Tetrapods (Greek tetrapoda = four feet) are vertebrate four-legged land animals. This kind of locomotion is called quadrupedal.

Amphibians, reptiles, dinosaurs (including their direct descendants, the birds), and mammals are all tetrapods. Even though snakes do not have limbs, they are tetrapods because they evolved from animals with four limbs.

Tetrapod

The earliest tetrapods evolved from the Sarcopterygii (lobe-finned fish) into air-breathing amphibians, perhaps in the Upper Devonian period. This means the transition took place in fish, before the land was the main habitat. This is typical of transitional fossils undergoing mosaic evolution.

Tetrapod

Evolution

Fishapods

Clear fossil tetrapod tracks from the mid-Devonian predate previous tetrapod records by 18 million years. These tracks are from the Middle Devonian of Poland, nearly 400 million years ago. The footprints were made in the mud of a tropical lagoon, and no animal of that time could have made the tracks except a tetrapod. The find strongly suggests that the animals were fish, not amphibia, when the transition to limbs occurred. The term fishapod is sometimes used. They would have been derived from lobe-finned fish (Sarcopterygii), but of a genus whose body fossil has not yet been found. The Polish team suggest the fish to tetrapod transition might have taken place as early as the Lower Devonian.

Tetrapod

The earliest tetrapods lived entirely in water. They could not live on land. Earlier, it was believed that fish had first moved onto land – either looking for food (like modern mudskippers) or to find water when the pond they lived in dried out. It was believed that they later evolved legs, lungs, and other body parts to live better on land.

Nine genera of Devonian tetrapods have been described. These earliest tetrapods were not terrestrial. They lived in swampy habitats like shallow wetlands, coastal lagoons, brackish river deltas, and even shallow marine sediments. There is much to suggest that these are the kind of environments in which the tetrapods evolved. Also, because fossils of early tetrapods are found widely in the Old Red Sandstone continent, they must have spread by following the coastal lines. This means they could not have lived only in freshwater.

Still, they may have spent brief periods out of water and would have used their legs to paw their way through the mud. The earliest confirmed terrestrial forms are known from the early Carboniferous deposits, some 20 million years later.

Tetrapods adapted to terrestrial environments over time and spent longer periods away from the water. They also spent more of their juvenile stage on land before returning to the water for the rest of their life. It is also possible that the adults started to spend some time on land to bask in the sun, close to the water's edge. The first true tetrapods that were adapted to moving on land were small. Only later did they increase in size.

Romer's gap

Between the lobe-finned fish tetrapods and the first amphibia and amniotes in the Middle Carboniferous lies a gap of 30 million years, with few satisfactory tetrapod fossils. This, noted in the early 1950s by Alfred Romer, is Romer's gap. Some new fossils were found in the 1990s, such as Pederpes, right in the middle of the Romer Gap. The gap still hides details of the transition from fish to tetrapod, but not so much as before.

So far, only two fossil sites from the Tournasian (earliest Carboniferous) are known. One is Horton Bluff Formation at Blue Beach, Nova Scotia. Much of this material has not yet been scientifically described.

The only other place a Tournasian tetrapod was found is near Dumbarton in the West of Scotland. The articulated (connected) skeleton of Pederpes was found there. Now comes news of new finds from this formation in five places in the Scottish Borders. The paper describes five new Tournaisian tetrapods, with information about their habitat. "Our analysis shows that the Tournaisian included a rich and diverse assemblage of taxa, which included close relatives of some Devonian forms on the tetrapod stem and basal members of the amphibian stem".

The authors say

"Although an extinction event at the end of the Devonian saw the demise of many archaic fish groups, our studies provide new perspectives on the recovery and diversification of surviving groups that went on to found the basis of modern vertebrate diversity".

Apparently, the split between the amphibians and the amniotes came soon after the extinction event as the tetrapods began to recover. "This accords with most molecular dates for the split, which place it at an average of 355 my ago, a date only 4 myr after the end-Devonian".