Tulerpeton
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''Tulerpeton'' is an extinct
genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...
of
Devonian The Devonian ( ) is a period (geology), geologic period and system (stratigraphy), system of the Paleozoic era (geology), era during the Phanerozoic eon (geology), eon, spanning 60.3 million years from the end of the preceding Silurian per ...
four-limbed vertebrate, known from a fossil that was found in the
Tula Region Tula Oblast () is a federal subject (an oblast) of Russia. It is geographically located in European Russia and is administratively part of the Central Federal District, covering an area of . It has a population of Tula is the largest city and t ...
of
Russia Russia, or the Russian Federation, is a country spanning Eastern Europe and North Asia. It is the list of countries and dependencies by area, largest country in the world, and extends across Time in Russia, eleven time zones, sharing Borders ...
at a site named Andreyevka. This genus, ''
Acanthostega ''Acanthostega'', from Ancient Greek ἄκανθα (''ákantha''), meaning "spine", and στέγη (''stégē''), meaning "roof", is an extinct genus of stem tetrapoda, stem-tetrapod, among the first vertebrates, vertebrate animals to have recogn ...
'', and ''
Ichthyostega ''Ichthyostega'', from Ancient Greek ἰχθύς (''ikthús''), meaning "fish", and στέγη (''stégē''), meaning "roof", is an Extinction, extinct genus of limbed tetrapodomorpha, tetrapodomorphs from the Devonian, Late Devonian of what is ...
'' represent the earliest
tetrapods A tetrapod (; from Ancient Greek τετρα- ''(tetra-)'' 'four' and πούς ''(poús)'' 'foot') is any four- limbed vertebrate animal of the clade Tetrapoda (). Tetrapods include all extant and extinct amphibians and amniotes, with the lat ...
.


Description

''Tulerpeton'' is considered one of the first "
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" (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 o ...
, 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'', from Ancient Greek ἄκανθα (''ákantha''), meaning "spine", and στέγη (''stégē''), meaning "roof", is an extinct genus of stem tetrapoda, stem-tetrapod, among the first vertebrates, vertebrate animals to have recogn ...
'' and ''
Ichthyostega ''Ichthyostega'', from Ancient Greek ἰχθύς (''ikthús''), meaning "fish", and στέγη (''stégē''), meaning "roof", is an Extinction, extinct genus of limbed tetrapodomorpha, tetrapodomorphs from the Devonian, Late Devonian of what is ...
'') 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 period (geology), geologic period and system (stratigraphy), system of the Paleozoic era (geology), era during the Phanerozoic eon (geology), eon, spanning 60.3 million years from the end of the preceding Silurian per ...
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 time scale, geologic period and System (stratigraphy), system of the Paleozoic era (geology), era that spans 60 million years, from the end of the Devonian Period Ma (million years ago) to the beginning of the ...
period that the first truly terrestrial pentadactyl
tetrapods A tetrapod (; from Ancient Greek τετρα- ''(tetra-)'' 'four' and πούς ''(poús)'' 'foot') is any four- limbed vertebrate animal of the clade Tetrapoda (). Tetrapods include all extant and extinct amphibians and amniotes, with the lat ...
– 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. The term is derived from the Greek word () meaning "finger." Sometimes the suffix "-dactylia" is used. The deri ...
. The polydactyl (more than five toes) condition of ''Tulerpeton'' caused considerable comment when the fossil was first discovered. Before the discovery, the pentadactyl five-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'', from Ancient Greek ἄκανθα (''ákantha''), meaning "spine", and στέγη (''stégē''), meaning "roof", is an extinct genus of stem tetrapoda, stem-tetrapod, among the first vertebrates, vertebrate animals to have recogn ...
'' and ''
Ichthyostega ''Ichthyostega'', from Ancient Greek ἰχθύς (''ikthús''), meaning "fish", and στέγη (''stégē''), meaning "roof", is an Extinction, extinct genus of limbed tetrapodomorpha, tetrapodomorphs from the Devonian, Late Devonian of what is ...
'' confirm that the pentadactyl 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 tetrapod vertebrate animals belonging to the clade Amniota, a large group that comprises the vast majority of living terrestrial animal, terrestrial and semiaquatic vertebrates. Amniotes evolution, evolved from amphibious Stem tet ...
s and all extinct tetrapods that were more closely related to amniotes than to
lissamphibia The Lissamphibia (from Greek λισσός (lissós, "smooth") + ἀμφίβια (amphíbia), meaning "smooth amphibians") is a group of tetrapods that includes all modern amphibians. Lissamphibians consist of three living groups: the Salientia ( ...
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 Lepospondyli is a diverse clade of early tetrapods. With the exception of one late-surviving lepospondyl from the Late Permian of Morocco ('' Diplocaulus minimus''), lepospondyls lived from the Visean stage of the Early Carboniferous to the Earl ...
,
seymouriamorphs Seymouriamorpha were a small but widespread group of limbed vertebrates (tetrapods). They have long been considered stem-amniotes (reptiliomorphs), and most paleontologists still accept this point of view, but some analyses suggest that seymouria ...
,
Embolomeri Embolomeri is an Order (biology), order of Tetrapod, tetrapods or Stem-group, stem-tetrapods, possibly members of Reptiliomorpha. Embolomeres first evolution, evolved in the Early Carboniferous (Mississippian age, Mississippian) Period and were th ...
,
temnospondyls Temnospondyli (from Greek τέμνειν, ''temnein'' 'to cut' and σπόνδυλος, ''spondylos'' 'vertebra') or temnospondyls is a diverse ancient order of small to giant tetrapods—often considered primitive amphibians—that flourished wo ...
, baphetids, colosteids and whatcheeriids were.
Michael Benton Michael James Benton (born 8 April 1956) is a British palaeontologist, and professor of vertebrate paleontology, vertebrate palaeontology in the School of Earth Sciences at the University of Bristol. His published work has mostly concentrated on ...
(2005) noted that, due to the fact that ''Tulerpeton'' had more than five 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 defi ...
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 Stegocephali Prehistoric lobe-finned fish genera Late Devonian sarcopterygians Devonian sarcopterygians of Europe Famennian genera