Nundasuchus songeaensis, a New Archosauriform from the Middle Triassic of Tanzania and its Implications for the Phylogeny of the Archosauria

Nesbitt, S. J., Sidor, C. A., Angielczyk, K. D., Smith, R. M. H., and L. A. Tsuji. 2014. A new archosaur from the Manda beds (Anisian, Middle Triassic) of southern Tanzania and its implications for character state optimizations at Archosauria and Pseudosuchia. Journal of Vertebrate Paleontology 34:1357-1382 DOI:10.1080/02724634.2014.859622

Abstract - The presence of early pseudosuchians and avemetatarsalians in Anisian beds of Africa demonstrates that the archosaur radiation was well underway by the beginning of the Middle Triassic. The rapid radiation produced a variety of forms, but many of the unique, well-diagnosed
Late Triassic clades (e.g., Aetosauria, Ornithosuchidae) lack diagnosable members from the Middle Triassic. Here, we introduce a new Middle Triassic archosaur, Nundasuchus songeaensis, gen. et sp. nov., with an unusual mix of apomorphic character states found within Pseudosuchia and just outside Archosauria. The holotype consists of partial skeleton, including representative postcranial elements and parts of the skull. We added Nundasuchus songeaensis, gen. et sp. nov., into the two most comprehensive early archosaur phylogenetic data sets available, and in both analyses the new taxon falls within Pseudosuchia. However, a number of plesiomorphic archosaurian character states (e.g., posterolaterally directed tuber of the calcaneum) optimize as local autapomorphies of the new taxon within Pseudosuchia in our analyses. Therefore, we tested alternative hypotheses of relationships for the new taxon by utilizing constraint trees. The analyses resulted in little change in the relationships and structure of other Triassic archosaur clades, but changed optimizations of certain character states and character support at the base of Pseudosuchia and Archosauria. Our analyses suggest that the
complex evolution of character-state changes at the base of Archosauria is inhibiting our understanding of the relationships of early Pseudosuchia and, in turn, Archosauria.

Asperoris mnyama, a New Archisauriform from the Middle Triassic of Tanzania

Nesbitt, S. J., Butler, R. J., and D. J. Gower. 2013. A New Archosauriform (Reptilia: Diapsida) from the Manda Beds (Middle Triassic) of Southwestern Tanzania. PLoS ONE 8(9): e72753. doi:10.1371/journal.pone.0072753

Abstract

Background

Archosauria and their closest relatives, the non-archosaurian archosauriforms, diversified in the Early and Middle Triassic, soon after the end-Permian extinction. This diversification is poorly documented in most Lower and Middle Triassic rock sequences because fossils of early groups of archosauriforms are relatively rare compared to those of other amniotes. The early Middle Triassic (? late Anisian) Manda beds of southwestern Tanzania form an exception, with early archosaur skeletons being relatively common and preserved as articulated or associated specimens. The Manda archosaur assemblage is exceptionally diverse for the Middle Triassic. However, to date, no non-archosaurian archosauriforms have been reported from these rocks.

Methodology/Principal Findings

Here, we name a new taxon, Asperoris mnyama gen. et sp. nov., from the Manda beds and thoroughly describe the only known specimen. The specimen consists of a well-preserved partial skull including tooth-bearing elements (premaxilla, maxilla), the nasal, partial skull roof, and several incomplete elements. All skull elements are covered in an autapomorphic highly rugose sculpturing. A unique combination of character states indicates that A. mnyama lies just outside Archosauria as a stem archosaur within Archosauriformes, but more precise relationships of A. mnyama relative to other early archosauriform clades (e.g., Erythrosuchidae) cannot be determined currently.

Conclusions/Significance

Asperoris mnyama is the first confirmed non-archosaurian archosauriform from the Manda beds and increases the morphological and taxonomic diversity of early archosauriforms known from the Middle Triassic. The direct association of A. mnyama with species referable to Archosauria demonstrates that non-archosaurian archosauriforms were present during the rise and early diversification of Archosauria. Non-archosaurian archosauriforms and archosaurs co-occur in fossil reptile assemblages across Pangaea from the late Early Triassic to the end of the Late Triassic.

Rediscovered Specimen Draws Dinosaur Origins Down Into the Middle Triassic

I've been peripherally involved in the recent renaissance regarding dinosaur origins since my discovery of the skeleton of Revueltosaurus callenderi in 2004 and the subsequent recognition that it was not a dinosaur. With my good friends and colleagues Sterling Nesbitt and Randall Irmis, I proceeded to reexamine much of the Late Triassic dinosaur record using an apomorphy-based approach emphasized by UT Austin professor Chris Bell and his colleagues for Neogene vertebrates. Our findings were that Triassic dinosaurs were actually rarer than previously believed, especially in North America. In addition, extensive fieldwork by Sterling and Randy led to the discoveries of Dromomeron romeri, Tawa hallae, and Asilisaurus kongwe, filling in some key gaps in our understanding of the early diversification of the Ornithodira.  Nonetheless it has become readily apparent that many important specimens crucial to this issue were not weathering out of outcrops in the field, but rather were sitting un- or misidentified in museum collections around the globe. Our new understanding of character states and polarities for early diverging dinosauriforms provided us with specific search criteria leading to the discoveries and/or reinterpretations of taxa such as Eucoelophysis baldwini, Dromomeron gregorii, Technosaurus smalli, Daemonosaurus chauliodus, and of course Effigia okeeffae, all from previously collected material including fossils collected for Edward Cope in the 1800s. Furthermore, Randy's work found that the rise of dinosaurs was diachronous, although the timing is still poorly understood.

