Guest Post - Roland Sookias Discusses His New Study Examining How Dinosaurs Came to Fill Most Ecological Niches During the Mesozoic

Why were dinosaurs, and other archosauromorphs (the group of animals including crocodiles, dinosaurs, pterosaurs and several other extinct groups), so big? Was natural selection for increasing size responsible for archosauromorphs’ dramatic rise to larger sizes and did selection for decreasing size drive therapsids’ (‘mammal-like reptiles’, which were the dominant land vertebrates before archosauromorphs) reduction in size during the Triassic (see picture)? These are the questions which my, Richard Butler’s and Roger Benson’s recent publication in Proceedings of the Royal Society B – “Rise of dinosaurs reveals major body size transitions are driven by passive processes of trait evolution” – attempts to answer.
Most of the work in the paper was done as part of my MSc thesis project, which Richard, Roger and Andrew Smith supervised. To carry out the project I spent a good deal of last summer collecting femur and skull length measurements (which we used as proxies for body mass) in the Natural History Museum Library, London. Though barely seeing daylight for a month or two, I managed to collect measurements for ~200 species, which, in combination with data from Benson et al. 2011 got us to >400 species in total. To answer the questions above we focused on getting data for archosauromorphs and therapsids from the Late Permian to Middle Jurassic. This allowed comparison between the two groups, and the interval brackets the rise of archosauromorphs to become the dominant terrestrial vertebrates, replacing therapsids. Thus it allowed us to look at body size evolutionary dynamics during a major faunal transition.

Archosauromorphs (dinosaurs, crocodiles, pterosaurs and their relatives) increased greatly in maximum size (black line, black triangles) from the Permian to Jurassic, and therapsids (‘mammal-like reptiles’) decreased. However in both this was due to expansion in the size variance (i.e. both size increase and decrease), but with subsequent extinction of larger therapsid species. Figure from Sookias et al. 2012, Proc Roy Soc B.

Once we’d got the data together we analysed them using maximum likelihood model fitting approaches. We tried both phylogenetic – i.e. incorporating evolutionary relationships – and time series (ignoring within-group evolutionary relationships and simply averaging size within time ‘bins’) models. Time series models confirmed that on average archosauromorphs tended to increase across the time interval, and that therapsids got smaller. However when we included phylogeny (evolutionary relationships) we found that there was no directional trend in either group along individual lineages. Thus the apparent trends through time were in fact due to ‘passive expansion’ in size, but as the original size was nearer the bottom than the top of the eventual size range the average size tended to increase (see picture). We thus can say that the long-repeated idea of “Cope’s rule” – that taxa in a clade tend to get larger over time due to within-lineage natural selection for larger body sizes – is not found in either archosauromorphs or therapsids during this time interval.

Our work excludes larger size in archosauromorphs as an explanation for their success, as if larger size was especially beneficial one would expect a directional evolutionary trend towards larger sizes. Instead, archosauromorphs probably replaced therapsids opportunistically, as many have hypothesized before. However, the exceptionally high growth, and thus reproductive, rates of archosauromorphs may have allowed them to re-fill empty ecological niches especially easily and rapidly after they went empty due to extinction of therapsids. Thus, while size and growth rate probably did not allow archosauromorphs to outcompete therapsids, it did allow them to fill up free niches quickly.

We also found that archosauromorph predators exceeded the size of the largest herbivores – anomodont therapsids – during the Middle-early Late Triassic. This finding – that the largest carnivores are larger than herbivores - is extremely rare in ecosystems throughout time. It demonstrates that extinct archosauromorphs really were exceptionally large, and that they were able to grow to larger sizes than therapsids given the same resources.

Well, there’s not much more to say about that paper except hope you enjoy it! However, we should be publishing some more work based on my MSc thesis in the near future, so stay tuned, and I’ve just started a PhD with Richard Butler on the early archosauromorph radiation, so hopefully I’ll be involved in answering a few more interesting questions in the coming years. Finally, a very big thank you to Bill Parker for giving us a guest slot here on the esteemed Chinleana.

