 = Temnospondyli =

 Temnospondyli ( from Greek τέμνειν ( temnein , " to cut " ) and σπόνδυλος ( spondylos , " vertebra " ) ) is a diverse subclass of extinct small to giant tetrapods — often considered primitive amphibians — that flourished worldwide during the Carboniferous , Permian , and Triassic periods . A few species continued into the Cretaceous . Fossils have been found on every continent . During about 210 million years of evolutionary history , they adapted to a wide range of habitats , including fresh water , terrestrial , and even coastal marine environments . Their life history is well understood , with fossils known from the larval stage , metamorphosis , and maturity . Most temnospondyls were semiaquatic , although some were almost fully terrestrial , returning to the water only to breed . These temnospondyls were some of the first vertebrates fully adapted to life on land . Although temnospondyls are considered amphibians , many had characteristics , such as scales , claws , and armour @-@ like bony plates , that distinguish them from modern amphibians .
 Temnospondyls have been known since the early 19th century , and were initially thought to be reptiles . They were described at various times as batrachians , stegocephalians , and labyrinthodonts , although these names are now rarely used . Animals now grouped in Temnospondyli were spread out among several amphibian groups until the early 20th century , when they were found to belong to a distinct taxon based on the structure of their vertebrae . Temnospondyli means " cut vertebrae " , as each vertebra is divided into several parts .
 Experts disagree over whether temnospondyls were ancestral to modern amphibians ( frogs , salamanders , and caecilians ) , or whether the whole group died out without leaving any descendants . Different hypotheses have placed modern amphibians as the descendants of temnospondyls , another group of early tetrapods called lepospondyls , or even as descendants of both groups ( with caecilians evolving from lepospondyls and frogs and salamanders evolving from temnospondyls ) . Recent studies place a family of temnospondyls called the amphibamids as the closest relatives of modern amphibians . Similarities in teeth , skulls , and hearing structures link the two groups .

