Edopoids
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Edopoidea is a
clade In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...
of primitive
temnospondyl Temnospondyli (from Greek language, Greek τέμνειν, ''temnein'' 'to cut' and σπόνδυλος, ''spondylos'' 'vertebra') or temnospondyls is a diverse ancient order (biology), order of small to giant tetrapods—often considered Labyrinth ...
amphibians including the genus ''
Edops ''Edops'' ('swollen face') is an extinct genus of temnospondyl amphibian from the Late Carboniferous - Early Permian periods. Unlike more advanced temnospondyls of the time, such as ''Eryops'', ''Edops'' exhibited an archaic pattern of palatal bo ...
'' and the family
Cochleosauridae Cochleosauridae is a family of edopoid temnospondyl amphibians, among the most basal of temnospondyls. Most members of this family are known from the late Carboniferous (Pennsylvanian) and early Permian (Cisuralian) of Europe and North America, t ...
. Edopoids are known from the
Late Carboniferous Late or LATE may refer to: Everyday usage * Tardy, or late, not being on time * Late (or the late) may refer to a person who is dead Music * Late (The 77s album), ''Late'' (The 77s album), 2000 * Late (Alvin Batiste album), 1993 * Late!, a pseudo ...
and
Early Permian 01 or 01 may refer to: * The year 2001, or any year ending with 01 * The month of January * 1 (number) Music * '01 (Richard Müller album), ''01'' (Richard Müller album), 2001 * 01 (Urban Zakapa album), ''01'' (Urban Zakapa album), 2011 * ''01011 ...
of North America and Europe, and the
Late Permian Late or LATE may refer to: Everyday usage * Tardy, or late, not being on time * Late (or the late) may refer to a person who is dead Music * ''Late'' (The 77s album), 2000 * Late (Alvin Batiste album), 1993 * Late!, a pseudonym used by Dave Groh ...
of Africa. They are among the most basal temnospondyls, and possess a number of primitive features that were lost in later members of the group.


Description

Edopoids are relatively large temnospondyls, with many species estimated to have grown several meters in length. The skull of ''Edops'' is broad while those of cochleosaurids are narrower and elongated. Distinguishing features of edopoids include the presence of an
intertemporal bone The intertemporal bone is a paired Skull, cranial bone present in certain Sarcopterygii, sarcopterygians (lobe-finned fish) and extinct amphibian-Evolutionary grade, grade tetrapods. It lies in the rear part of the skull, behind the eyes. Many lin ...
that is absent in all other temnospondyls, and the lack of a
pineal foramen A parietal eye (third eye, pineal eye) is a part of the epithalamus in some vertebrates. The eye is at the top of the head; is photoreceptive; and is associated with the pineal gland, which regulates circadian rhythmicity and hormone production ...
, a small hole on 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, including land-living vertebrates. The bones are derived from dermal bone and are part of the dermatocranium. In com ...
of many early
tetrapod A tetrapod (; from Ancient Greek :wiktionary:τετρα-#Ancient Greek, τετρα- ''(tetra-)'' 'four' and :wiktionary:πούς#Ancient Greek, πούς ''(poús)'' 'foot') is any four-Limb (anatomy), limbed vertebrate animal of the clade Tetr ...
s (young individuals still possess this hole). Relative to other temnospondyls, edopoids also have enlarged
premaxilla The premaxilla (or praemaxilla) is one of a pair of small cranial bones at the very tip of the upper jaw of many animals, usually, but not always, bearing teeth. In humans, they are fused with the maxilla. The "premaxilla" of therian mammals h ...
e,
maxilla In vertebrates, the maxilla (: maxillae ) is the upper fixed (not fixed in Neopterygii) bone of the jaw formed from the fusion of two maxillary bones. In humans, the upper jaw includes the hard palate in the front of the mouth. The two maxil ...
e, and
nasal bone The nasal bones are two small oblong bones, varying in size and form in different individuals; they are placed side by side at the middle and upper part of the face and by their junction, form the bridge of the upper one third of the nose. Eac ...
s in the snout region, which constrict the nostrils to small holes and push them to the sides of the skull. Most edopoids lacked grooves in the skull called sensory sulci, which presumably supported a
lateral line The lateral line, also called the lateral line organ (LLO), is a system of sensory organs found in fish, used to detect movement, vibration, and pressure gradients in the surrounding water. The sensory ability is achieved via modified epithelia ...
system in other temnospondyls. The lack of sensory sulci suggests that most edopoids were adapted to terrestrial lifestyles, as lateral lines are characteristic of aquatic animals. ''
Nigerpeton ''Nigerpeton'' (''Niger'', for the country, and ''herpeton'' (Greek), meaning crawler)Sidor, C.A., O’Keefe, F.R., Damiani, R., Steyer, J.S., Smith, R.M.H., Larsson, H.C.E., Sereno, P.C., Ide, O., Maga, A., 2005. Permian tetrapods from the Sahara ...
'' is the only edopoid to possess sensory sulci, but only in its adult form. The skulls of edopoids have only one
occipital condyle The occipital condyles are undersurface protuberances of the occipital bone in vertebrates, which function in articulation with the superior facets of the Atlas (anatomy), atlas vertebra. The condyles are oval or reniform (kidney-shaped) in shape ...
connecting them to the vertebrae of the neck, whereas more derived temnospondyls have two occipital condyles.