The discovery of the silesaurid Asilisaurus kongwe (published in 2010) pulled the split between Silesauridae and Dinosauria into the Middle Triassic creating a significant ghost lineage for Dinosauria as the earliest known bona fide dinosaurs do not appear until the end of the Carnian stage of the Late Triassic. Amazingly it appears that we did not need to wait very long for this ghost lineage to be filled.

Today's issue of Biology Letters has a paper by Sterling Nesbitt, Paul Barrett, Sarah Werning, Christian Sidor, and the late Alan Charig on a probable new dinosaur from the Middle Triassic of Tanzania. Even more amazing is that this is not a new specimen, but was actually collected in the 1930s, and never described, except in Charig's 1950s dissertation, until today. Charig named the new specimen Nyasasaurus parringtoni and until now this name has been a nomen nudum.

Known from a partial humerus and several vertebrate (three cervical, five presacral and three sacral), reanalysis places Nyasasaurus as either a dinosaur or as the sister taxon to Dinosauria. The humerus bears a ventrally elongated deltopectoral crest with a deflected apex, both synapomorphies of Dinosauria. The cervical vertebrae are elongate and possess deep lateral fossae, consistent with character states found in dinosaurs. The presence of three sacral vertebrae, although not restricted to Dinosauria, also supports this placement. This interpretation is supported not only by a phylogenetic analysis but also by bone histology, which shows high, continuous growth rates similar to that of early diverging dinosaurs. 

This material suggests that dinosaurs probably appeared in the fossil record 10 to 15 million years earlier than we expected. Furthermore, Nesbitt et al. argue that this strongly supports the hypothesis that dinosaurs were not a dominant group during their early history. Finally, Nyasasaurus also provides more support for a Gondwanan origin of dinosaurs.

One thing that is now definitely clear is that workers interested in dinosaur origins will need to spend more time in Middle Triassic terrestrial units. Back in 2004 I don't think any of us fathomed what discoveries and interpretations the next decade would bring. As I often state, it is not necessarily what we already know that dirves our work, but what is still out there for us to learn.

Nesbitt, S. J., Barrett, P. M., Werning, S., Sidor, C. A., and A. J. Charig. 2012. The oldest dinosaur? A Middle Triassic dinosauriform from Tanzania. Biology Letters.
 

Abstract - The rise of dinosaurs was a major event in vertebrate history, but the timing of the origin and early diversification of the group remain poorly constrained. Here, we describe Nyasasaurus parringtoni gen. et sp. nov., which is identified as either the earliest known member of, or the sister-taxon to, Dinosauria. Nyasasaurus possesses a unique combination of dinosaur character states and an elevated growth rate similar to that of definitive early dinosaurs. It demonstrates that the initial dinosaur radiation occurred over a longer timescale than previously thought (possibly 15 My earlier), and that dinosaurs and their immediate relatives are better understood as part of a larger Middle Triassic archosauriform radiation. The African provenance of Nyasasaurus supports a southern Pangaean origin for Dinosauria.

Redescription of the Archosaur Parringtonia gracilis from the Middle Triassic Manda Beds of Tanzania

Here is another redescription of another of the enigmatic archosauriforms from the Middle Triassic Manda Formation. This study confirms the affinity of Parringtonia gracilis and Erpetosuchus granti, and reinforces the importance of redescribing these older taxa using an apomorphy-based approach and placing them in a modern phylogenetic analysis. Unfortunately the exact position of Erpetosuchidae with Archosauria was not able to be precisely determined.

Nesbitt, S. J., and R. J. Butler. 2012. Redescription of the archosaur Parringtonia gracilis from the Middle Triassic Manda beds of Tanzania, and the antiquity of Erpetosuchidae. Geological Magazine (FirstView article), 14p. doi:10.1017/S0016756812000362 

Abstract - Parringtonia gracilis Huene, 1939 is represented by both cranial and postcranial material collected from the lower Middle Triassic (Anisian) Lifua Member of the Manda beds in southwestern Tanzania. This aberrant taxon was previously proposed to have affinities with pseudosuchian archosaurs, and specifically with the enigmatic Erpetosuchus granti from the Upper Triassic of Scotland. Here, we confirm the close affinities of Parringtonia gracilis and Erpetosuchus granti based on the following unambiguous synapomorphies: mediolaterally expanded posterior portion of the maxilla, alveoli present only in the anterior half of the maxilla, and absence of tooth serrations. Furthermore, the two taxa share osteoderms with deep sculpturing, a deep fossa on the dorsal margin of the neural spines and a heavily waisted shaft of the scapula. We added both Parringtonia gracilis and Erpetosuchus granti into a comprehensive phylogenetic analysis of early archosaurs and found that these taxa are clearly referable to Archosauria but that relationships are poorly resolved at the base of this clade. However, our analysis demonstrates that Erpetosuchus granti is not closely related to Crocodylomorpha, as has been hypothesized previously. The Erpetosuchidae are a clade of small-bodied archosaurs that have a poor fossil record but have members from both northern and southern Pangaea, ranging temporally from the Middle to Late Triassic. Thus, Erpetosuchidae is part of the early archosaurian radiation.