Temporal range: Mid Devonian to Recent
|Life restoration of Ichthyostega after Ahlberg, 2005.|
Tetrapods (Greek tetrapoda = four feet) are vertebrate land animals. The basic tetrapod plan is four legs and feet. This kind of locomotion is quadrupedal. Amphibians, reptiles, dinosaurs, birds, and mammals are all tetrapods. Even though snakes do not have limbs, they are tetrapods because they evolved from animals with four limbs.
The earliest tetrapods evolved from the Sarcopterygii, or 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.
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, dated to 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 becoming 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 > tetrapod transition might have taken place as early as the Lower Devonian.
Research by Jennifer Clack showed that the earliest tetrapods lived entirely in water. They could not live on land. Before this, 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.
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 1950, 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 obscures the details of the tetrapod transition.
The cleidoic egg[change]
Whereas amphibia lay their eggs in water, all other tetrapods (the amniotes) lay cleidoic eggs. These eggs are like private little ponds, protecting and nourishing the embryo until it grows into a hatchling. This was a key evolutionary 'invention', which allowed the amniotes to invade the land. Once the amniotes were truly land animals, there followed a huge adaptive radiation. This was one of the most significant advances in vertebrate evolution.
The Tetrapod clade[change]
The tetrapod clade separated from fish in the Upper Devonian. The amniotes were the earliest to lay cleidoic eggs. The Synapsids (> mammals) and the sauropsids (> reptiles) are sister clades, in particular, reptiles did not give rise to mammals.
- frogs and toads, newts and salamanders, and caecilians
- birds and modern reptiles
A partial taxonomy of the tetrapods:
- Phylum Chordata
- Class Sarcopterygii
- Superclass Tetrapoda
- Clack, Jennifer 2012. Gaining ground: the origin and early evolution of tetrapods. 2nd ed, Bloominton, Indiana: Indiana Umiversity Press. ISBN 978-0-35675-8
- Janvier, Phillipe and Clément, Gaël 2010. Muddy tetrapod origins. Nature 463, p40.
- Niedzwiedzki G. Szrek P. Narkiewitcz K & M and Ahlberg P.E. 2010. Nature 463, p43.
- formed by the joining of North America and northwestern Europe
- Romer A.S. 1949. The Vertebrate Body. Saunders, Philadelphia. 2nd ed. 1955; 3rd ed. 1962; 4th ed. 1970
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