The paper’s full citation is:


Sookias, R. B., Butler, R. J., Benson, R. B. J. (2012). Rise of dinosaurs reveals major body size transitions are driven by passive processes of trait evolution. Proceedings of the Royal Society B.
doi: 10.1098/rspb.2011.2441 



Abstract- A major macroevolutionary question concerns how long-term patterns of body-size evolution are underpinned by smaller scale processes along lineages. One outstanding long-term transition is the replacement of basal therapsids (stem-group mammals) by archosauromorphs, including dinosaurs, as the dominant large-bodied terrestrial fauna during the Triassic (approx. 252–201 million years ago). This landmark event preceded more than 150 million years of archosauromorph dominance. We analyse a new body-size dataset of more than 400 therapsid and archosauromorph species spanning the Late Permian–Middle Jurassic. Maximum-likelihood analyses indicate that Cope’s rule (an active within-lineage trend of body-size increase) is extremely rare, despite conspicuous patterns of body-size turnover, and contrary to proposals that Cope’s rule is central to vertebrate evolution. Instead, passive processes predominate in taxonomically and ecomorphologically more inclusive clades, with stasis common in less inclusive clades. Body-size limits are clade-dependent, suggesting intrinsic, biological factors are more important than the external environment. This clade-dependence is exemplified by maximum size of Middle–early Late Triassic archosauromorph predators exceeding that of contemporary herbivores, breaking a widely accepted ‘rule’ that herbivore maximum size greatly exceeds carnivore maximum size. Archosauromorph and dinosaur dominance occurred via opportunistic replacement of therapsids following extinction, but were facilitated by higher archosauromorph growth rates.



Popular press coverage:

http://news.discovery.com/animals/how-dinosaurs-got-so-big-120131.html

http://news.sciencemag.org/sciencenow/2012/01/the-secret-of-dinos-success.html?ref=hp

Pampadromaeus barberenai, a New Basal Sauropodomorph from the Late Triassic of Brazil

For those of you who were at SVP this year, this is the new dinosaur taxon from the Triassic of Brazil that Max Langer presented on.

Cabreira, S. F., Schultz, C. L., Bittencourt, J. S., Soares, M. B., Fortier, D. C., Silva, L. R., and M. C. Langer. 2011. New stem-sauropodomorph (Dinosauria, Saurischia) from the Triassic of Brazil. Naturwissenschaften (advance online publication) DOI: 10.1007/s00114-011-0858-0

Abstract - Post-Triassic theropod, sauropodomorph, and ornithischian dinosaurs are readily recognized based on the set of traits that typically characterize each of these groups. On the contrary, most of the early members of those lineages lack such specializations, but share a range of generalized traits also seen in more basal dinosauromorphs. Here, we report on a new Late Triassic dinosaur from the Santa Maria Formation of Rio Grande do Sul, southern Brazil. The specimen comprises the disarticulated partial skeleton of a single individual, including most of the skull bones. Based on four phylogenetic analyses, the new dinosaur fits consistently on the sauropodomorph stem, but lacks several typical features of sauropodomorphs, showing dinosaur plesiomorphies together with some neotheropod traits. This is not an exception among basal dinosaurs, the early radiation of which is characterized by a mosaic pattern of character acquisition, resulting in the uncertain phylogenetic placement of various early members of the group.

The Buck-toothed Demon Reptile, Daemonosaurus chauliodus from the Upper Triassic of New Mexico

Historically it was long thought that the Upper Triassic Coelophysis Quarry at Ghost Ranch New Mexico contained, with the exception of an odd phytosaur, almost exclusively skeletons of the neotheropod dinosaur Coelophysis bauri. However, the recent discoveries of Effigia and Vancleavea demonstrate a much greater diversity of vertebrates present in the quarry. Now we can add to the list a second theropod dinosaur. Daemonosaurus chauliodus ('prominent toothed demon reptile') is known from only a single skull and several associated vertebrae found in the Carnegie Museum block (C-4-81) collected from the quarry in the early 1980s.

