Further Investigating the Biogeographic Origins of Dinosauria

New paper out in Palaeontology.

Marsola, J. C., Ferreira, G. S., Langer, M. C., Button, D. J., and R. J. Butler. 2018. Increases in sampling support the southern Gondwanan hypothesis for the origins of dinosaurs. Palaeontology. Early Online. https://doi.org/10.1111/pala.12411

Abstract - Dinosaurs were ubiquitous in terrestrial ecosystems through most of the Mesozoic and are still diversely represented in the modern fauna in the form of birds. Recent efforts to better understand the origins of the group have resulted in the discovery of many new species of early dinosaur and their closest relatives (dinosauromorphs). In addition, recent re‐examinations of early dinosaur phylogeny have highlighted uncertainties regarding the interrelationships of the main dinosaur lineages (Sauropodomorpha, Theropoda and Ornithischia), and questioned the traditional hypothesis that the group originated in South Gondwana and gradually dispersed over Pangaea. Here, we use an historical approach to examine the impact of new fossil discoveries and changing phylogenetic hypotheses on biogeographical scenarios for dinosaur origins over 20 years of research time, and analyse the results in the light of different fossil record sampling regimes. Our results consistently optimize South Gondwana as the ancestral area for Dinosauria, as well as for more inclusive clades including Dinosauromorpha, and show that this hypothesis is robust to increased taxonomic and geographic sampling and divergent phylogenetic results. Our results do not find any support for the recently proposed Laurasian origin of dinosaurs and suggest that a southern Gondwanan origin is by far the most plausible given our current knowledge of the diversity of early dinosaurs and non‐dinosaurian dinosauromorphs.

An interesting new study out today that uses phylogeny based biogeographical analyses to test the hypothesis of Baron et al. (2017a,b) that stem-dinosaurs originated in Laurasia. Their results reject that hypothesis in favor of the long standing hypothesis of a Gondwanan origin. I couldn't access the supplemental data in Dryad because the article isn't officially out. I'm not suggesting that the conclusions are wrong, but do have a few questions/comments about the data and how specimen sampling issues from Western North America could affect a rerun of the analysis. 

1) Some of the separation of taxa into biogeographical bins is confusing. For example most of the Chinle Formation taxa Chindesaurus, Tawa,  and Eucoelophysis are shown in green depicting the 'Equatorial Belt' as the ancestral zone, yet taxa from the same formation and localities therein such as Dromomeron gregorii and D. romeri are shown in yellow from the 'Euroamerica' zone. Why are they separated?

2) I am not aware of any Rhaetian occurrences of Eucoelophysis baldwini. This taxon occurs in a couple of quarries from around Ghost Ranch, New Mexico and one of them, the Hayden Quarry, is solidly dated in the middle-late Norian at about 212 Ma (Irmis et al., 2011). There is a purported occurrence (Rinehart et al., 2009) of Eucoelophysis from the Coelophysis Quarry at Ghost Ranch that is most likely Rhaetian in age; however, this occurrence is based on the rejected hypothesis that Eucoelophysis remains a neotheropod dinosaur with the holotype representing a highly weathered individual (Rinehart et al., 2009). This referred specimen is simply another specimen of Coelophysis. Thus all presently known occurrences of Eucoelophysis are Norian in age.

3) Recent fieldwork in the Chinle Formation, especially at Petrified Forest National Park, has recovered a significant amount of early dinosaur material. This includes the earliest known dated neotheropod specimen and early occurrences of dinosauriforms. This material is presently under study. Combined with already published accounts of silesaurids and coelophysids (e.g., Padian, 1986; Parker et al., 2006), these show a robust record of dinosauromorphs in the early-middle Norian of Arizona. Thus a specimen based study using autapomorphy-based identifications would pull Equatorial Laurasian silesaurids and neotheropods down into the early-middle Norian. Also important is a recently mentioned occurrence of Dromomeron gregorii from the Otischalkian Boren Quarry in the Dockum Group of Texas which is older than any of the Chinle Formation occurrences and pulls these occurrences down even further (Lessner et al., 2018).

4) Many of the specimens mentioned above are of interest because they originate from some of the lowest fossil bearing beds in the Chinle Formation, the Blue Mesa Member. To date no diagnostic vertebrate fossils from the Chinle Formation are known from below the middle of the Blue Mesa Member, thus the vertebrate faunas of the lower Chinle (lower Blue Mesa, Mesa Redondo, Shinarump) are unknown. This is significant because these units represent the earliest Norian based on the 'long-Norian' hypothesis (227-208 Ma). Coupled with the Moenkopi/Chinle unconformity much of the well-sampled Triassic of the western U.S. is apparently lacking the Carnian and earliest Norian. This is a bias that should not be ignored. Possible Carnian rocks elsewhere such as the base of the Dockum Group in Texas and units in Wyoming need to be better sampled.  

It's difficult to say how these details would affect these early dinosaur biogeographical studies, but there are data out there that should be included in future analyses.

REFERENCES

Baron, M. G., Norman, D. B. and Barrett, P. M. 2017a. A new hypothesis of dinosaur relationships and early dinosaur evolution. Nature, 543, 501–506.

Baron, M. G., Norman, D. B. and Barrett, P. M. 2017b. Baron et al. reply. Nature, 551, E4–E5.

Irmis, R. B., Mundil, R., Martz, J. W., and W. G. Parker. 2011. High resolution U-Pb ages from the Upper Triassic Chinle Formation (New Mexico, U.S.A.) support a diachronous rise of dinosaurs. Earth and Planetary Science Letters 309:258-267.

Lessner, E. J., Parker, W. G., Marsh, A. D., Nesbitt, S. J., Irmis, R. B. and B. Mueller. 2018. New insights into Late Triassic dinosauromorph-bearing assemblages from Texas using apomorphy-based identifications. PaleoBios, 35.ucmp_paleobios_39960.

Padian, K.. 1986. On the type material of Coelophysis Cope (Saurischia: Theropoda), and a new specimen from the Petrified Forest of Arizona (Late Triassic: Chinle Formation), p. 45-60. In K. Padian (ed.), The beginning of the Age of Dinosaurs: Faunal change across the Triassic-Jurassic boundary. Cambridge University Press, Cambridge.

Parker, W. G., Irmis, R. B., and S. J. Nesbitt. 2006. Review of the Late Triassic dinosaur record from Petrified Forest National Park, Arizona. Museum of Northern Arizona Bulletin 62:160-161.

Rinehart, L. F., Lucas, S. G., Heckert, A. B., Spielmann, J. A., and M. D. Celeskey. 2009. The paleobiology of Coelophysis bauri (Cope) from the Upper Triassic (Apachean) Whitaker Quarry, New Mexico, with detailed analysis of a single quarry block. New Mexico Museum of Natural History and Science Bulletin 45.