters seen in its closest relatives. The Cleveland skull certainly does not have any adult features - of any derived tyrannosauroid - that I can see.
On a related note, the original manuscript for Carr (1999) did not have the summary table of characters. One of the insightful reviewers required that I put the reams of features mentioned in the text in one, easy-to-follow, table. I like the outcome: the table of growth characters is a useful tool that people can use for a quick assessment of the relative maturity of any new tyrannosaurid specimens.
In the table, each column of characters can be thought of as analogous to a series of lines of arrested growth. I have since tested the data for hierarchical structure using parsimony analysis for T. rex (Carr and Williamson, 2004) and A. sarcophagus (Carr, 2010), which was found to be present. A hierarchy from this sort of data is best explained by ontogeny. As such, the table – and the cladistic analyses drawn from it – summarizes the hard-wired sequence of growth changes the members of the clade inherited, and altered, from their recent common ancestor.
#5. A related point is that Larson agrees Nanotyrannus could be the sister taxon of Tyrannosaurus, so your long list of characters they share works well for both hypotheses. We don't need to assume Bakker's odd phylogeny with a basal Nanotyrannus convergent with Tyrannosaurus for the genus to be valid.
Could Nanotyrannus could be a juvenile specimen of the sister taxon of T. rex? For that to be defensible, it has to be shown that the purported new taxon is diagnostic. The weight of evidence shows that it is a juvenile T. rex, so that avenue of thought doesn’t rescue the identity of the Cleveland skull from T. rex.
With regard to Bakker et al.’s (1988) tyrannosaurid phylogeny, I do think that context matters. Their arrangement complements what they say in the text, together showing the deductions that led to their taxonomic decision. It is an excellent article in that the data are absolutely clear.
#6. Further, it implies a taxon can be invalidated merely by finding the original diagnosis wanting. Let's see how that works for Tyrannosaurus, based on Osborn's (1905) original diagnosis.
1. "Carnivorous Dinosaurs attaining very large size." Tarbosaurus got as large as the average Tyrannosaurus, so this is invalid even within Tyrannosauridae.
2. "Humerus believed to be of large size and elongate (Brown)." This was based on an incorrectly identified bone.
3. "No evidence of bony dermal plates (Brown)." This was to distinguish it from Dynamosaurus, whose holotype had incorrectly referred plates.
Well, guess Tyrannosaurus is an invalid taxon we can't refer the Cleveland skull to...
Is there any serious reason to doubt the validity of T. rex as there is for Nanotyrannus? The diagnosis issue has to be taken on a case-by-case basis; there is no reason to think that T. rex is invalid, given the quality of specimens (including the type) and sample size that has been amassed. Certainly, the time has come to bring the diagnoses for T. rex and many other taxa up to date, as I did for Albertosaurus sarcophagus (Carr, 2010). I have this in the works for other tyrannosaurids, including T. rex.
#7. Finally, couldn't the Cleveland skull be a juvenile and a valid genus, just without known adult specimens? That's what you argue for in Alioramus, which is (1) based on two specimens of juvenile tyrannosaurid, (2) has an original diagnosis which relies entirely on characters claimed to be ontogenetic in the case of Nanotyrannus (long snout, high tooth count, compressed teeth, etc.), (3) has derived characters only known in the sympatric Tarbosaurus, so is by your reasoning (4) therefore juvenile Tarbosaurus. Yes, Alioramus has additional suggested apomorphies, but so does Nanotyrannus and they seem to be of equal import (e.g. additional pneumatic features in each). So I'd like to know why you came to divergent conclusions in these similar cases.
I was not at the helm of that project, which accounts for the difference in results (this also pertains to your recent follow-up comment on Alioramus).
#8. Is that really fair to paleontologists in 1988? Almost no theropod specimens were even recognized as juveniles, besides some Gallimimus and Allosaurus individuals. The few known juvenile tyrannosaurids (e.g. the then unnamed Jordan theropod and Shanshanosaurus) were thought to be small basal tyrannosauroids or even dromaeosaurids. Similarly, there was no consensus the smaller Nemegt tyrannosaurids were young bataar. AFAIK, no one had thought to look for immature bone grain in any Mesozoic dinosaur yet, and there was no consensus of (Alberto(Daspleto(Tarbo,Tyranno))) like we have now to use to find nested traits that suggest synonymy. The sample size of Tyrannosaurus was low enough to make the number of specimens a notable media figure, when now we have more than most species of theropod known from more than just their holotype. The argument Nanotyrannus is a juvenile Tyrannosaurus thus depends on a lot of data which didn't exist in 1988 or wasn't organized yet. Rozhdestvensky (1965) was remarkably prescient decades earlier regarding some of these issues, though his same methodology incorrectly found Lufengosaurus to be young Yunnanosaurus, so we might question whether his seemingly correct tyrannosaurid ideas were validly justified or at least partly luck.
Yes, it is fair because sufficient data was at hand (e.g., Rozhdestvensky, 1965), and there were two growth series (A. libratus, T. bataar) available with which to make comparisons. Bakker et al. (1988) had the opportunity to define tyrannosaurid ontogeny, despite the stumble over suture closure in the Cleveland skull.
#9. It's that old standing on the shoulders of giants cliche, and while you are a major giant here, suggesting Currie, Russell, Molnar, Paul, Carpenter, Bakker, etc., basically every theropod worker of the 80s, was unreasonable seems unfair.
 I did not cast shade over the entire lot; I just just think Bakker et al. (1988) were unreasonable in their approach to tyrannosaurid taxonomy; they split taxa before assessing ontogeny. Aside from that, sincerely, it is an excellent osteological work.
#10. A valid criticism is that Bakker et al. (1988) never cite or mention Molnar's (1980) description of what would become the type of Dinotyrannus, that was referred to lancensis at the time. It was published 8 years earlier in a well read journal, so should have gotten some comment. 
That’s a good point! I have to admit that I was late on the scene as well (Carr and Williamson, 2004).
#11. The supposed unreasonableness of Bakker et al. (1988) also affects your main argument, as some of it depends upon the truth of our current consensus. Accepting Nanotyrannus as a juvenile Tyrannosaurus seems more plausible in a world where Maleevosaurus is just a young Tarbosaurus, Stygivenator is a juvenile Tyrannosaurus, etc.. But if the well-studied Cleveland skull is valid based on newly recognized characters, what does this mean for Maleevosaurus, which has never been studied firsthand since its description to my knowledge? Larson considers Stygivenator a younger Nanotyrannus, for instance, so the former can no longer be used as an example of what juvenile Tyrannosaurus are like.
The bottom line is that the relative maturity of specimens and their taxonomic identity must be considered separately, so a change in the decision regarding the Cleveland skull does not reverse the decision on Maleevosaurus; it doesn’t change the fact that both have the same constellation of juvenile features that are seen in all other derived tyrannosauroids, nor does it change the fact that the Cleveland skull has features that are only seen in T. rex. Williamson and I (2004) showed that ‘Stygivenator’ is referable to T. rex, but Larson (2013) did not engage with the evidence that we presented.
In the same fashion, the Larson (2013) article did not deal with the diagnosis issue in Bakker et al. (1988), or the evidence presented in Carr (1999). Therefore, the taxonomic decisions in that article carry no weight because the validity of Nanotyrannus is uncritically assumed to be valid despite all of the evidence to the contrary.
I trust all of that satisfies your query.