Classification

Edopoidea was named as a superfamily of temnospondyls by American paleontologist
Alfred Romer Alfred Sherwood Romer (December 28, 1894 – November 5, 1973) was an American paleontologist and biologist and a specialist in vertebrate evolution. Biography Alfred Romer was born in White Plains, New York, the son of Harry Houston Romer an ...
in the second edition of his textbook ''
Vertebrate Paleontology Vertebrate paleontology is the subfield of paleontology that seeks to discover, through the study of fossilized remains, the behavior, reproduction and appearance of extinct vertebrates (animals with vertebrae and their descendants). It also t ...
'', published in 1945. He recognized a close relationship between the families Edopidae (which includes only ''Edops'') and Cochleosauridae. In the late 1980s and early 1990s, the relationship between these two groups was supported by many
phylogenetic In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as phylogenetic inference. It infers the relationship among organisms based on empirical dat ...
analyses. One phylogenetic analysis separated ''Edops'' and cochleosaurids, finding the cochleosaurids to group with more derived temnospondyls like ''
Archegosaurus ''Archegosaurus'' is a genus of temnospondyl amphibian which lived during the Asselian to Wuchiapingian stages of the Permian, around 299-253 million years ago. The remains of this animal, consisting of at least 90 partial skeletons (mostly skull ...
''. However, the skull characteristics used in this analysis are common to almost all temnospondyls with elongated skulls, and are not strong evidence of evolutionary relationships because long snouts could easily have appeared through
evolutionary convergence Convergent evolution is the independent evolution of similar features in species of different periods or epochs in time. Convergent evolution creates analogous structures that have similar form or function but were not present in the last comm ...
. Most recent phylogenetic analyses support a
sister group In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and ...
relationship between ''Edops'' and Cochleosauridae, meaning that they are each other's closest relatives. Edopoids are usually placed at the base of Temnospondyli along with other primitive forms like '' Dendrerpeton'', '' Balanerpeton'', and '' Capetus''. Below is a
cladogram A cladogram (from Greek language, Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an Phylogenetic tree, evolutionary tree because it does not s ...
showing the relationships of edopoids from Sidor ''et al.'' (2006):


Paleobiology

Most edopoids are known from the
Late Carboniferous Late or LATE may refer to: Everyday usage * Tardy, or late, not being on time * Late (or the late) may refer to a person who is dead Music * Late (The 77s album), ''Late'' (The 77s album), 2000 * Late (Alvin Batiste album), 1993 * Late!, a pseudo ...
and
Early Permian 01 or 01 may refer to: * The year 2001, or any year ending with 01 * The month of January * 1 (number) Music * '01 (Richard Müller album), ''01'' (Richard Müller album), 2001 * 01 (Urban Zakapa album), ''01'' (Urban Zakapa album), 2011 * ''01011 ...
of Europe and North America, which at the time formed a larger continent called
Euramerica Laurasia () was the more northern of two large landmasses that formed part of the Pangaea supercontinent from around ( Mya), the other being Gondwana. It separated from Gondwana (beginning in the late Triassic period) during the breakup of Pa ...
. '' Procochleosaurus'', the oldest edopoid, is known from Ireland, while ''Edops'', the most basal edopoid, is known from the United States, strongly suggesting that the group originated in Euramerica. Tropical and subtropical environments were likely widespread across Euramerica during the Carboniferous and Early Permian, meaning that edopoids could easily travel between what are now North America and Europe. The edopoid ''Nigerpeton'' is known from the
Late Permian Late or LATE may refer to: Everyday usage * Tardy, or late, not being on time * Late (or the late) may refer to a person who is dead Music * ''Late'' (The 77s album), 2000 * Late (Alvin Batiste album), 1993 * Late!, a pseudonym used by Dave Groh ...
of Africa, extending the time span of edopoids by about 40 million years and expanding their geographic range outside Euramerica. It lived in a mountainous tropical region near the equator that is thought to have been a refugium for temnospondyls during the end of the Permian. At this time, the equatorial region was likely bounded by deserts to the north and south, which were too arid for amphibious animals like edopoids.


References

{{Taxonbar, from=Q5340197 Carboniferous temnospondyls Permian temnospondyls Prehistoric amphibians of Africa Prehistoric amphibians of Europe Prehistoric amphibians of North America Pennsylvanian first appearances Lopingian extinctions Taxa named by Alfred Romer Vertebrate superfamilies