 = = Description = =

 Many temnospondyls are much larger than living amphibians , and superficially resemble crocodiles . Others are smaller and resemble salamanders . Most have broad , flat heads that are either blunt ( brevirostrine ) or elongated ( longirostrine ) . The skulls are rounded or triangular in shape when viewed from above , and are usually covered in pits and ridges . The rugged surfaces of bones may have supported blood vessels , which could transfer carbon dioxide to the bones to neutralize acidic build up in the blood ( early semiaquatic tetrapods would have had difficulty expelling carbon dioxide from their bodies while on land , and these dermal bones may have been an early solution to the problem ) . Many temnospondyls also have canal @-@ like grooves in their skulls called sensory sulci . The sulci , which usually run around the nostrils and eye sockets , are part of a lateral line system used to detect vibrations in water . As semiaquatic animals , most temnospondyls have small limbs with four toes on each front foot and five on each hind foot . Terrestrial temnospondyls have larger , thicker limbs , and some even have claws . One unusual terrestrial temnospondyl , Fayella , has relatively long limbs for its body , and probably lived as an active runner able to chase prey .
 Homologues of most of the bones of temnospondyls are also seen in other early tetrapods , aside from a few bones in the skull , such as interfrontals , internasals , and interparietals , that have developed in some temnospondyl taxa . Most temnospondyls have tabular horns in the backs of their skulls , rounded projections of bone separated from the rest of the skull by indentations called otic notches ; in some temnospondyls , such as Zatrachys , they are pointed and very prominent . Among the most distinguishing features of temnospondyls are the interpterygoid vacuities , two large holes in the back of the palate . Another pair of holes , choanae , are present in front of these vacuities , and connect the nasal passage with the mouth . Temnospondyls often have teeth on their palates , as well as in their jaws . Some of these teeth are so large , they are referred to as tusks . In some temnospondyls , such as Nigerpeton , tusks in the lower jaw pierce the palate and emerge through openings in the top of the skull .
 Very little is known of the soft tissue of temnospondyls . A block of sandstone , described in 2007 from the Early Carboniferous Mauch Chunk Formation of Pennsylvania , included impressions of the bodies of three temnospondyls . These impressions show , when alive , they had smooth skin , robust limbs with webbed feet , and a ridge of skin on their undersides . Trackways referable to small temnospondyls have also been found in Carboniferous and Permian rocks . The trackways , called batrachichni , are usually found in strata deposited around freshwater environments , suggesting the animals had some ties to the water .
 Unlike modern amphibians , many temnospondyls are covered in small , closely packed scales . The undersides of most temnospondyls are covered in rows of large ventral plates . During early stages of development , they first have only small , rounded scales . Fossils show , as the animals grew , the scales on the undersides of their bodies developed into large , wide ventral plates . The plates overlap each other in a way that allows a wide range of flexibility . Later semiaquatic temnospondyls , such as trematosaurs and capitosaurs , have no evidence of scales . They may have lost scales to make movement easier under water or to allow cutaneous respiration , the absorption of oxygen through the skin .
 Several groups of temnospondyls have large bony plates on their backs . One temnospondyl , Peltobatrachus , has armour @-@ like plating that covers both its back and underside . The temnospondyl Laidleria also has extensive plating on its back . Most members of the family Dissorophidae also have armor , although it only covers the midline of the back with two narrow rows of plates . Other temnospondyls , such as Eryops , have been found with small , disc @-@ like bony scutes that were in life probably embedded in the skin . All of these temnospondyls were adapted to a terrestrial lifestyle . Armor may have offered protection from predators in the case of Peltobatrachus . The scutes may have provided stability for the spine , as they would have limited flexibility and may have been connected by strong ligaments . Temnospondyls such as Sclerothorax and Eryops that may have been at least partly terrestrial also have long neural spines on top of their vertebrae that would have stabilized the spine . Bony scutes are also seen in plagiosaurs , but unlike Peltobatrachus , Laidleria , Eryops , and dissorophids , these animals are thought to have been fully aquatic . Plagiosaurs may have inherited their armor from a terrestrial ancestor , as both Peltobatrachus and Laidleria have been considered close relatives of the group .
 Temnospondyls ' vertebrae are divided into several segments . In living tetrapods , the main body of the vertebra is a single piece of bone called the centrum , but in temnospondyls , this region was divided into a pleurocentrum and intercentrum . Two types of vertebrae are recognized in temnospondyls : stereospondylous and rhachitomous vertebrae . In rhachitomous vertebrae , the intercentra are large and wedge @-@ shaped , and the pleurocentra are relatively small blocks that fit between them . Both elements support a spine @-@ like neural arch , and well @-@ developed interlocking projections called zygapophyses strengthen the connections between vertebrae . The strong backbone and strong limbs of many ratchitomous temnospondyls allowed them to be partially , and in some cases fully , terrestrial . In stereospondylous vertebrae , the pleurocentra have been lost entirely , with the intercentra enlarged as the main body of the vertebrae . This weaker type of backbone indicates the stereospondylous temnospondyls spent more time in water .

