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.


Whatcheeria deltae here was an early tetrapod from the Early Carboniferous, about 340 million years ago, descended from the earlier fish-like forms and closely related to the ancestors of modern amphibians and amniotes.

Hundreds of fossils of this species have been found in Iowa, USA. Most represent juveniles, but rare larger specimens suggest fully-grown adults reached at least 2m long (6’6″).

Its large chunky limbs and flat feet seem to have been well-adapted for walking, with body proportions similar to later temnospondyl amphibians. But its cartilaginous ankles and the presence of lateral lines on its skull suggest it was still primarily aquatic, possibly walking along on the bottom of the ancient lakes, rivers, and swamps it inhabited.

It also had an unusually long neck and oddly-shaped skull for such an early tetrapod – most other known species had rather wide and flat skulls, but Whatcheeria‘s head was instead proportionally taller and narrower. Along with heavily reinforced sutures between the bones of its skull, it would have had a very powerful bite and been able to resist the twisting forces of large struggling prey in its jaws, suggesting it was a specialized crocodile-like predator.


Some of the earliest large terrestrial herbivores on Earth were the edaphosaurids – a very early-branching group of synapsids, the evolutionary lineage whose only modern surviving members are mammals. Like their more famous cousin Dimetrodon these animals sported huge elaborate sails on their backs formed from highly elongated vertebral spines, but despite the similarity in appearance they actually seem to have evolved these structures completely independently.

Known from a single partial skeleton discovered in southern New Mexico, USA, the edaphosaurid Gordodon kraineri dates to around the very end of the Carboniferous or the very earliest Permian, about 299 million years ago.

It was fairly small for an edaphosaurid at about 1.5m long (~5′), and seems to have had transitional anatomy between earlier and later members of the group. Its sail spines were thicker than those of earlier species but still less heavyset than those of later forms, and while each spine had numerous side projections these structures were small, thorn-like, and randomly distributed, unlike the more organized thick crossbars seen in Edaphosaurus.

Its head was proportionally small compared to its body, but still relatively large for an edaphosaurid, and it had an unusually long neck for an early synapsid. But its most distinctive features were its jaws and teeth – it had a narrow snout with a pair of large incisor-like teeth at the front of both its upper and lower jaws, followed by a large toothless gap (a diastema) and then a short row of small peg-like teeth. Like Edaphosaurus it also would have had batteries of grinding tooth plates inside its upper and lower jaws, but probably not as extensively.

Overall its tooth arrangment looked more like a modern herbivorous mammal than an early synapsid, much more highly specialized than anything else known to be alive at the time – the next synapsid known to convergently evolve similar teeth lived around 90 million years later!

It probably had a very different diet to its relatives, with its specialized teeth and fairly slender body suggesting it may have been a selective feeder, cropping the softer more nutritious parts of plants like the fleshy seeds and cones of gymnosperm plants.

Its discovery also hints that herbivorous edaphosaurids in general were much more diverse than we previously thought, and there may be even more surprising forms out there still to be discovered.


Named after a legendary Scandinavian serpent, Joermungandr bolti here was a recumbisrostran “microsaur” – part of a group of animals that were traditionally considered to be lepospondyl amphibians, but more recently have been proposed to in fact be a lineage of early reptiles.

Discovered in the Mazon Creek fossil beds in Illinois, USA, this species dates to the late Carboniferous period around 310 million years ago. A single near-complete specimen about 5cm long (~2″) preserves impressions of the body outline and numerous tiny scales, giving us a pretty good idea of what it looked like in life.

Joermungandr had a long streamlined tubular body with small limbs and a short tapering tail, and a stubby snout with fused bones heavily reinforcing its skull. Along with microscopic ridges on its body scales that resemble the dirt-repelling scales of some modern reptiles, this combination of features suggest it was a headfirst burrower that wriggled its way through soil with snakelike motions.


A group of early tetrapods known as diadectids were some of the first land animals to become specialized herbivores, developing adaptations for the bulk processing of tough fibrous vegetation. They had powerful jaws, chisel-like front teeth, and grinding cheek teeth, and they grew to relatively large sizes for their time with bulky bodies supporting voluminous plant-fermenting guts.

Although usually considered to be reptilomorphs – “amphibian-grade” animals more closely related to amniotes than to modern amphibians – some studies have instead placed these early plant-eaters as being true amniotes related to the synapsids. Fossil trackways show they may have had amniote-like claws on their feet, and that their highly flexible lizard-like ankle joints allowed them to walk much more efficiently than other early tetrapods, possibly using a semi-upright gait, but these may be convergently evolved features. Since we don’t know whether they laid amniote-like eggs or if they instead spawned amphibian-style in water, it’s currently hard to tell for certain just what they really were.