Daemonosaurus differs from Coelophysis bauri in the presence of the extreme protruding teeth, a significantly larger prefrontal, and in the dorsoventrally deep premaxilla. A phylogenetic analysis of basal dinosaurs recovers Daemonosaurus outside of Neotheropoda as the sister taxon to Tawa hallae + Neotheropoda.

One possibility I find interesting is that we now have four theropods known from the Upper Triassic in the Ghost Ranch area, Tawa, a coelophysoid, and Chindesaurus from the Hayden Quarry (Petrified Forest Member) and Coelophysis and Daemonosaurus from the Coelophysis Quarry (siltstone member). Currently Chindesaurus lacks a preserved cranium, whereas Daemonosaurus is known almost exclusively from the skull. Thus, it is possible that they could represent the same or a similar taxon. Only the discovery of more complete material of either taxon can test this idea.

Holotype specimen of Daemonosaurus chauliodus in left lateral view

Life reconstruction of Daemonosaurus chauliodus by Jeff Martz

Sues, H.-D., Nesbitt, S. J., Berman, D. S., and A. C., Henrici. 2011. A late-surviving basal theropod dinosaur from the latest Triassic of North America. Proceedings of the Royal Society B. doi:10.1098/rspb.2011.0410 [Free download]

Abstract - The oldest theropod dinosaurs are known from the Late Carnian of Argentina. However, the evolutionary diversification of this group after its initial radiation but prior to the Triassic–Jurassic boundary is still poorly understood because of a sparse fossil record near that boundary. Here, we report on a new basal theropod, Daemonosaurus chauliodus gen. et sp. nov., from the latest Triassic ‘siltstone member’ of the Chinle Formation of the Coelophysis Quarry at Ghost Ranch, New Mexico. Based on a comprehensive phylogenetic analysis, Daemonosaurus is more closely related to coeval neotheropods (e.g. Coelophysis bauri) than to Herrerasauridae and Eoraptor. The skeletal structure of Daemonosaurus and the recently discovered Tawa bridge a morphological gap between Eoraptor and Herrerasauridae on one hand and neotheropods on the other, providing additional support for the theropod affinities of both Eoraptor and Herrerasauridae and demonstrating that lineages from the initial radiation of Dinosauria persisted until the end of the Triassic. Various features of the skull of Daemonosaurus, including the procumbent dentary and premaxillary teeth and greatly enlarged premaxillary and anterior maxillary teeth, clearly set this taxon apart from coeval neotheropods and demonstrate unexpected disparity in cranial shape among theropod dinosaurs just prior to the end of the Triassic.

skeletal outline of Daemonosaurus chauliodus with a human for scale

News coverage of the find here, here, and this "gem". Alas there appears to be a preponderance of "mis**ng l**k" quotes in the many articles posted today.

Citation and Abstract for Nothronychus graffami

Part II of my earlier post. Here is the abstract of the new paper.

Zanno, L.E, Gillette, D.D., Albright, L.B., and A.L. Titus. 2009. A new North American therizinosaurid and the role of herbivory in ‘predatory’ dinosaur evolution.Proceedings of the Royal Society B. Published online before print July 15, 2009, doi: 10.1098/rspb.2009.1029

Historically, ecomorphological inferences regarding theropod (i.e. ‘predatory’) dinosaurs were guided by an assumption that they were singularly hypercarnivorous. A recent plethora of maniraptoran discoveries has produced evidence challenging this notion. Here, we report on a new species of maniraptoran theropod, Nothronychus graffami sp. nov. Relative completeness of this specimen permits a phylogenetic reassessment of Therizinosauria—the theropod clade exhibiting the most substantial anatomical evidence of herbivory. In the most comprehensive phylogenetic study of the clade conducted to date, we recover Therizinosauria as the basalmost maniraptoran lineage. Using concentrated changes tests, we present evidence for correlated character evolution among herbivorous and hypercarnivorous taxa and propose ecomorphological indicators for future interpretations of diet among maniraptoran clades. Maximum parsimony optimizations of character evolution within our study indicate an ancestral origin for dietary plasticity and facultative herbivory (omnivory) within the clade. These findings suggest that hypercarnivory in paravian dinosaurs is a secondarily derived dietary specialization and provide a potential mechanism for the invasion of novel morpho- and ecospace early in coelurosaurian evolution—the loss of obligate carnivory and origin of dietary opportunism.

