Polydactyly in stem-tetrapods

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Limb evolution in Devonian tetrapodomorphs from pectoral fins to forelimbs. From left to right: Eusthenopteron, Panderichthys, Tiktaalik, Elpistostege, Acanthostega Fins to hands.png
Limb evolution in Devonian tetrapodomorphs from pectoral fins to forelimbs. From left to right: Eusthenopteron , Panderichthys , Tiktaalik , Elpistostege , Acanthostega

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 earliest stegocephalians during the evolution of terrestriality. The polydactyly in these largely aquatic animals is not to be confused with polydactyly in the medical sense, i.e. it was not an anomaly in the sense it was not a congenital condition of having more than the typical number of digits for a given taxon. [1] Rather, it appears to be a result of the early evolution from a limb with a 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." [2] Tetrapods evolved from animals with fins such as found in lobe-finned fishes. 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. [3] The condition is thought to have arisen from the loss of the fin ray-forming proteins actinodin 1 and actinodin 2 or modification of the expression of HOXD13. [4] [5] 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." [6]

Devonian taxa were polydactylous. Acanthostega had eight digits on both the hindlimbs and forelimbs. Ichthyostega , which was both more derived and more specialized, had seven digits on the hindlimb, though the hand is unknown. The yet-more-derived Tulerpeton had six toes on both the hindlimbs and forelimbs. [1]

It is unclear whether polydactylous tetrapods survived to the Carboniferous. Crassigyrinus , from the fossil-poor Romer's gap in the early Carboniferous, is usually thought to have had five digits to each foot. The anthracosaurs, which may be stem-tetrapods [7] [8] or reptiliomorphs, [9] retained the five-toe pattern still found in amniotes. Further reduction had taken place in the temnospondyls, leaving the forefoot with four toes and the hind foot with five, a pattern still found in modern amphibians. [10] 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. The number of digits was once thought to have been reduced in amphibians and reptiles independently, [1] [11] but more recent studies suggest that a single reduction occurred, along the tetrapod stem, in the Early Carboniferous. [12] [13] Even the early ichthyostegalians like Acanthostega and Ichthyostega appear to have had the forward ossified bony toes combined in a single stout digit, making them effectively five-toed.

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<span class="mw-page-title-main">Tetrapod</span> Superclass of the first four-limbed vertebrates and their descendants

A tetrapod is any four-limbed vertebrate animal of the superclass Tetrapoda. Tetrapods include all extant and extinct amphibians and amniotes, with the latter in turn evolving into two major clades, the sauropsids and synapsids. Some tetrapods, such as snakes, legless lizards, and caecilians, have evolved to become limbless via mutations of the Hox gene. Nevertheless, these limbless groups still qualify as tetrapods through their ancestry, and some retain a pair of vestigial spurs that are remnants of the hindlimbs.

<span class="mw-page-title-main">Dactyly</span> Arrangement of digits on hands and feet

In biology, dactyly is the arrangement of digits on the hands, feet, or sometimes wings of a tetrapod animal. The term is derived from the Greek word δακτυλος meaning "finger."

<i>Ichthyostega</i> Extinct genus of tetrapodomorphs

Ichthyostega is an extinct genus of limbed tetrapodomorphs from the Late Devonian of what is now Greenland. It was among the earliest four-limbed vertebrates ever in the fossil record and was one of the first with weight-bearing adaptations for terrestrial locomotion. Ichthyostega possessed lungs and limbs that helped it navigate through shallow water in swamps. Although Ichthyostega is often labelled a 'tetrapod' because of its limbs and fingers, it evolved long before true crown group tetrapods and could more accurately be referred to as a stegocephalian or stem tetrapod. Likewise, while undoubtedly of amphibian build and habit, it is not a true member of the group in the narrow sense, as the first modern amphibians appeared in the Triassic Period. Until finds of other early stegocephalians and closely related fishes in the late 20th century, Ichthyostega stood alone as a transitional fossil between fish and tetrapods, combining fish and tetrapod features. Newer research has shown that it had an unusual anatomy, functioning more akin to a seal than a salamander, as previously assumed.

<span class="mw-page-title-main">Jenny Clack</span> English paleontologist and evolutionary biologist (1947–2020)

Jennifer Alice Clack, was an English palaeontologist and evolutionary biologist. She specialised in the early evolution of tetrapods, specifically studying the "fish to tetrapod" transition: the origin, evolutionary development and radiation of early tetrapods and their relatives among the lobe-finned fishes. She is best known for her book Gaining Ground: the Origin and Early Evolution of Tetrapods, published in 2002 and written with the layperson in mind.

<span class="mw-page-title-main">Labyrinthodontia</span> Paraphyletic group of tetrapodomorphs

"Labyrinthodontia" is an informal grouping of extinct predatory amphibians which were major components of ecosystems in the late Paleozoic and early Mesozoic eras. Traditionally considered a subclass of the class Amphibia, modern classification systems recognize that labyrinthodonts are not a formal natural group (clade) exclusive of other tetrapods. Instead, they consistute an evolutionary grade, ancestral to living tetrapods such as lissamphibians and amniotes. "Labyrinthodont"-grade vertebrates evolved from lobe-finned fishes in the Devonian, though a formal boundary between fish and amphibian is difficult to define at this point in time.

<i>Acanthostega</i> Extinct genus of tetrapodomorphs

Acanthostega is an extinct genus of stem-tetrapod, among the first vertebrate animals to have recognizable limbs. It appeared in the late Devonian period about 365 million years ago, and was anatomically intermediate between lobe-finned fishes and those that were fully capable of coming onto land.

