On the rise of the archosauromorphs

Proterosaurus speneri at Teyler’s Museum.

In the aftermath of the devastating Permo-Triassic mass extinction (~252 Ma), synapsid groups such as anomodonts and gorgonopsians and parareptiles such as pareiasaurs, were decimated and largely displaced by the archosauromorphs. The group, which include the ‘ruling reptiles’ (crocodylians, pterosaurs, dinosaurs, and their descendants, birds), originated during the middle–late Permian. The most basal archosauromorphs are Aenigmastropheus and Protorosaurus.

During the Triassic, the archosauromorphs achieved high morphological diversity, including aquatic or semi aquatic forms, highly specialized herbivores, massive predators, armoured crocodile-like forms, and gracile dinosaur precursors. The group constitutes an excellent empirical case to shed light on the recovery of terrestrial faunas after a mass extinction.

The Permian-Triassic boundary at Meishan, China (Photo: Shuzhong Shen)

The massive volcanic eruptions in Siberia at the end of the Permian, covered more than 2 millions of km 2 with lava flows, releasing more carbon in the atmosphere. High amounts of fluorine and chlorine increased the climatic instability, which means that the Mesozoic began under extreme hothouse conditions. Isotope studies and fossil record, indicates that temperatures in Pangaea interiors during the Early Triassic oscillated between 30 and 40 degrees Celsius, with heat peaks in the Induan and during the Early and Late Olenekian. It was suggested that during that time there was a moderate oxygen depletion that caused the low body size of the amphibian and reptilian life-forms found in those rocks.

After the mass extinction event, a distributed archosauromorph ‘disaster fauna’ dominated by proterosuchids, established for a short time. In South Africa, Proterosuchus occurs only between 5 and 14 m above the PT boundary and a similar pattern has been documented for the synapsid Lystrosaurus. During the Olenekian (1–5 million years after the extinction), archosauromorphs underwent a major phylogenetic diversification with the origins or initial diversification of major clades such as rhynchosaurs, archosaurs, erythrosuchids and tanystropheids.

Stenaulorhynchus stockleyi, a rhynchosaur from the Middle Triassic (From Wikimedia Commons)

The Mid Triassic is marked by the return of conifer-dominated forests, and the end of an interval of intense carbon perturbations, suggesting the recovery and stabilization of global ecosystems. The Anisian (5–10 Myr after the extinction) is characterized by a high diversity among the archosauromorphs with the appearance of large hypercarnivores, bizarre and highly specialized herbivores, long-necked marine predators, and gracile and agile dinosauromorphs. This phylogenetic diversity of archosauromorphs by the Middle Triassic paved the way for the ongoing diversification of the group (including the origins of dinosaurs, crocodylomorphs, and pterosaurs) in the Late Triassic, and their dominance of terrestrial ecosystems for the next 170 million years.




Ezcurra MD, Butler RJ. 2018 The rise of the ruling reptiles and ecosystem recovery from the Permo-Triassic mass
extinction. Proc. R. Soc. B 285: 20180361. http://dx.doi.org/10.1098/rspb.2018.0361

Ezcurra MD. (2016) The phylogenetic relationships of basal archosauromorphs, with an emphasis on the systematics of proterosuchian archosauriforms. PeerJ 4:e1778 https://doi.org/10.7717/peerj.1778

Holz, M., Mesozoic paleogeography and paleoclimates – a discussion of the diverse greenhouse and hothouse conditions of an alien world, Journal of South American Earth Sciences (2015), doi: 10.1016/j.jsames.2015.01.001


Introducing Shringasaurus indicus

Cranial anatomy of Shringasaurus indicus (From Sengupta et al., 2017)

In the aftermath of the Permo-Triassic mass extinction (~252 Ma), well diversified archosauromorph groups appear for the first time in the fossil record, including aquatic or semi aquatic forms, highly specialized herbivores, and massive predators. Allokotosaurians, meaning “strange reptiles” in Greek, comprise a bizarre suite of herbivorous archosauromorphs with a high disparity of craniodental features.

Shringasaurus indicus, from the early Middle Triassic of India, is a new representative of the Allokotosauria. The generic name is derived from ‘Śṛṅga’ (Shringa), horn (ancient Sanskrit), and ‘sauros’ (σαῦρος), lizard (ancient Greek), referring to the horned skull.  The species name ‘indicus’, refers to the country where it was discovered. The holotype ISIR (Indian Statistical Institute, Reptile, India) 780, consist of a partial skull roof (prefrontal, frontal, postfrontal, and parietal) with a pair of large supraorbital horns. The fossil bones have been collected from the Denwa Formation of the Satpura Gondwana Basin. At least seven individuals of different ontogenetic stages were excavated in the same area. Most of them were disarticulated, with exception of a partially articulated skeleton.

Skeletal anatomy of Shringasaurus indicus (From Sengupta et al., 2017)

Shringasaurus reached a relatively large size (3–4 m of total length) that distinctly exceeds the size range of other Early-Middle Triassic archosauromorphs. This new species shows convergences with sauropodomorph dinosaurs, including the shape of marginal teeth, and a relative long neck.  

Shringasaurus has a proportionally small skull with a short, rounded snout and confluent external nares. The premaxilla lacks a prenarial process and has four tooth positions. The prefrontal, nasal, frontal, and postfrontal of each side of the skull are fused to each other in large individuals. But the most striking feature of Shringasaurus indicus is the presence of a pair of large supraorbital horns, ornamented by tangential rugosities and grooves. Individuals of Shringasaurus of different ontogenetic stages indicate the size and robustness of the horns were exacerbated towards the adulthood, with a distinct variability in their orientation and anterior curvature in large individuals. Several amniotes have horns very similar to those of Shringasaurus (e.g. bovid mammals, chamaeleonid lepidosaurs). The independent evolution of similar horn shapes and robustness among different groups can be explained as the result of sexual selection.


Saradee Sengupta, Martín D. Ezcurra and Saswati Bandyopadhyay. 2017. A New Horned and Long-necked Herbivorous Stem-Archosaur from the Middle Triassic of India. Scientific Reports. 7, Article number: 8366. DOI: s41598-017-08658-8

Ezcurra MD. (2016The phylogenetic relationships of basal archosauromorphs, with an emphasis on the systematics of proterosuchian archosauriformsPeerJ 4:e1778 https://doi.org/10.7717/peerj.1778