References cited
Bakker, R.T., Williams, M., and Currie, P.J.  1988.  Nanotyrannus, a new genus of pygmy tyrannosaur, from the latest Cretaceous of Montana. Hunteria 1: 1-30.

Carr, T. D.  1999.  Craniofacial ontogeny in Tyrannosauridae (Dinosauria, Theropoda). Journal of Vertebrate Paleontology 19:497-520.
Carr, T. D. 2010. A taxonomic assessment of the type series of Albertosaurus sarcophagus and the identity of Tyrannosauridae (Dinosauria, Coelurosauria) in the Albertosaurus bonebed from the Horseshoe Canyon Formation (Campanian–Maastrichtian, Late Cretaceous). Canadian Journal of Earth Sciences 47:1213-1226.
Carr, T. D. and T. E. Williamson. 2004. Diversity of Late Maastrichtian Tyrannosauridae from western North America. Zoological Journal of the Linnean Society 142:479-523.
Currie, P.J. 2003. Cranial anatomy of tyrannosaurid dinosaurs from the Late Cretaceous of Alberta, Canada. Acta Palaeontologica Polonica 48(2): 191-226.
Larson, P. L. 2013.  The case for Nanotyrannus in J. M. Parrish, R. A. Molnar, P. J. Currie., and E. B. Koppelhus (eds.) Tyrannosaurid Paleobiology, University of Indiana Press, Bloomington and Indianapolis, pp. 15-53.
Rozhdestvensky, A. K. 1965. [Growth changes in Asian dinosaurs and some problems of their taxonomy]. Palaeontological Zhurnal 1965:95-100. (Translated from the Russian.)