 = = History of study = =

 Temnospondyli was named by German palaeontologist Karl Alfred von Zittel in his second edition of Handbuch der Palaeontologie , published in 1888 . Temnospondyl remains were known since the early part of the 19th century , however . The earliest described temnospondyl was Mastodonsaurus , named by Georg Friedrich Jaeger in 1828 . Jaeger named Mastodonsaurus from a single tooth , and considered it a reptile . Mastodonsaurus means " breast tooth lizard " after the nipple @-@ like shape of the tip of the tooth .
 The naming of these first specimens was disputed , however . Leopold Fitzinger named the animal Batrachosaurus in 1837 . In 1841 , English palaeontologist Richard Owen referred to the genus as Labyrinthodon to describe its highly folded or labyrinthine teeth . Owen thought the name Mastodonsaurus " ought not to be retained , because it recalls unavoidably the idea of the mammalian genus Mastodon , or else a mammilloid form of the tooth ... and because the second element of the word , saurus , indicates a false affinity , the remains belonging , not to the Saurian , but to the Batrachian order of Reptiles . " Owen recognized the animal was not a " saurian " reptile , yet he also referred Jaeger 's Phytosaurus to the genus . Although the two genera both have similarly sized conical teeth , Phytosaurus was later found to be a crocodile @-@ like reptile . Additional material , including skulls , firmly placed Labyrinthodon as an amphibian . Jaeger also named Salamandroides giganteus in 1828 , basing it on partial occiput , or back portion of the skull . In 1833 , he described a complete skull of S. giganteus that had the same teeth as his Mastodonsaurus , making it the first known complete skull of a temnospondyl . Because Mastodonsaurus was named first , it has precedence over the other names as a senior subjective synonym . Batrachosaurus is still used as the name of an unrelated brachyopid temnospondyl .
 Mastodonsaurus and other similar animals were referred to as labyrinthodonts , named like Labyrinthodon for teeth that were highly folded in cross section . Owen 's " Labyrinthodon Jaegeri " was later found at Guy 's Cliffe , England by paleontologist William Buckland . Other specimens were found in the red sandstone of Warwickshire . As more fossils were uncovered in England , Owen depicted these labyrinthodonts as the " highest " form of batrachian and compared them to crocodiles , which he considered the highest form of reptiles . He also noted the large labyrinthodonts of the Keuper ( a unit of rocks that dates to the Late Triassic ) were younger than more advanced reptiles in the Magnesian and Zechstein , which are Late Permian in age . Owen used these fossils to counter the notion that reptiles evolved from a sequential progression from early amphibians ( what he called " metamorphosed fishes " ) .
 In addition to Mastodonsaurus , some of the earliest named genera included Metopias and Rhombopholis in 1842 , Zygosaurus in 1848 , Trematosaurus in 1849 , Baphetes and Dendrerpeton in 1853 , Capitosaurus in 1858 , and Dasyceps in 1859 . Baphetes is now placed as an early tetrapod outside Temnospondyli , and Rhombopholis is now considered a prolacertiform reptile .
 Later in the 19th century , temnospondyls were classified as various members of Stegocephalia , a name coined by American paleontologist Edward Drinker Cope in 1868 . Cope placed stegocephalians in the class Batrachia , the name then used for Amphibia . Stegocephalia means " roof @-@ headed " in Greek , a reference to the wide , flat heads of temnospondyls and other early tetrapods . During this time , palaeontologists considered temnospondyls to be amphibians because they possessed three main features : gill arches in juvenile skeletons , indicating they were amphibious for at least the first part of their lives ; ribs that do not connect at the underside of the rib cage ; and deep pits in the skull that were interpreted as space for mucous glands .
 Several suborders of stegocephalians were recognized in the late 19th and early 20th centuries . Animals now regarded as temnospondyls were primarily labyrinthodonts , but some were classified in the Branchiosauria . Branchiosaurs were small @-@ bodied and had simple conical teeth , while labyrinthodonts were larger and had complex , folded dentin and enamel in their teeth . Branchiosauria included only a few forms , such as Branchiosaurus from Europe and Amphibamus from North America , that had poorly developed bones , external gills , and no ribs . Some skeletons of Amphibamus were later found with long ribs , prompting its reassignment to Microsauria ( although more detailed studies found it to be a temnospondyl ) . Soft tissue , such as scales and external gills , were found in many well @-@ preserved branchiosaur fossils from Germany . In the early 20th century , branchiosaurs would be recognized as larval forms of temnospondyls lacking many of the typical features that define the group , and is no longer recognized as a distinct group .
 Other animals that would later be classified as temnospondyls were placed in a group called Ganocephala , characterized by plate @-@ like skull bones , small limbs , fish @-@ like scales , and branchial arches . Unlike labyrinthodonts , they did not have parietal foramena , small holes in their skulls behind their eye sockets . Archegosaurus , Dendrerpeton , Eryops and Trimerorhachis were placed in this group and were considered to be the most primitive members of Reptilia . Their rhachitomous vertebrae , notochord , and lack of occipital condyles ( which attached the head to the neck ) were features that were also shared with fishes . Thus , they were considered a link between early fishes and more advanced forms such as stegocephalians .
 Another group called Microsauria was named by Cope in 1868 . Cope classified Microsauria as a subgroup of Labyrinthodontia , placing many small , amphibian @-@ like animals within it . Among them were Dendrerpeton , once placed in Ganocephala . Dendrerpeton was later placed as a labyrinthodont with other temnospondyls , but confusion existed for many years over the classification of small amphibians .
 By the end of the 19th century , most of what are today regarded as temnospondyls were placed in the suborder Labyrinthodonta . American paleontologist Ermine Cowles Case called it Labyrinthodonta vera or " true labyrinthodonts " . The names Stegocephalia and Labyrinthodontia were used interchangeably to refer to the order in which it belonged . The labyrinthodontian suborders Microsauria and Branchiosauria , both of which contain temnospondyls , were distinct from Labyrinthodonta . Within Labyrinthodonta were the groups Rhachitomi , Labyrinthodonti , and Embolerimi . Members of Rhachitomi , such as Archegosaurus and Eryops , had rhachitomous vertebrae with enlarged intercentra that displaced the pleurocentra . Labyrinthodonti , such as Mastodonsaurus , Trematosaurus , and Micropholis , had lost their pleurocentra , and the intercentra made up the entire body of the vertebrae . Embolerimi had intercentra and pleurocentra that were of equal size . Embolomeres are now identified as reptiliomorphs distantly related to temnospondyls .
 In 1888 , von Zittel divided stegocephalians among three taxa : Lepospondyli , Temnospondyli , and Stereospondyli . He placed microsaurs in Lepospondyli , a group which he characterized as having simple , spool @-@ shaped vertebral centra . Temnospondyli included forms with the centra divided into pleurocentra and intercentra . All members of Stereospondyli had amphicoelous centra composed only of the intercentra . Cope objected to von Zittel 's classification , considering the vertebrae of lepospondyls and stereospondyls indistinguishable because each had a simple spool shape . He continued to use Ganocephala and Labyrinthodonta ( which he alternatively referred to as Rhachitomi ) to distinguish animals based on the absence or presence of occipital condyles .
 Temnospondyli became a commonly used name at the turn of the century . Paleontologists included both embolomeres and rhachitomes in the group . Cope 's Ganocephala and Labyrinthodonta fell out of use . In 1919 , British paleontologist D. M. S. Watson proposed that the evolutionary history of these large amphibians could be seen through changes in their vertebrae . Embolomerous forms in the Carboniferous graded into rhachitomous forms in the Permian , and finally into stereospondyls in the Triassic . More importantly , Watson began using the term Labyrinthodontia to refer to these groups . The name Temnospondyli was rarely used in the decades that followed . Swedish paleontologist Gunnar Säve @-@ Söderbergh removed embolomeres from the group , narrowing its scope to rhachitomes and stereospondyls . His classification of labyrinthodonts was based heavily on characteristics of the skull rather than the vertebrae .
 American paleontologist Alfred Romer brought the name Temnospondyli back into use in the later 20th century . Säve @-@ Söderbergh used the name Labyrinthodontia in a strict sense ( sensu stricto ) to refer to Rhachitomi and Stereospondyli , excluding Embolomeri . Romer agreed with this classification , but used the name Temnospondyli to avoid confusion with Labyrinthodontia in its wider sense ( sensu lato ) . Unlike modern temnospondyl classification , however , Romer included the primitive Ichthyostegalia in the group .