New North American Therizinosaur Nothronychus graffami

Alright...now I'm up into the Cretaceous rather than the Triassic or even the Jurassic. However, it is good to see this beastie finally published after a lot of hard work and many trials and tribulations. Today (July 15) a paper (I don't have the actual ref yet) comes out in Procedings of the Royal Society B by David Gillette and Lindsay Zanno describing a new therizinosaur from the Tropic Shale of Utah. I was a graduate student at Northern Arizona University when Merle Graffam discovered the holotype specimen. Dave Gillette, one of my graduate advisors, collected the specimen in 1999 and laboriously prepared the specimen in the lab where I was working on the skeleton of Desmatosuchus that was the focus of my Masters thesis.

The therizinosaur skeleton was difficult to prepare due to the matrix and diagenetic crushing of the elements, but at the time it was collected it represented the first evidence of therizinosaurs from North America. Doug Wolfe and colleagues had not yet figured out that part of the frill of Zuniceratops was actually the ischium of what would later be named Nothronychus mckinleyi; and the Crystal Geyser Quarry, which would produce Falcarius, had not yet yielded any recognized therizinosaur material. What made the MNA fossil even more spectacular was its stratigraphic control. It was found in a marine deposit and the bones were actually covered with ammonites!

Shortly afterwards the MNA had some problems and at one point the Geology program was terminated. Fortunately due to public outcry (among other things) the department was reestablished; however, these circumstances caused delays in the study of this specimen. A couple of years ago; however, it was the focus of an excellent new exhibit at the MNA and now the description is finally out. As I have not yet seen the paper I cannot comment on the content but there are some early news reports up here and here. Congrats to Dave Gillette (and to Lindsay Zanno) on the release of the results of this long awaited study. I know that he put a ton of work into this specimen.



You can read an earlier account of this excavation and specimen that came out in Arizona Geology back in the summer of 2007. There is also a good photo gallery of Utah fossils here which includes a reconstruction of N. graffami by artist Victor Leshyk (see photo above), pictures of a display of some of the bones, as well as a picture of Dave in front of a full scale reconstruction done by Rob Gaston.

Dinosaur Diversity and the Rock Record

Barrett, P.M., McGowan, A.J., and V. Page. 2009. Dinosaur diversity and the rock record. Procedings of the Royal Society B published online 29 April 2009. doi: 10.1098/rspb.2009.0352

Abstract - Palaeobiodiversity analysis underpins macroevolutionary investigations, allowing identification of mass extinctions and adaptive radiations. However, recent large-scale studies on marine invertebrates indicate that geological factors play a central role in moulding the shape of diversity curves and imply that many features of such curves represent sampling artefacts, rather than genuine evolutionary events. In order to test whether similar biases affect diversity estimates for terrestrial taxa, we compiled genus-richness estimates for three Mesozoic dinosaur clades (Ornithischia, Sauropodomorpha and Theropoda). Linear models of expected genus richness were constructed for each clade, using the number of dinosaur-bearing formations available through time as a proxy for the amount of fossiliferous rock outcrop. Modelled diversity estimates were then compared with observed patterns. Strong statistically robust correlations demonstrate that almost all aspects of ornithischian and theropod diversity curves can be explained by geological megabiases, whereas the sauropodomorph record diverges from modelled predictions and may be a stronger contender for identifying evolutionary signals. In contrast to other recent studies, we identify a marked decline in dinosaur genus richness during the closing stages of the Cretaceous Period, indicating that the clade decreased in diversity for several million years prior to the final extinction of non-avian dinosaurs at the Cretaceous-Palaeocene boundary.

