Edopoids
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Edopoidea is a clade of primitive
temnospondyl Temnospondyli (from Greek τέμνειν, ''temnein'' 'to cut' and σπόνδυλος, ''spondylos'' 'vertebra') is a diverse order of small to giant tetrapods—often considered primitive amphibians—that flourished worldwide during the Carbo ...
amphibians including the genus ''
Edops ''Edops'' ('swollen face') is an extinct genus of temnospondyl amphibian from the Early Permian Period. Unlike more advanced temnospondyls of the time, such as ''Eryops'', ''Edops'' exhibited an archaic pattern of palatal bones, and still posses ...
'' and the family Cochleosauridae. Edopoids are known from the
Late Carboniferous Late may refer to: * LATE, an acronym which could stand for: ** Limbic-predominant age-related TDP-43 encephalopathy, a proposed form of dementia ** Local-authority trading enterprise, a New Zealand business law ** Local average treatment effect, ...
and Early Permian of North America and Europe, and the
Late Permian Late may refer to: * LATE, an acronym which could stand for: ** Limbic-predominant age-related TDP-43 encephalopathy, a proposed form of dementia ** Local-authority trading enterprise, a New Zealand business law ** Local average treatment effect, ...
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 that is absent in all other temnospondyls, and the lack of a
pineal foramen A parietal eye, also known as a third eye or pineal eye, is a part of the epithalamus present in some vertebrates. The eye is located at the top of the head, is photoreceptive and is associated with the pineal gland, regulating circadian rhyth ...
, a small hole on the skull roof of many early
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 t ...
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 mammal has ...
e,
maxilla The maxilla (plural: ''maxillae'' ) in vertebrates 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. T ...
e, and nasal bones 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 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'' 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 vertebra. The condyles are oval or reniform (kidney-shaped) in shape, and their anteri ...
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 animals with vertebrae or a notochord. It also tries to connect, by us ...
'', 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 (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
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 sk ...
''. 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 com ...
. Most recent phylogenetic analyses support a sister group 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 ''Dendrerpeton'' (from el, δένδρον , 'tree' and el, ἑρπετόν , 'creeping thing') is a genus of an extinct group of temnospondyl amphibians. Its fossils have been found primarily in the Joggins Formation of Eastern Canada and in ...
'', '' Balanerpeton'', and '' Capetus''. 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 ...
showing the relationships of edopoids from Sidor ''et al.'' (2006):


Paleobiology

Most edopoids are known from the
Late Carboniferous Late may refer to: * LATE, an acronym which could stand for: ** Limbic-predominant age-related TDP-43 encephalopathy, a proposed form of dementia ** Local-authority trading enterprise, a New Zealand business law ** Local average treatment effect, ...
and Early Permian of Europe and North America, which at the time formed a larger continent called Euramerica. '' 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 may refer to: * LATE, an acronym which could stand for: ** Limbic-predominant age-related TDP-43 encephalopathy, a proposed form of dementia ** Local-authority trading enterprise, a New Zealand business law ** Local average treatment effect, ...
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