The Zachelmie trackways are a series of Middle Devonian-age trace fossils in Poland, purportedly the oldest evidence of terrestrial vertebrates (tetrapods) in the fossil record. These trackways were discovered in the Wojciechowice Formation, an Eifelian-age carbonate unit exposed in the Zachełmie Quarry of the Świętokrzyskie Mountains (Holy Cross Mountains). The discovery of these tracks has complicated the study of tetrapod evolution. Morphological studies suggest that four-limbed vertebrates ("tetrapods" in the broad sense, also known as stegocephalians) are descended from a specialized type of tetrapodomorph fish, the elpistostegalians. This hypothesis was supported further by the discovery and 2006 description of Tiktaalik, a well-preserved elpistostegalian from the Frasnian of Nunavut. Crucial to this idea is the assumption that tetrapods originated in the Late Devonian, after elpistostegalians appear in the fossil record near the start of the Frasnian. The Zachelmie trackways, however, appear to demonstrate that tetrapods were present prior to the Late Devonian. The implications of this find has led to several different perspectives on the sequence of events involved in tetrapod evolution.
Zachelmie trackways in the context of Tetrapod
A tetrapod (/ˈtɛtrəˌpɒd/; from Ancient Greek τετρα- (tetra-) 'four' and πούς (poús) 'foot') is any vertebrate animal of the clade Tetrapoda (/tɛˈtræpədə/). Tetrapods include all extant and extinct amphibians and amniotes, with the latter in turn evolving into two major clades, the sauropsids (reptiles, including dinosaurs and therefore birds) and synapsids (extinct "pelycosaurs", therapsids and all extant mammals, including humans). Hox gene mutations have resulted in some tetrapods becoming limbless (snakes, legless lizards, and caecilians) or two-limbed (cetaceans, sirenians, some lizards, kiwis, and the extinct moa and elephant birds). Nevertheless, they still qualify as tetrapods through their ancestry, and some retain a pair of vestigial spurs that are remnants of the hindlimbs.
Tetrapods evolved from a group of semiaquatic animals within the tetrapodomorphs which, in turn, evolved from ancient lobe-finned fish (sarcopterygians) around 390 million years ago in the Middle Devonian period. Early tetrapodomorphs were transitional between lobe-finned fishes and true four-limbed tetrapods, though most still fit the body plan expected of other lobe-finned fishes. The oldest fossils of four-limbed vertebrates (tetrapods in the broad sense of the word) are trackways from the Middle Devonian, and body fossils became common near the end of the Late Devonian, around 370–360 million years ago. These Devonian species all belonged to the tetrapod stem group, meaning that they did not belong to any modern tetrapod group.
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