New Paper on the Triassic Dinosaur Staurikosaurus pricei

Bittencourt, J.S., and A.W.A. Kellner. 2009. The anatomy and phylogenetic position of the Triassic dinosaur Staurikosaurus pricei Colbert, 1970. Zootaxa 2079:1-56.

Abstract: We redescribe the holotype of the saurischian dinosaur Staurikosaurus pricei Colbert, 1970 from Late Triassic Santa Maria Formation (southern Brazil), following additional preparation that revealed new anatomical features. A revised diagnosis is proposed and the published synapomorphies for Dinosauria and less inclusive clades (e.g. Saurischia) are evaluated for this species. Some characters previously identified as present in the holotype, including the intramandibular joint, hyposphene-hypantrum articulations in dorsal vertebrae, and a cranial trochanter and trochanteric shelf on the femur, cannot be confirmed due to poor preservation or are absent in the available material. In addition, postcranial characters support a close relationship between S. pricei and Herrerasaurus ischigualastensis Reig, 1963 (Late Triassic, Argentina), forming the clade Herrerasauridae. Several pelvic and vertebral characters support the placement of S. pricei as a saurischian dinosaur. Within Saurischia, characters observed in the holotype, including the anatomy of the dentition and caudal vertebrae, support theropod affinities. However, the absence of some characters observed in the clades Theropoda and Sauropodomorpha suggests that S. pricei is not a member of Eusaurischia. Most morphological characters discussed in previous phylogenetic studies cannot be assessed for S. pricei because of the incompleteness of the holotype and only known specimen. The phylogenetic position of S. pricei is constrained by that of its sister taxon H. ischigualastensis, which is known from much more complete material.

New Long Necked Stegosaur Miragaia longicollum

Not Triassic but still really cool. This is a new advance paper in Proceedings of the Royal Society B on a new long necked stegosaur from the Late Jurassic of Portugal. Appropriately named Miragaia longicollum it has at least 17 cerival vertebrate due to "cervicalization" of the dorsal column. Very cool. Even cooler is that it is available for free from the Royal Society website. The photo below is from here.

REFERENCE

Mateus, O., Maidment, S.C.R., and N.A. Christiansen. 2009. A new long-necked ‘sauropod-mimic’ stegosaur and the evolution of the plated dinosaurs. Proceedings of the Royal Society B, Published online on February 25, 2009. DOI 10.1098/rspb.2008.1909.

What exactly is going on with the dinosaurs in the Early Jurassic?

It is considered by some to be, and should have been, a classic example of adaptive radiation. At the end of the Triassic the majority of pseudosuchians go extinct, removing the biggest competitors of the dinosaurs and leaving the door open for an evolutionary explosion of the dinosaurs. Interestingly, however, this is not what happened according to a new study by Steve Brusatte and colleagues (Brusatte et al., 2008b) who found, in the continuation of their research comparing morphospace disparity between ornithidirans and pseudosuchians (Brusatte et al., 2008a), that dinosaur disparity remained relatively unchanged through the Triassic/Jurassic boundary. It would be expected that once the extinction of the pseudosuchians freed up a large amount of morphospace, the dinosaur record (with whom the pseudosuchians occupied a lot of the same niches and had similar body plans) would show a strong response, yet the dinosaurs show only a "slight non-significant increase" (Brusatte et al., 2008b). Thus, these authors argue, "different aspects of dinosaur radiation (diversity, disparity, and abundance) were decoupled, and the overall macroevolutionary pattern of the first 50 Myr of dinosaur evolution is more complex than often considered (Brusatte et al., 2008b).

