Elpistostegalia or Panderichthyida is an order of prehistoric

lobe-finned fishes 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 includ ...

which lived during the

Middle Devonian The Devonian ( ) is a geologic period and system of the Paleozoic era, spanning 60.3 million years from the end of the Silurian, million years ago (Mya), to the beginning of the Carboniferous, Mya. It is named after Devon, England, wher ...

Late Devonian The Devonian ( ) is a geologic period and system of the Paleozoic era, spanning 60.3 million years from the end of the Silurian, million years ago (Mya), to the beginning of the Carboniferous, Mya. It is named after Devon, England, wher ...

period Period may refer to: Common uses * Era, a length or span of time * Full stop (or period), a punctuation mark Arts, entertainment, and media * Period (music), a concept in musical composition * Periodic sentence (or rhetorical period), a concept ...

(about 385 to 374 million years ago). They represent the advanced

tetrapodomorph The Tetrapodomorpha (also known as Choanata) are a clade of vertebrates consisting of tetrapods (four-limbed vertebrates) and their closest sarcopterygian relatives that are more closely related to living tetrapods than to living lungfish. Advance ...

stock, the fishes more closely related to

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 than the

osteolepiform Osteolepiformes, also known as Osteolepidida, is a group of prehistoric lobe-finned fishes which first appeared during the Devonian period. The order contains the families Canowindridae, Megalichthyidae, Osteolepididae and Tristichopteridae. Th ...

fishes. The earliest elpistostegalians, combining fishlike and tetrapod-like characters, are sometimes called fishapods, a phrase coined for the advanced elpistostegalian ''

Tiktaalik ''Tiktaalik'' (; Inuktitut ) is a monospecific genus of extinct sarcopterygian (lobe-finned fish) from the Late Devonian Period, about 375 Mya (million years ago), having many features akin to those of tetrapods (four-legged animals). It may ha ...

''. Through a strict

cladistic Cladistics (; ) is an approach to biological classification in which organisms are categorized in groups (" clades") based on hypotheses of most recent common ancestry. The evidence for hypothesized relationships is typically shared derived char ...

view, the order includes the terrestrial tetrapods.

Palaeobiology of the elpistostegalians

A rise in global oxygen content allowed for the evolution of large, predatory fish that were able to exploit the shallow tidal areas and swamplands as top predators. Several groups evolved to fill these niches, the most successful were the elpistiostegalians. In such environments, they would have been challenged by periodic oxygen deficiency. In comparable modern aquatic environments like shallow

eutrophic Eutrophication is the process by which an entire body of water, or parts of it, becomes progressively enriched with minerals and nutrients, particularly nitrogen and phosphorus. It has also been defined as "nutrient-induced increase in phytoplank ...

lakes and swampland, modern

lungfish Lungfish are freshwater vertebrates belonging to the order Dipnoi. Lungfish are best known for retaining ancestral characteristics within the Osteichthyes, including the ability to breathe air, and ancestral structures within Sarcopterygii, i ...

and some genera of

catfish Catfish (or catfishes; order Siluriformes or Nematognathi) are a diverse group of ray-finned fish. Named for their prominent barbels, which resemble a cat's whiskers, catfish range in size and behavior from the three largest species alive, ...

also rely on the more stable, atmospheric source of oxygen. Being shallow-water fishes, the elpistostegalians evolved many of the basic adaptions that later allowed the tetrapods to become terrestrial animals. The most important ones were the shift of main propulsion apparatus from the tail fin to the pectoral and pelvic fins, and a shift to reliance on lungs rather than gills as the main means of obtaining oxygen. Both of these appear to be a direct result of moving to an inland freshwater mode of living. The elpistostegalians gave rise to the

s in the

Eifelian The Eifelian is the first of two faunal stages in the Middle Devonian Epoch. It lasted from 393.3 ± 1.2 million years ago to 387.7 ± 0.8 million years ago. It was preceded by the Emsian Stage and followed by the Givetian Stage. North American s ...

(early middle

Devonian The Devonian ( ) is a geologic period and system of the Paleozoic era, spanning 60.3 million years from the end of the Silurian, million years ago (Mya), to the beginning of the Carboniferous, Mya. It is named after Devon, England, whe ...

) around 395 million years ago. While the early tetrapods flourished and diversified over the next 30 million years, the non-tetrapod elpistostegalians disappear from the fossil record fairly quickly in the early

Frasnian The Frasnian is one of two faunal stages in the Late Devonian Period. It lasted from million years ago to million years ago. It was preceded by the Givetian Stage and followed by the Famennian Stage. Major reef-building was under way during th ...

around 380 million years ago, leaving the tetrapods the sole survivors of their line.

