Polydactyly in stem-tetrapods should here be understood as having more than five digits to the finger or foot, a condition that was the natural state of affairs in the very first

stegocephalians Stegocephali (often spelled Stegocephalia) is a group containing all four-limbed vertebrates. It is equivalent to a broad definition of Tetrapoda: under this broad definition, the term "tetrapod" applies to any animal descended from the first ve ...

during the evolution of terrestriality. The polydactyly in these largely

aquatic animal An aquatic animal is any animal, whether invertebrate or vertebrate, that lives in water for most or all of its lifetime. Many insects such as mosquitoes, mayflies, dragonflies and caddisflies have aquatic larvae, with winged adults. Aquatic anim ...

s is not to be confused with

polydactyly Polydactyly or polydactylism (), also known as hyperdactyly, is an anomaly in humans and animals resulting in supernumerary fingers and/or toes. Polydactyly is the opposite of oligodactyly (fewer fingers or toes). Signs and symptoms In humans ...

in the medical sense, i.e. it was not an anomaly in the sense it was not a

congenital condition A birth defect, also known as a congenital disorder, is an abnormal condition that is present at birth regardless of its cause. Birth defects may result in disabilities that may be physical, intellectual, or developmental. The disabilities can r ...

of having more than the typical number of digits for a given

taxon In biology, a taxon (back-formation from ''taxonomy''; plural taxa) is a group of one or more populations of an organism or organisms seen by taxonomists to form a unit. Although neither is required, a taxon is usually known by a particular nam ...

. Rather, it appears to be a result of the early

evolution Evolution is change in the heritable characteristics of biological populations over successive generations. These characteristics are the expressions of genes, which are passed on from parent to offspring during reproduction. Variation ...

from a limb with a

fin A fin is a thin component or appendage attached to a larger body or structure. Fins typically function as foils that produce lift or thrust, or provide the ability to steer or stabilize motion while traveling in water, air, or other fluids. Fin ...

rather than digits. "Living tetrapods, such as the frogs, turtles, birds and mammals, are a subgroup of the tetrapod lineage. The lineage also includes finned and limbed tetrapods that are more closely related to living tetrapods than to living lungfishes."

Tetrapod Tetrapods (; ) are four-limbed vertebrate animals constituting the superclass Tetrapoda (). It includes extant and extinct amphibians, sauropsids ( reptiles, including dinosaurs and therefore birds) and synapsids (pelycosaurs, extinct theraps ...

s evolved from animals with fins such as found in

lobe-finned fish Sarcopterygii (; ) — sometimes considered synonymous with Crossopterygii () — is a taxon (traditionally a class or subclass) of the bony fishes known as the lobe-finned fishes. The group Tetrapoda, a mostly terrestrial superclass includi ...

es. From this condition a new pattern of limb formation evolved, where the development axis of the limb rotated to sprout secondary axes along the lower margin, giving rise to a variable number of very stout skeletal supports for a paddle-like foot. The condition is thought to have arisen from the loss of the

fin ray Fins are distinctive anatomical features composed of bony spines or rays protruding from the body of a fish. They are covered with skin and joined together either in a webbed fashion, as seen in most bony fish, or similar to a flipper, as see ...

-forming proteins actinodin 1 and actinodin 2 or modification of the expression of

HOXD13 Homeobox protein Hox-D13 is a protein that in humans is encoded by the ''HOXD13'' gene. This gene belongs to the homeobox family of genes. The homeobox genes encode a highly conserved family of transcription factors that play an important role in ...

. It is still unknown why exactly this happens. " SHH is produced by the mesenchymal cells of the zone of polarizing activity (ZPA) found at the posterior margin of the limbs of all vertebrates with paired appendages, including the most primitive chondrichthyian fishes. Its expression is driven by a well-conserved limb-specific enhancer called the ZRS (zone of polarizing region activity regulatory sequence) that is located approximately 1 Mb upstream of the coding sequence of ''Shh''." Early groups like ''

Acanthostega ''Acanthostega'' (meaning "spiny roof") is an extinct genus of stem-tetrapod, among the first vertebrate animals to have recognizable limbs. It appeared in the late Devonian period (Famennian age) about 365 million years ago, and was anatomic ...