Adam Yates had discussed this (and his hypothesis) a few weeks ago at Dracovenator and I had provided some follow-up discussion here. Nonetheless, despite the timing of the extinction it is apparent that not to much is going on for the dinosaurs immediately after the TR/J extinction (which by the way took out the non-dinosaurian dinosauromorphs). Sure to the record of coelophysoids and sauropodomorphs you add a few large theropods such as Dilophosaurus and you see the first good records of heterodontosaurids and the earliest thyreophorans (including the first ornithischians and sauropodomorphs in N. America), but you do not see a true explosion of dinosaur diversity until you get to the Late Jurassic. How much of this is a sampling and/or preservation problem is unclear, but simply look at the Weishampel et al. (2004) chapter on dinosaur distribution in the 2nd edition of The Dinosauria and compare the faunal lists for these epochs. You really have to clean up the Late Triassic portion removing many of the Ornithischia references, indeterminate theropods (could be shuvosaurids), and all of the footprint evidence (no ornithischian or sauropodomorph tracks in N. America; the "theropod" tracks worldwide could be made by convergent dinosauriforms, and pseudosuchians), not to mention the really messed up stratigraphy for the Chinle and Dockum which caused some duplicate entries. Now compare the Late Triassic, Early Jurassic, and Middle Jurassic lists to the rest of the chapter. Surprised? I commend Brusatte et al. (2008a, 2008b) for setting the stage and providing a baseline framework for some much needed future research to address this enigma.

REFERENCES

Brusatte, S.L., Benton, M.J., Ruta, M., and G.T. Lloyd. 2008a. Superiority, competition, and opportunism in the evolutionary radiation of dinosaurs. Science 321:1485-1488.

Brusatte, S.L., Benton, M.J., Ruta, M., and G.T. Lloyd. 2008b. The first 50 Myr of dinosaur evolution: macroevolutionary pattern and morphological disparity. Biology Letters, doi:10.1098/rsbl.2008.0441, published online.

Weishampel, D. B., Barrett, P. M., Coria, R. E., Le Loeuff, J., Gomani, E. S., Zhao Z., Xu X., Sahni, A., and C. Noto. 2004. Dinosaur distribution. In: Weishampel, D. B., Dodson, P., and Osmólska, H. eds. The Dinosauria. 2nd edition. Univ. California Press, Berkeley. pp. 517-606.

Dinosauria vs. Pseudosuchia - New paper in Science

Very recently there has been a resurgence of interest in the early appearance and diversification of the Dinosauria mainly due to the recognition that there exists strong convergence between early dinosaurs and pseudosuchian archosaurs such as Revueltosaurus and Shuvosaurus, and that dinosaur precursors such as Dromomeron and Silesaurus not only survived into the Late Triassic but also coexisted with the dinosaurs for millions of years (Dzik, 2003; Ezcurra, 2006; Irmis et al. 2007b; Nesbitt et al., 2007; Parker et al., 2005; Nesbitt and Norell, 2006). This has been accompanied by studies demonstrating that in some faunas (especially those of North America) dinosaurs were neither dominant or diverse, and that in fact there is no unambiguous evidence of Triassic ornithischians or sauropodomorphs in North America, and that the global record of Triassic ornithischians is extremely poor (Irmis et al., 2007a; Nesbitt et al., 2007). These and other studies have also demonstrated that Late Triassic pseudosuchians were extremely diverse and that their occurrence together with ornithodirans in most Late Triassic assemblages demonstrates that they were filling similar ecological roles. Thus, one of the biggest mysteries is why the majority of pseudosuchian lineages die out at the end of the Triassic, while the more conservative dinosaurs go on to have great success for the next 140 million years.

Today in the new issue of Science, Brusatte et al. provide the results of a multifaceted study addressing this question. They provide a new phylogenetic analysis of the Archosauria (supplementary materials) and compare evolutionary rates and morphological disparity between pseudosuchians and ornithodirans. Interestingly they found that the dinosaurs had lower disparity and represented a lesser amount of morphospace occupation compared to the pseudosuchians. Furthermore rates of character evolution between the two groups were indistinguishable. Previous hypotheses that the dinosaurs were more successful due to physiological superiority and were “preordained for success” are discounted (as was also argued by Irmis et al., 2007b). Instead Brusatte et al., suggest that the “dinosaurs were the beneficiaries of two mass extinction events – and some good luck”.