 = = Evolutionary history = =

 = = = Carboniferous and Early Permian = = =

 Temnospondyls first appeared in the Early Carboniferous around 330 million years ago ( Mya ) . During the Carboniferous , temnospondyls included basal medium @-@ sized forms such as Dendrerpeton or large semiaquatic forms such as Cochleosaurus . Other , more derived temnospondyls , such as the amphibamids , were smaller and more terrestrial . They resembled salamanders , and some taxa , such as the genus Branchiosaurus , even retained external gills like the modern @-@ day axolotl . During the latest Carboniferous and Early Permian around 300 Mya , several groups , such as the dissorophids and trematopids evolved strong , robust limbs and vertebrae and became adapted to life on land while others such as the eryopids , developed into large semiaquatic predators . The dvinosaurs , a group of small aquatic temnospondyls , evolved from terrestrial ancestors in the Late Carboniferous .

 = = = Late Permian = = =

 During the Late Permian , increasing aridity and the diversification of reptiles contributed into a decline in terrestrial temnospondyls , but semiaquatic and fully aquatic temnospondyls continued to flourish , including the large Melosaurus of Eastern Europe . Other temnospondyls , such as archegosaurids , developed long snouts and a close similarity to crocodiles , although they lacked the armor characteristic of the latter group . These temnospondyls included the largest known amphibian , the 9 @-@ m @-@ long Prionosuchus of Brazil .