'' had eight digits, while the more-derived ''

Ichthyostega ''Ichthyostega'' (from el, ἰχθῦς , 'fish' and el, στέγη , 'roof') is an extinct genus of limbed tetrapodomorphs from the Late Devonian of Greenland. It was among the earliest four-limbed vertebrates in the fossil record, and was on ...

'' had seven digits; the yet-more-derived ''

Tulerpeton ''Tulerpeton'' is an extinct genus of Devonian four-limbed vertebrate, known from a fossil that was found in the Tula Region of Russia at a site named Andreyevka. This genus and the closely related ''Acanthostega'' and ''Ichthyostega'' represent ...

'' had six toes. ''

Crassigyrinus ''Crassigyrinus'' (from la, crassus, 'thick' and el, γυρίνος el, gyrínos, 'tadpole') is an extinct genus of carnivorous stem tetrapod from the Early Carboniferous Limestone Coal Group of Scotland and possibly Greer, West Virginia. ...

'' from the fossil-poor

Romer's gap Romer's gap is an example of an apparent gap in the tetrapod fossil record used in the study of evolutionary biology. Such gaps represent periods from which excavators have not yet found relevant fossils. Romer's gap is named after paleontologist ...

in the early

Carboniferous The Carboniferous ( ) is a geologic period and system of the Paleozoic that spans 60 million years from the end of the Devonian Period million years ago ( Mya), to the beginning of the Permian Period, million years ago. The name ''Carbonifero ...

is usually thought to have had five digits to each foot. The anthracosaurs, which may be stem-tetrapods or reptiliomorphs, retained the five-toe pattern still found in

amniote Amniotes are a clade of tetrapod vertebrates that comprises sauropsids (including all reptiles and birds, and extinct parareptiles and non-avian dinosaurs) and synapsids (including pelycosaurs and therapsids such as mammals). They are disti ...

s, while further reduction had taken place on other

labyrinthodont "Labyrinthodontia" (Greek, 'maze-toothed') is an informal grouping of extinct predatory amphibians which were major components of ecosystems in the late Paleozoic and early Mesozoic eras (about 390 to 150 million years ago). Traditionally consid ...

lines, leaving the forefoot with four toes and the hind foot with five, a pattern still found in modern amphibians. The increasing knowledge of labyrinthodonts from Romer's gap has led to the challenging of the hypothesis that pentadactyly, as displayed by most modern tetrapods, is

plesiomorphic In phylogenetics, a plesiomorphy ("near form") and symplesiomorphy are synonyms for an ancestral character shared by all members of a clade, which does not distinguish the clade from other clades. Plesiomorphy, symplesiomorphy, apomorphy, and ...

. The number of digits was once thought to have been reduced in

amphibians Amphibians are four-limbed and ectothermic vertebrates of the class Amphibia. All living amphibians belong to the group Lissamphibia. They inhabit a wide variety of habitats, with most species living within terrestrial, fossorial, arbore ...

and

reptiles Reptiles, as most commonly defined are the animals in the Class (biology), class Reptilia ( ), a paraphyletic grouping comprising all sauropsid, sauropsids except birds. Living reptiles comprise turtles, crocodilians, Squamata, squamates (lizar ...

independently, but more recent studies suggest that a single reduction occurred, along the tetrapod stem, in the Early Carboniferous. However, even the early

ichthyostegalia Ichthyostegalia is an order of extinct amphibians, representing the earliest landliving vertebrates. The group is thus an evolutionary grade rather than a clade. While the group are recognized as having feet rather than fins, most, if not all, ...

ns like ''Acanthostega'' and ''Ichthyostega'' appear to have had the forward ossified bony toes combined in a single stout digit, making them effectively five-toed.

References

{{Commons category, Polydactyly_in_early_tetrapods Comparative anatomy Evolution of tetrapods Limbs (anatomy) Animal anatomy

See also

References