stem-amniote – Nix Illustration

Tyrannoroter

Tyrannoroter heberti lived in a lush tropical coal forest covering what is now Nova Scotia, Canada during the late Carboniferous, about 307 million years ago.

It was a member of the pantylid family in the recumbirostran lineage, a group of tetrapods whose evolutionary relationships are still a little uncertain. Traditionally they were classified as lepospondyl “amphibians”, but more recently some studies have found them to be either very early sauropsids or convergently reptile-like stem-amniotes.

Known only from a partial skull and jaw, based on the proportions of its pantylid relatives Tyrannoroter was probably around 30cm long (~1′) and would have resembled a squat lizard with a large blunt triangular head.

It would have had small bony scales within its skin, with interlocking polygonal “armor” on the underside of its jaw and chest, and irregular pebbly scales on other parts of its body. It may also have had claws on its toes, and potentially was capable of burrowing using a combination of its stout limbs and its shovel-like snout.

The roof of its mouth and the inside of its jaw were covered in extensive “batteries” of blunt teeth that show evidence of shearing and grinding motions – suggesting it may have been primarily processing tough plant matter, and making it one of the earliest known herbivorous tetrapods.

Anthracodromeus

Anthracodromeus longipes was an early reptile* that lived in what is now Ohio, USA, during the late Carbonifeorus about 307-305 million years ago.

(*or possibly a very reptile-like stem–amniote)

Around 20cm in total length (~8″), it had a rather lizard-like shape with a long body and a short tail. The digits on all four of its limbs were highly elongated with hooked claws, which appears to have been an adaptation for climbing.

It inhabited a coal forest dominated by lycopsid trees and seed ferns, and as one of the earliest known tetrapods to develop climbing behavior it would have had some ecological advantages over its relatives, being better able to escape from predators and access new food sources.

Araeoscelis

Araeoscelis gracilis was a superficially lizard-like animal that lived during the mid-Permian, around 275 million years ago, in what is now Texas, USA. About 60cm long (~2′), it had a slender body, proportionally long legs, and a solidly-built skull with strong teeth, suggesting that it was a fast runner that specialized in cracking open the carapaces of thick-shelled prey.

It was one of the last known members of a lineage known as araeoscelidians, which are usually considered to be very early members of the diapsid reptiles – but a recent study has proposed they might have even more ancient roots than that, possibly being a branch of stem–amniotes instead.

Termonerpeton

During the Early Carboniferous, around 330 million years ago, the region that is now the East Kirkton Quarry in Scotland was located close to the equator, with a lush tropical climate and volcanic hot springs dotting the landscape. It preserves fossils of some of the earliest known fully terrestrial tetrapods, and a recent discovery shows how some of these animals were already experimenting with the shapes of their feet to better get around on land.

Termonerpeton makrydactylus is only known from a partial skeleton, and shows a mix of anatomical features that make identifying its exact evolutionary relationships rather difficult – but it was probably a very early reptilomorph, closer related to amniotes than to lissamphibians. It may also have been very closely related to the equally enigmatic Eldeceeon and Silvanerpeton from the same region, but was almsot twice their size with a estimated total length of around 70cm (2’4″).

It would have resembled a rather heavily-built lizard-like or salamander-like animal, with fairly stumpy legs and probably lacking claws on its digits. While it would have had spindle-shaped scales on its underside, and possibly small rounded scales along its sides and back, these were bony structures embedded in its skin and probably weren’t very visible externally in life.

But Termonerpeton‘s most surprising feature was its proportionally large feet with especially elongated fourth toes, which would have helped to extend its stride length for energy-efficient terrestrial locomotion and to stabilize its movement on unstable surfaces – a much more “advanced” amniote-like arrangement than expected in such an early reptilomorph, and convergently similar to to the foot shapes of some modern lizards. Its fourth toe was also unusually chunky, suggesting it may even have been bearing most of its weight on just that one digit when walking.