 = = = Mesozoic = = =

 As temnospondyls continued to flourish and diversify in the Late Permian ( 260 @.@ 4 - 251 @.@ 0 Mya ) , a major group called Stereospondyli became more dependent on life in the water . The vertebrae became weak , the limbs small , and the skull large and flat , with the eyes facing upwards . During the Triassic period , these animals dominated the freshwater ecosystems , evolving in a range of both small and large forms . During the Early Triassic ( 251 @.@ 0 - 245 @.@ 0 Mya ) one group of successful long @-@ snouted fish @-@ eaters , the trematosauroids , even adapted to a life in the sea , the only known amphibians to do so with the exception of the modern crab @-@ eating frog . Another group , the capitosauroids , included medium- and large @-@ sized animals 2 @.@ 3 to 4 m ( 7 @.@ 5 to 13 @.@ 1 ft ) in length , with large and flat skulls that could be over a meter long in the largest forms such as Mastodonsaurus . These animals spent most or all their lives in water as aquatic predators , catching their prey by a sudden opening of the upper jaw and sucking in fish or other small animals .
 In the Carnian stage of the Late Triassic ( 228 @.@ 0 - 216 @.@ 5 Mya ) , capitosauroids were joined by the superficially very similar Metoposauridae . Metoposaurids are distinguished from capitosauroids by the positioning of their eye sockets near the front of their skulls . Another group of stereospondyls , the plagiosaurs , had wide heads with external gills , and adapted to life at the bottom of lakes and rivers . By this time , temnospondyls had become a common and widespread component of semiaquatic ecosystems . Some temnospondyls , such as Cryobatrachus and Kryostega , even inhabited Antarctica , which was covered in temperate forests at the time .
 Triassic temnospondyls were often the dominant semiaquatic animals in their environments . Large assemblages of metoposaurs with hundreds of individuals preserved together have been found in the southwestern United States . They have often been interpreted as mass death events caused by droughts in floodplain environments . Recent studies show these dense assemblages were instead probably the result of currents accumulating dead individuals in certain areas . These environments seem to have had little diversity , as they were inhabited almost exclusively by metoposaurs .
 The Triassic @-@ Jurassic extinction event around 199 @.@ 6 Mya led to the extinction of most Mesozoic temnospondyls . The brachyopoids survived , as well as a few capitosauroids and trematosauroids . While the latter two groups soon became extinct , brachyopoids persisted and grew to large sizes during the Jurassic . Among brachyopoids , the brachyopids flourished in China and the chigutisaurids became common in Gondwana . The most recent known temnospondyl was the giant chigutisaurid Koolasuchus , known from the Early Cretaceous of Australia . It survived in rift valleys that were too cold in the winter for crocodiles that normally would have competed with them . Koolasuchus was one of the largest of the brachyopoids , with an estimated weight of 500 kg ( 1 @,@ 100 lb ) .

 = = Classification = =

 Originally , temnospondyls were classified according to the structure of their vertebrae . Early forms , with complex vertebrae consisting of a number of separate elements , were placed in the suborder Rachitomi , and large Triassic aquatic forms with simpler vertebrae were placed in the suborder Stereospondyli . With the recent growth of phylogenetics , this classification is no longer viable . The basic rhachitomous condition is found in many primitive tetrapods , and is not unique to one group of temnospondyls . Moreover , the distinction between rhachitomous and stereospondylous vertebrae is not entirely clear . Some temnospondyls have rhachitomous , semirhachitomous , and sterospondylous vertebrae at different points in the same vertebral column . Other taxa have intermediate morphologies that do not fit into any category . Rachitomi is no longer recognized as a group , but Stereospondyli is still considered valid . Below is a simplified taxonomy of temnospondyls showing currently recognized groups :
 Class Amphibia
 Order Temnospondyli
 Superfamily Edopoidea
 Family Cochleosauridae ( Chenoprosopidae )
 Family Edopidae
 Family Dendrerpetontidae
 Suborder Euskelia
 Superfamily Dissorophoidea
 Family Amphibamidae
 Family Branchiosauridae
 Family Dissorophidae
 Family Micromelerpetontidae
 Superfamily Eryopoidea
 Family Eryopidae
 Family Parioxyidae
 Family Zatrachydidae
 Clade Limnarchia
 Clade Stereospondylomorpha
 Superfamily Archegosauroidea
 Family Actinodontidae
 Family Archegosauridae
 Family Intasuchidae ( placement is uncertain )
 Family Sclerocephalidae
 Suborder Stereospondyli
 Family Peltobatrachidae
 Family Lapillopsidae
 Family Rhinesuchidae
 Family Lydekkerinidae
 Clade Capitosauria
 Superfamily Mastodonsauroidea ( Capitosauroidea )
 Family Heylerosauridae
 Family Mastodonsauridae
 Family Stenotosauridae
 Infraorder Trematosauria
 Superfamily Trematosauroidea
 Superfamily Metoposauroidea
 Superfamily Plagiosauroidea
 Superfamily Brachyopoidea
 Superfamily Rhytidosteoidea

