Tulerpeton
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''Tulerpeton'' is an extinct
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 ...
of
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 ...
four-limbed vertebrate, known from a fossil that was found in the
Tula Region Tula Oblast (russian: Ту́льская о́бласть, ''Tulskaya oblast'') is a federal subject (an oblast) of Russia. It is geographically in the European Russia region of the country and is part of the Central Federal District, covering a ...
of
Russia Russia (, , ), or the Russian Federation, is a List of transcontinental countries, transcontinental country spanning Eastern Europe and North Asia, Northern Asia. It is the List of countries and dependencies by area, largest country in the ...
at a site named Andreyevka. This genus and the closely related ''
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 ...
'' and ''
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 ...
'' represent the earliest
tetrapods 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 therapsids ...
.


Description

''Tulerpeton'' is considered one of the first "
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" (in the broad sense of the word) to have evolved. It is known from a fragmented skull, the left side of the
pectoral 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 ...
, and the entire right forelimb and right hindlimb along with a few belly scales. This species is differentiated from the less derived "aquatic tetrapods" (such as ''
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 ...
'' and ''
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 ...
'') by a strengthened limb structure. These limbs consist of 6 toes and fingers. Additionally, its limbs appear to have evolved for powerful paddling rather than walking. The fossil fragments also indicate that its head was disconnected from the pectoral girdle. From the absence of the rough postbranchial lamina of the pectoral girdle, it has been determined that ''Tulerpeton'' had no gills and was therefore entirely dependent on breathing air.


Era

''Tulerpeton'' lived approximately 365 million years ago, in the 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, whe ...
period when the climate was fairly warm and there were no glaciers. Land had already been colonized by plants. But it was only during the following
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 ...
period that the first truly terrestrial pentadactyl
tetrapods 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 therapsids ...
– the ancestors of present-day lissamphibians, mammals, birds, and reptiles – first began to appear.


Lifestyle

Even though ''Tulerpeton'' breathed air, it lived mainly in shallow marine water. The Andreyevka fossil bed where it was discovered was at least 200 km from the nearest landmass during this era. The fossils of plants in the area tell us that the salinity of the waters where it lived fluctuated wildly, indicating that the waters were quite shallow. Because the bones of the neck and the pectoral girdle were disconnected, ''Tulerpeton'' could lift its head. Therefore, in shallow water, it had a considerable advantage over the other animals whose heads only moved side to side. The later land animals that descended from ''Tulerpeton''’s relatives needed this head flexion on land, but the condition probably evolved because of the advantage that this gave it in shallow marine waters, not for land. In the book “Vertebrate Life”, authors Pough, Janis, and Heiser say that,” The development of a distinct neck, with the loss of the opercular bones and the later gain of a specialized articulation between the skull and the vertebral column (not yet present in the earliest tetrapods), may be related to lifting the snout out of the water to air or to snap at prey items.” The six fingered hands and toes were stronger than the fins from which they developed, therefore ''Tulerpeton'' had an advantage in propelling itself through shallow and brackish water, but the limbs do not yet seem strong enough for extensive use on land.


Significance of the find

''Tulerpeton'' is one of the early transition tetrapods – a marine animal capable of living on land. The separation of the pectoral-shoulder girdle from the head allowed the head to move up and down, and the strengthening of the legs and arms allowed the early tetrapods to propel themselves on land. ''Tulerpeton'' is important in the study of
dactyly In biology, dactyly is the arrangement of digits (fingers and toes) on the hands, feet, or sometimes wings of a tetrapod animal. It comes from the Greek word δακτυλος (''dáktylos'') = "finger". Sometimes the ending "-dactylia" is used ...
. The polydactyl (more than 5 toes) condition of ''Tulerpeton'' caused considerable comment when the fossil was first discovered. Before the discovery, the pentyldactyl, the 5-fingered condition that is ancestral to all living tetrapods, was thought to have developed before the first terrestrial tetrapods appeared. But the discoveries of ''
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 ...
'' and ''
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 ...
'' confirm that the pentyldactyl ancestor came later in the development of tetrapods. Phylogenetic studies conducted by Lebedev and Coates (1995) and Coates (1996) indicated that ''Tulerpeton'' was the earliest and basalmost member of the clade containing
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 and all extinct tetrapods that were more closely related to amniotes than to
lissamphibia The Lissamphibia is a group of tetrapods that includes all modern amphibians. Lissamphibians consist of three living groups: the Salientia (frogs, toads, and their extinct relatives), the Caudata (salamanders, newts, and their extinct relatives), ...
ns (living amphibians). However, later phylogenetic studies recovered ''Tulerpeton'' outside the least inclusive clade containing amniotes and lissamphibians, finding it to be more distantly related to amniotes than such extinct tetrapods as lepospondyls,
seymouriamorphs Seymouriamorpha were a small but widespread group of limbed vertebrates (tetrapods). They have long been considered reptiliomorphs, and most paleontologists may still accept this point of view, but some analyses suggest that seymouriamorphs are s ...
,
Embolomeri Embolomeri is an order of tetrapods or stem-tetrapods, possibly members of Reptiliomorpha. Embolomeres first evolved in the Early Carboniferous ( Mississippian) Period and were the largest and most successful predatory tetrapods of the Late Carb ...
,
temnospondyls 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 ...
, baphetids, colosteids and whatcheeriids were.
Michael Benton Michael James Benton One or more of the preceding sentences incorporates text from the royalsociety.org website where: (born 8 April 1956) is a British palaeontologist, and professor of vertebrate palaeontology in the School of Earth Sciences ...
(2005) noted that, due to the fact that ''Tulerpeton'' had more than 5 toes, establishing its exact phylogenetic position is important to our understanding of digital reduction in tetrapods. If it indeed was more closely related to amniotes than lissamphibians are, it would mean that "reduction to five digits had to happen twice, once on the line to amphibians and once on the
reptiliomorph Reptiliomorpha (meaning reptile-shaped; in PhyloCode known as ''Pan-Amniota'') is a clade containing the amniotes and those tetrapods that share a more recent common ancestor with amniotes than with living amphibians ( lissamphibians). It was de ...
line"; on the other hand, if it lies outside the least inclusive clade containing amniotes and lissamphibians, then "digital reduction happened once, between ''Tulerpeton'' and later tetrapods".Michael J. Benton (2005), ''Vertebrate Paleontology'', 3rd ed. Blackwell Science Ltd 2005, p. 80.


Notes


References


The Devonian Times reports on TulerpetonGordon, M.S. and Long, J.A. (2004) The Greatest Step In Vertebrate History: A Paleobiological Review of the Fish-Tetrapod Transition. Physiological and Biochemical Zoology. 77(5):700-719.The postcranial skeleton of the Devonian tetrapod Tulerpeton curtum Lebedev Stegocephalians: Tetrapods and other digit-bearing vertebrates Famennian Tetrapods Worldwide Tulerpeton
{{Taxonbar, from=Q133989 Late Devonian animals Basal tetrapods of Europe Ichthyostegalia