I admit that I am not surprised at all by their findings, but am probably biased because this trend is readily apparent in North America (where I work) where with the exception of the Hayden and Coelophysis Quarries at Ghost Ranch New Mexico (and trackways in the youngest Triassic units) there is a marked paucity of Triassic dinosaur fossils and an abundance of diverse pseudosuchians. I am a bit flummoxed over the basal positioning of Revueltosaurus in their phylogeny, but this is based on an incomplete coding which I have not thoroughly reviewed.

Overall I find the paper to be a useful contribution in the attempt to discern why such a wonderful diversity of crocodile-line archosaurs lineages was extinguished at the end Triassic. Their data helps quantify some of the trends seen by other workers, especially that the competition model is most likely untenable. However, disproving the competition scenario does not necessarily support the "lucky break" hypothesis. Furthermore, I have not seen strong evidence for a Carnian-Norian terrestrial extinction in the fossil record, a claim that is even more weakened by the recent announcement of a Rhaetian dicynodont, which supports known Norian dicynodonts in Arizona and rhynchosaurs in Brazil and Argentina. Recent published and unpublished studies revising the Late Triassic timescale demonstrate that much of the hypothesized Carnian terrestrial strata worldwide is probably actually Norian, thus at best there are very few Carnian age terrestrial assemblages (e.g., Muttoni et al., 2004; Furin et al., 2004). There is still much work to be done on this mystery and I for one am not quite ready yet to simply attribute it a “lucky break”; however if this is the case then I truly rue what would appear to be a cruel twist of fate, and can only wonder what might have come to pass if the pendulum had swung the other way.

REFERENCES

Brusatte, S.L., Benton, M.J., Ruta, M., and G.T. Lloyd. 2008. Superiority, competition, and opportunism in the evolutionary radiation of dinosaurs. Science 321:1485-1488.

Dzik, J. A beaked herbivorous archosaurs with dinosaur affinities from the early Late Triassic of Poland. Journal of Vertebrate Paleontology 23:556-574.

Ezcurra, M.D. 2007. A review of the systematic position of the dinosauriform archosaur Eucoelophysis baldwini Sullivan & Lucas, 1999 from the Upper Triassic of New Mexico, USA. Geodiversitas 28:649-684.

Furin, S., Preto, N., Rigo, M., Roghi, G., Gianolla, P., Crowley, J.L., and S. A. Bowring. 2006. High-precision U-Pb zircon age from the Triassic of Italy: Implications for the Triassic time scale and the Carnian origin of calcareous nannoplankton and dinosaurs. Geology 34:1009-1012.

Irmis, R.B., Parker, W.G., Nesbitt, S.J., and J. Liu, 2007a. Early ornithischian dinosaurs: the Triassic Record. Historical Biology 19:3-22.

Irmis, R.B., Nesbitt, S.J., Padian, K., Smith, N.D., Turner, A.H., Woody, D., and A. Downs. 2007b. A Late Triassic dinosauromorph assemblage from New Mexico and the rise of dinosaurs. Science 317:358-361.

Muttoni, G., Kent, D. V., Olsen, P. E., DiStefano, P., Lowrie, W., Bernasconi, S. M., and F. M. Hernández. 2004. Tethyan magnetostratigraphy from Pizzo Mondello (Sicily) and correlation to the Late Triassic Newark astrochronological polarity timescale. Geological Society of America Bulletin 116:1043-1058.

Nesbitt, S.J, and M.A. Norell. 2006. Extreme convergence in the body plans of an
early suchian (Archosauria) and ornithomimid dinosaurs (Theropoda). Proceedings of the Royal Society of London Series B 273: 1045–1048.

Nesbitt, S.J., Irmis, R.B., and W.G. Parker, 2007. A critical reevaluation of the Late Triassic dinosaur taxa of North America. Journal of Systematic Palaeontology 5:209-243.

Parker, W.G., Irmis, R.B., Nesbitt, S.N., Martz, J. W., and L. S. Browne, 2005. The pseudosuchian Revueltosaurus callenderi and its implications for the diversity of early ornithischian dinosaurs. Proceedings of the Royal Society London B 272:963-969.