 = = = Phylogeny = = =

 In one of the earliest phylogenetic analyses of the group , Gardiner ( 1983 ) recognized five characteristics that made Temnospondyli a clade : a bone at the back of the skull , the parasphenoid , is connected to another bone on the underside of the skull , the pterygoid ; large openings called interpterygoid vacuities are present between the pterygoids ; the stapes ( a bone involved in hearing ) is connected to the parasphenoid and projects upward ; the cleithrum , a bone in the pectoral girdle , is thin ; and part of the vertebra called the interdorsal attaches to the neural arch . Additional features were given by Godfrey et al . ( 1987 ) , including the contact between the postparietal and exoccipital at the back of the skull , small projections ( uncinate processes ) on the ribs , and a pelvic girdle with each side having a single iliac blade . These shared characteristics are called synapomorphies .
 Temnospondyls are placed as basal tetrapods in phylogenetic analyses , with their exact positioning varying between studies . Depending on the classification of modern amphibians , they are either included in the crown group Tetrapoda or the stem of Tetrapoda . Crown @-@ group tetrapods are descendants of the most recent common ancestor of all living tetrapods and stem tetrapods are forms that are outside the crown group . Modern amphibians have recently been suggested as descendants of temnospondyls , which would place them within crown Tetrapoda . Below is a cladogram from Ruta et al . ( 2003 ) placing Temnospondyli within crown Tetrapoda :
 Other studies place modern amphibians as the descendants of lepospondyls and place temnospondyls in a more basal position within the stem of Tetrapoda . Below is a cladogram from Laurin and Reisz ( 1999 ) placing Temnospondyli outside crown Tetrapoda :
 Most phylogenetic analyses of temnospondyl interrelationships focus on individual families . One of the first broad @-@ scale studies of temnospondyl phylogeny was conducted by paleontologist Andrew Milner in 1990 . A 2007 study made a " supertree " of all temnospondyl families , combining the family @-@ level trees of previous studies . The following cladogram is modified from Ruta et al . ( 2007 ) :
 1 Temnospondyli , 2 Edopoidea , 3 Dvinosauria , 4 Euskelia , 5 Eryopoidea , 6 Dissorophoidea , 7 Limnarchia , 8 Archegosauroidea , 9 Stereospondyli , 10 Rhytidostea , 11 Brachyopoidea , 12 Capitosauria , 13 Trematosauria , 14 Metoposauroidea
 The most basal group of temnospondyls is the superfamily Edopoidea . Edopoids have several primitive or plesiomorphic features , including a single occipital condyle and a bone called the intertemporal that is absent in other temnospondyls . Edopoids include the Late Carboniferous genus Edops and the family Cochleosauridae . Dendrerpetontidae has also been included in Edopoidea , and is the oldest known temnospondyl family . Balanerpeton woodi is the oldest species , having been present over 330 million years ago during the Viséan stage of the Early Carboniferous . Recent analyses place Dendrerpetontidae outside Edopoidea in a more derived position . Other primitive temnospondyls include Capetus and Iberospondylus . Saharastega and Nigerpeton , both described in 2005 from Niger , are also primitive yet come from the Late Permian . They are almost 40 million years younger than other basal temnospondyls , implying a long ghost lineage of species that are not yet known in the fossil record .
 In 2000 , paleontologists Adam Yates and Anne Warren produced a revised phylogeny of more derived temnospondyls , naming several new clades . Two major clades were Euskelia and Limnarchia . Euskelia includes the temnospondyls that were once called rhachitomes and includes two subfamilies , the Dissorophoidea and the Eryopoidea . Dissorophoids include small , mostly terrestrial temnospondyls that may be the ancestors of modern amphibians . Eryopoids include larger temnospondyls like Eryops . The second major clade , Limnarchia , includes most Mesozoic temnospondyls , as well as some Permian groups . Within Limnarchia are the superfamily Archegosauroidea and the most derived temnospondyls , the stereospondyls .
 Yates and Warren also named Dvinosauria , a clade of small aquatic temnospondyls from the Carboniferous , Permian , and Triassic . They placed Dvinosauria within Limnarchia , but more recent studies disagree on their position . For example , a 2007 study places them even more basal than euskelians , while a 2008 study keeps them as basal limnarchians .
 Within Stereospondyli , Yates and Warren erected two major clades : Capitosauria and Trematosauria . Capitosaurs include large semiaquatic temnospondyls like Mastodonsaurus with flat heads and eyes near the back of the skull . Trematosaurs include a diversity of temnospondyls , including large marine trematosauroids , aquatic plagiosaurs , brachyopoids that survived into the Cretaceous , and metoposauroids with eyes near the front of their heads . In 2000 , paleontologists Rainer Schoch and Andrew Milner named a third major clade of stereospondyls , the Rhytidostea . This group included more primitive stereospondyls that could not be placed in either Capitosauria or Trematosauria , and included groups like Lydekkerinidae , Rhytidosteidae , and Brachyopoidea . While Capitosauria and Trematosauria are still widely used , Rhytidostea is not often supported as a true clade in recent analyses . Rhytidosteids and brachyopoids are now grouped with trematosaurians , but lydekkerinids are still considered to be a primitive family of stereospondyls .
 A new phylogeny of temnospondyls was offered by paleontologist Rainer Schoch in 2013 . It supported many of the clades that were found by Yates and Warren , but it did not find support for their division of derived stereospondyls into Euskelia and Limnarchia . Eryopids were found to be more closely related to stereospondyls than to dissorophoids , which were grouped with dvinosaurs . The clade including Eryopidae and Stereospondylomorpha was named Eryopiformes . In addition , Schoch named the clade containing all temnospondyls except edopoids Eutemnospondyli and reinstated the name Rhachitomi for the clade containing all temnospondyls except edopoids and dendrerpetontids . Below is the cladogram from Schoch 's analysis :