  1. Can't wait reading your articles!

  2. Very we commented upon! I too cannot wait till your future Tyrannosaurid papers are published. Thank you Thomas and Mickey!

  3. For context, Mickey's original comment was here in response to Tom Carr's post.

    That said, I really appreciate the level of engagement taking place on this blog. More to say in a bit on the actual post...

  4. Fusion between elements was mentioned as possibly reason to think that "N" is fully grown but fusion is not a reliable indicator of the termination of growth or age in archosaurs-- and fusion is apparently variable within individuals. See papers by Ikejiri, Brochu (and also new SVP abstracts for more) -- and this is apparently the case in turtles as well at least in the postcrania (pers. obs.). Even in mammals with pathological fusion, the elements still continue to grow so this may not be as hard and fast as we assume (Johnson and Southwick 1960; Ritsila and Alhopuro 1975). We need to be careful as well - we don't necessarily know what could be happening during growth with hormones in dinosaurs - hormones can influence different tissue types to transform variably between each other (Steinetz et al. 1965)

    to be honest, this makes sense if we think about elements which fuse early with each other, esp those that form a complex bone with different ossification types and origins (parasphenoid-basisphenoid for example).

  5. I'm enjoying this a lot. Because my reply was so extensive, I made it a post on my own blog- . The basic points seem to be- we have different philosophical viewpoints; most debate on Nanotyrannus can't move forward until your papers are published so that you can comment on Larson's (2013) details; we can discuss the high tooth count though; and your views on Alioramus are vague but intriguing.

  6. I really enjoy this healthy discussion about this subject and look forward to the new papers.
    I was just wandering: What is the actual status of the "Tinker" specimen ? It was originaly described as a juvenile T.rex if I remeber correctly. Do we have more date about his specimen and its implications ?
    Thanks in advance.

    1. My website has the following info-

      (private coll.; Tinker) (~8 m; subadult) premaxillae, maxillae, partial nasal, jugal, parietal, squamosal, quadratojugals, quadrate, palatine, pterygoid, dentary, splenial, coronoids, surangular, angular, preartcular, articulars, teeth, two cervical ribs, five dorsal ribs, rib fragments, twenty partial caudal vertebrae, twelve chevrons, partial scapulae, coracoid, humeri, manual ungual, incomplete ilia, pubes, ischium (650 mm), tibia (670 mm), pedal ungual (Larson, 2008)

      In 1998, a subadult Tyrannosaurus was discovered in the Hell Creek Formation of South Dakota and nicknamed Tinker. Although touted as a juvenile in the press releases, Tinker is much larger than the 'Nanotyrannus' specimens CMNH 7541 and BMRP 2002.4.1, almost the size of the Dinotyrannus holotype. It is therefore unsurprising it possesses a low number of mediolaterally thick teeth characteristic of older tyrannosaurids, instead of the narrower more numerous teeth of 'Nanotyrannus' specimens. Interestingly, the latter type of tooth was found associated with Tinker, perhaps suggesting scavenging by younger Tyrannosaurus individuals or social behavior. Blasing (DML 2006) stated that another young Tyrannosaurus (nicknamed Belle) and remains of an adult were present in the jackets with Tinker. Unfortunately, Tinker was not deposited in a museum and the hired preparator declared bankruptcy, so the specimen is in storage in Pennsylvania as of 2006. For a time, Harding County, SD owned Tinker, as the lease between it and the people working on the fossil was declared invalid, though the collector's regained ownership in February 2008. It's unknown if or when Tinker will be available for scientific study.

  7. Carr, in SVP 2013, Larson has stated that Nanotyrannus has third digit manus phalanges. If this is true, I would believe that this is not a juvenile T.rex and even not a tyrannosaurid.
    But I have seen nothing about this before -even in Larson's 2013 article "The case for Nanotyrannus". And I have never seen any third digit manus phalanges in Jane specimen and even in dueling tyrannosaur photos!
    Is it true that Nanotyrannus specimens have third digit manus phalanges? I know that tyrannosaurids have third digit manus metacarpal, but a phalanges? That looks really weird to me..
    And anyways, I have really enjoyed reading your SVP 2013 article!

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