Traits

Paleontologist and professor Per E. Ahlberg has identified the following traits as synapomorphic for Elpistostegalia (and thus Tetrapoda): * The

endocranium The endocranium in comparative anatomy is a part of the skull base in vertebrates and it represents the basal, inner part of the cranium. The term is also applied to the outer layer of the dura mater in human anatomy. Structure Structurally, t ...

is hinged, the hinge forming the profundus nerve foramen. The cranial kinesis is also visible in the

skull roof The skull roof, or the roofing bones of the skull, are a set of bones covering the brain, eyes and nostrils in bony fishes and all land-living vertebrates. The bones are derived from dermal bone and are part of the dermatocranium. In comparati ...

, between the

parietal bone The parietal bones () are two bones in the Human skull, skull which, when joined at a fibrous joint, form the sides and roof of the Human skull, cranium. In humans, each bone is roughly quadrilateral in form, and has two surfaces, four borders, an ...

s and the

postparietal Postparietals are cranial bones present in fish and many tetrapods. Although initially a pair of bones, many lineages possess postparietals which were fused into a single bone. The postparietals were dermal bones situated along the midline of the ...

bones. * A rather small

shoulder girdle The shoulder girdle or pectoral girdle is the set of bones in the appendicular skeleton which connects to the arm on each side. In humans it consists of the clavicle and scapula; in those species with three bones in the shoulder, it consists of t ...

is present. * The anal and posterior dorsal fin supported by a basal plate and three unjointed radials. * The pectoral fin skeleton is composed of bones homologous to the tetrapod

humerus The humerus (; ) is a long bone in the arm that runs from the shoulder to the elbow. It connects the scapula and the two bones of the lower arm, the radius and ulna, and consists of three sections. The humeral upper extremity consists of a roun ...

ulna The ulna (''pl''. ulnae or ulnas) is a long bone found in the forearm that stretches from the elbow to the smallest finger, and when in anatomical position, is found on the medial side of the forearm. That is, the ulna is on the same side of t ...

, and

radius In classical geometry, a radius ( : radii) of a circle or sphere is any of the line segments from its center to its perimeter, and in more modern usage, it is also their length. The name comes from the latin ''radius'', meaning ray but also the ...

, followed by a host of smaller bones anchoring the fin rays; the pelvic fin skeleton similarly has

femur The femur (; ), or thigh bone, is the proximal bone of the hindlimb in tetrapod vertebrates. The head of the femur articulates with the acetabulum in the pelvic bone forming the hip joint, while the distal part of the femur articulates with ...

tibia The tibia (; ), also known as the shinbone or shankbone, is the larger, stronger, and anterior (frontal) of the two bones in the leg below the knee in vertebrates (the other being the fibula, behind and to the outside of the tibia); it connects ...

, and

fibula The fibula or calf bone is a leg bone on the lateral side of the tibia, to which it is connected above and below. It is the smaller of the two bones and, in proportion to its length, the most slender of all the long bones. Its upper extremity is ...

Phylogeny

The name, originally coined around the

genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...

Elpistostege ''Elpistostege'' is an extinct genus of finned tetrapodomorphs that lived during the late Givetian to early Frasnian ages of the Late Devonian epoch. Its only known species, ''E. watsoni'', was first described in 1938 by the British palaeontolog ...

'', later become a synonym for Panderichthyida. In most analyses, the group as traditionally imagined is actually an

evolutionary grade A grade is a taxon united by a level of morphological or physiological complexity. The term was coined by British biologist Julian Huxley, to contrast with clade, a strictly phylogenetic unit. Definition An evolutionary grade is a group of ...

, the last "fishes" of the tetrapod stem line, though Chang and Yu (1997) treated them as the sister

clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...

to Tetrapoda. Elpistostegalia was re-defined as a

containing ''

Panderichthys ''Panderichthys'' is a genus of extinct sarcopterygian (lobe-finned fish) from the late Devonian period, about 380 Mya. ''Panderichthys'', which was recovered from Frasnian (early Late Devonian) deposits in Latvia, is represented by two species. ...

'' and tetrapods. Below is a

cladogram A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to d ...

from Swartz, 2012.

References

{{Taxonbar, from=Q150821 Prehistoric lobe-finned fish Prehistoric fish orders Elpistostegalians Late Devonian animals Late Devonian fish Devonian bony fish