 = = = Relationship to modern amphibians = = =

 Modern amphibians ( frogs , salamanders , and caecilians ) are classified in Lissamphibia . Lissamphibians appear to have arisen in the Permian . Molecular clock estimates place the first lissamphibian in the Late Carboniferous , but the first member of Batrachia ( frogs and salamanders , but not caecilians ) is estimated to have appeared in the Middle Permian using the same technique . Using fossil evidence , there are three main theories for the origin of modern amphibians .
 One is that they evolved from dissorophoid temnospondyls . Another is that they evolved from lepospondyls , most likely the lysorophians . A third hypothesis is that caecilians descended from lepospondyls and frogs and salamanders evolved from dissorophoids .
 Recently , the theory that temnospondyls were the ancestors of all lissamphibians has gained wide support . The skull morphology of some small temnospondyls has been compared to those of modern frogs and salamanders , but the presence of bicuspid , pedicellate teeth in small , paedomorphic or immature temnospondyls has been cited as the most convincing argument in favor of the temnospondyl origin of lissamphibians . Seen in lissamphibians and many dissorophoid temnospondyls , pedicellate teeth have calcified tips and bases . During the development of most tetrapods , teeth begin to calcify at their tips . Calcification normally proceeds downward to the base of the tooth , but calcification from the tip stops abruptly in pedicellate teeth . Calcification resumes at the base , leaving an area in the center of the tooth uncalcified . This pattern is seen in living amphibians and fossils .
 The dissorophoid family Amphibamidae is thought to be most closely related to Lissamphibia . In 2008 , an amphibamid called Gerobatrachus hottoni was named from Texas and was nicknamed the " frogamander " for its frog @-@ like head and salamander @-@ like body . It was thought to be the most closely related temnospondyl to lissamphibians and was placed as the sister taxon of the group in a phylogenetic analysis . Another species of amphibamid called Doleserpeton annectens is now thought to be even more closely related to lissamphibians . Unlike Gerobatrachus , Doleserpeton was known since 1969 , and the presence of pedicellate teeth in its jaws has led some paleontologists to conclude soon after its naming that it was a relative of modern amphibians . It was first described as a " protolissamphibian " , and the specific name annectens means " connecting " in reference to its inferred transitional position between temnospondyls and lissamphibians . The structure of its tympanum , a disk @-@ like membrane that functions like an ear drum , is similar to that of frogs and has also been used as evidence for a close relationship . Other features including the shape of the palate and the back of the skull , the short ribs , and the smooth skull surface also point to it being a closer relative of lissamphibians than is Gerobatrachus . Below is a cladogram modified from Sigurdsen and Bolt ( 2010 ) showing the relationships of Gerobatrachus , Doleserpeton , and Lissamphibia :