Diasparactus zenos (sometimes alternately known as Diadectes zenos) was a diadectid that lived during the early Permian in New Mexico, USA, about 296 million years ago. Around 1.3m long (4’3”), it was only about half the size of its largest relatives, but it’s notable for having unusually high neural spines on its vertebrae – not quite long enough to be considered a sail, but more of a “high back” that may have supported powerful musculature or fatty deposits.


Spathicephalus mirus here was part of a group of amphibian-like animals called the baphetoids, a lineage that weren’t quite true tetrapods themselves but were still very closely related to them.

Living in Scotland during the mid-Carboniferous period, about 326 million years ago, this 1.5m long (~5′) stem-tetrapod had an incredibly unusual head compared to its relatives – wide and flat, almost square in shape, with its jaws lined with hundreds of tiny chisel-like teeth.

Most other stem-tetrapods had deep skulls with large teeth, adapted for fish-eating, so clearly Spathicephalus was specialized for a very different diet. Some comparisons have been made to flat-headed ambush predator plagiosaurid temnospondyls like Gerrothorax, but a better ecological comparison might actually be filter-feeders like “pancake crocs“.


The lysorophians were a group of Carboniferous and Permian tetrapods with highly elongated bodies and tiny limbs, usually thought to be part of the lepospondyl amphibians – but some recent studies have instead placed them and some of their close relatives as possibly being very early members of the reptile branch of amniotes.

Brachydectes newberryi here is one of the best-known lysorophians, represented by a good amount of fossil material compared to many of its relatives. Living in the Midwestern United States during the late Carboniferous and early Permian, around 310-290 million years ago, it had a proportionally tiny head and reached lengths of around 60-70cm (2′-2’4″).

Its wide shovel-shaped snout and thickened reinforced bones around its braincase suggest it was adapted for headfirst digging, and some specimens have actually been found preserved inside their burrows. The roof of its skull also developed extensive “sculpturing” as individuals aged, with juveniles having smooth bone surfaces and larger adults having a distinct rough bumpy texture.

So I’ve depicted it here with a speculative keratinous “head shield”.


Ever since the earliest tetrapods crawled onto land and developed limbs and digits, some lineages have just… decided the whole “legs” thing was overrated and lost them entirely.

And the earliest known group to do this were the aïstopods. These highly elongated amphibian-like animals had specialized lightly-built skulls with large jaw muscles, and they may have filled a similar ecological niche to modern snakes, hunting small terrestrial invertebrates.

Lethiscus stocki was one of the first members of this snake-like group, living in Scotland during the Early Carboniferous about 340 million years ago. Growing to at least 50cm long (~20″), it was already a very specialized animal despite its basal position among the aïstopods, with eyes set far forward on its face and no trace of vestigial limbs.

CT scans of its skull have shown some surprisingly fish-like anatomy, especially in its braincase, features that were lost very early in tetrapod evolution. This suggests that aïstopods weren’t part of the lepospondyl amphibians like previously thought, but actually originated much much earlier in the tetrapod evolutionary tree – potentially placing them somewhere among the “fishapods” between Ichthyostega and Crassigyrinus.


Echinerpeton intermedium here was one of the earliest known members of the synapsids, the lineage that includes all mammals along with other “reptile-like” stem-mammals such as the famous sailbacked Dimetrodon.

Living during the Late Carboniferous in Nova Scotia, Canada, this 60-70cm long (2′-2’4″) distant cousin to modern mammals was previously known only from the fossilized remains of juveniles – with all known specimens showing slightly elongated spines on their vertebrae that gave it a sort of high-backed “proto-sail” appearance.

But a newly described fossil has completely changed what we know about this animal.

A single vertebrae identified as belonging to Echinerpeton shows a much much longer spine than anything we’ve ever seen before, and confirms that this species actually had a large elaborate true sailback – making it the earliest known tetrapod to experiment with this type of anatomy.

This individual seems to have been older than the other known specimens, but still not fully grown, leaving the possibility that fully mature Echinerpeton may have had even larger sails than this.

Temnospondyl Toes

The evolutionary origins of modern amphibians are still a bit murky, but one of the most likely possibilities is that they evolved from a group of temnospondyls known as amphibamiformes. (Or, at least, that frogs-and-salamanders evolved from them. Caecilians might be a different type of temnospondyl.)

And a recent discovery adds a little bit more evidence to that hypothesis.

A new specimen from the 309-million-year-old Late Carboniferous Mazon Creek fossil deposits in Illinois, USA, shows some soft-tissue impressions around the body of a terrestrial amphibamiform* — most notably showing its toes, with chunky rounded fleshy pads at the end like those seen in many modern amphibians.

Fossil trackways already suggested that some terrestrial temnospondyls had chunky toes, but up until now all known soft-tissue impressions only showed the slender tapering toes of aquatic forms. This is the first direct fossil evidence of toe pads, and hints that a lot of modern amphibians’ soft-tissue features may have actually had a very ancient origin.

(*A more precise identification couldn’t be made, but it shows some similarities to both Doleserpeton and Pasawioops.)