<i>Eusthenopteron</i> Extinct genus of tetrapodomorphs

Eusthenopteron is a genus of prehistoric sarcopterygian fish known from several species that lived during the Late Devonian period, about 385 million years ago. It has attained an iconic status from its close relationship to tetrapods. Early depictions of animals of this genus show them emerging onto land, but paleontologists now think that eusthenopteron species were strictly aquatic animals, though this is not completely known.

<span class="mw-page-title-main">Polydactyl cat</span> Cats with genetic anomaly that causes extra toes

A polydactyl cat is a cat with a congenital physical anomaly called polydactyly, which causes the cat to be born with more than the usual number of toes on one or more of its paws. Cats with this genetically inherited trait are most commonly found along the East Coast of North America and in South West England and Wales.

<i>Hynerpeton</i> Extinct genus of tetrapodomorphs

Hynerpeton is an extinct genus of early four-limbed vertebrate that lived in the rivers and ponds of Pennsylvania during the Late Devonian period, around 365 to 363 million years ago. The only known species of Hynerpeton is H. bassetti, named after the describer's grandfather, city planner Edward Bassett. Hynerpeton is known for being the first Devonian four-limbed vertebrate discovered in the United States, as well as possibly being one of the first to have lost internal (fish-like) gills.

<i>Tiktaalik</i> Extinct genus of tetrapodomorphs

Tiktaalik is a monospecific genus of extinct sarcopterygian from the Late Devonian Period, about 375 Mya, having many features akin to those of tetrapods. Tiktaalik is estimated to have had a total length of 1.25–2.75 metres (4.1–9.0 ft) based on various specimens.

<i>Crassigyrinus</i> Extinct genus of tetrapodomorphs

Crassigyrinus is an extinct genus of carnivorous stem tetrapod from the Early Carboniferous Limestone Coal Group of Scotland and possibly Greer, West Virginia.

<i>Ventastega</i> Extinct genus of tetrapodomorphs

Ventastega is an extinct genus of stem tetrapod that lived during the Upper Fammenian of the Late Devonian, approximately 372.2 to 358.9 million years ago. Only one species is known that belongs in the genus, Ventastega curonica, which was described in 1996 after fossils were discovered in 1933 and mistakenly associated with a fish called Polyplocodus wenjukovi. ‘Curonica’ in the species name refers to Curonia, the Latin name for Kurzeme, a region in western Latvia. Ventastega curonica was discovered in two localities in Latvia, and was the first stem tetrapod described in Latvia along with being only the 4th Devonian tetrapodomorph known at the time of description. Based on the morphology of both cranial and post-cranial elements discovered, Ventastega is more primitive than other Devonian tetrapodomorphs including Acanthostega and Ichthyostega, and helps further understanding of the fish-tetrapod transition.

<i>Tulerpeton</i> Extinct genus of tetrapodomorphs

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 the earliest tetrapods.

<span class="mw-page-title-main">Limb development</span>

Limb development in vertebrates is an area of active research in both developmental and evolutionary biology, with much of the latter work focused on the transition from fin to limb.

<span class="mw-page-title-main">Stegocephali</span> Clade of tetrapodomorphs

Stegocephali is a clade of vertebrate animals containing all fully limbed tetrapodomorphs. It is equivalent to a broad definition of the superclass Tetrapoda: under this broad definition, the term "tetrapod" applies to any animal descended from the first vertebrate with four limbs each with digits in the extremity (pentadactyly), rather than fins of their sarcopterygian relatives.

<span class="mw-page-title-main">Diplopodia</span>

Diplopodia is a congenital anomaly in tetrapods that involves duplication of elements of the foot on the hind limb. It comes from the Greek roots diplo = "double" and pod = "foot". Diplopodia is often found in conjunction with other structural abnormalities and can be lethal. It is more extreme than polydactyly, the presence of extra digits.

<span class="mw-page-title-main">Ichthyostegalia</span> Paraphyletic group of tetrapodomorphs

Ichthyostegalia is an order of extinct Tetrapodomorphs, 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, had internal gills in adulthood and lived primarily as shallow water fish and spent minimal time on land.

<span class="mw-page-title-main">Stem tetrapoda</span>

The Stem Tetrapoda are a cladistically defined group, consisting of all animals more closely related to extant four-legged vertebrates than to their closest extant relatives, but excluding the crown group Tetrapoda. They are thus paraphyletic, though acceptable in phylogenetic nomenclature as the group is defined by strict reference to phylogeny rather than to traits as in traditional systematics. Thus, some finned sarcopterygians are considered to be stem tetrapods.

<span class="mw-page-title-main">Evolution of tetrapods</span> Evolution of four legged vertebrates and their derivatives

The evolution of tetrapods began about 400 million years ago in the Devonian Period with the earliest tetrapods evolved from lobe-finned fishes. Tetrapods are categorized as animals in the biological superclass Tetrapoda, which includes all living and extinct amphibians, reptiles, birds, and mammals. While most species today are terrestrial, little evidence supports the idea that any of the earliest tetrapods could move about on land, as their limbs could not have held their midsections off the ground and the known trackways do not indicate they dragged their bellies around. Presumably, the tracks were made by animals walking along the bottoms of shallow bodies of water. The specific aquatic ancestors of the tetrapods, and the process by which land colonization occurred, remain unclear. They are areas of active research and debate among palaeontologists at present.


Innovations conventionally associated with terrestrially first appeared in aquatic elpistostegalians such as Panderichthys rhombolepis, Elpistostege watsoni, and Tiktaalik roseae. Phylogenetic analyses distribute the features that developed along the tetrapod stem and display a stepwise process of character acquisition, rather than abrupt. The complete transition occurred over a period of 30 million years beginning with the tetrapodomorph diversification in the Middle Devonian.

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