 = = Paleobiology = =

 = = = Feeding = = =

 Although the earliest temnospondyls were primarily semiaquatic , they had the ability to feed on land . Later , eryopoids and dissorophoids , some well adapted to terrestrial life , also fed on land . Some eryopoids became better adapted toward life in water , and shifted their diets toward aquatic organisms . The first primarily aquatic feeders were archegosaurs in the Permian . Trematosaurs and capitosaurs became independently aquatic and also returned to this type of feeding .
 Most aquatic stereospondyls have flattened heads . When feeding , they probably opened their mouths by lifting their skulls instead of lowering their lower jaws . The jaw mechanics of the plagiosaurid Gerrothorax is well known , and is one of the most highly adapted . Gerrothorax is thought to have lifted its skull to around 50 ° above horizontal through the flexing of the atlanto @-@ occipital joint between the occipital condyles of the skull and the atlas vertebra of the neck . As the skull is raised , the quadrate bone pushes forward and causes the lower jaw to protrude outward . Other stereospondyls probably also lifted their skulls , but they are not as well adapted for such movement . D.M.S. Watson was the first to suggest skull lifting as a means of feeding in temnospondyls . He envisioned that Mastodonsaurus , a much larger temnospondyl than Gerrothorax , was able to make the same movement . Paleontologist A.L. Panchen also supported the idea in 1959 , suggesting that Batrachosuchus also fed in this way . At the time it was thought that these temnospondyls lifted their heads with strong jaw muscles , but it is now thought that they used larger muscles in the neck that were attached to the large pectoral girdle . Plagiosuchus , a close relative of Gerrothorax , also has a hyobranchial skeleton that muscles may have attached to . Plagiosuchus has very small teeth and a large area for muscle attachment behind the skull , suggesting that it could suction feed by rapidly opening its mouth .
 Unlike semiaquatic temnospondyls , terrestrial temnospondyls have skulls that are adapted for biting land @-@ living prey . The sutures between the bones of the skull in the dissorophoid Phonerpeton are able to withstand a high degree of compression . Compressive forces would have been experienced when biting down on prey . Earlier aquatic tetrapods and tetrapod ancestors differ from temnospondyls like Phonerpeton in that their skulls were also built to withstand tension . This tension would have been experienced during suction feeding underwater . Temnospondyls like Phonerpeton were among the first tetrapods that were almost exclusively terrestrial and fed by biting .

 = = = Reproduction = = =

 Temnospondyls , like all amphibians , reproduced in aquatic environments . Most temnospondyls probably reproduced through external fertilization . Like most living frogs , female temnospondyls would have laid masses of eggs in water while males released sperm to fertilize them . Several fossils were described from the Early Permian of Texas in 1998 that may be egg masses of dissorophoid temnospondyls . They were the first known fossils of amphibian eggs . The fossils consist of small disks with thin membranes that are probably vitelline membranes and halo @-@ like areas surrounding them that are most likely mucous coatings . They are attached to plant fossils , suggesting that these temnospondyls laid eggs on aquatic plants much like modern frogs . The mucous membranes show that the eggs were laid by amphibians , not fish ( their eggs lack mucous ) , but the type of amphibian that laid them cannot be known because no body fossils are preserved with the eggs . The eggs are thought to be from dissorophoids because they are likely to be close relatives of modern amphibians , and probably had similar reproductive strategies . They are also the most common amphibians from the deposit in which the eggs were found .
 One temnospondyl , the dvinosaur Trimerorhachis , may have brooded young in an area between the gills called the pharyngeal pouch . Small bones belonging to younger Trimerorhachis individuals have been found in these pouches . The living Darwin 's Frog is also a mouth brooder and would be the closest modern analogue to Trimerorhachis if it cared for its young in this way . An alternative possibility is that Trimerorhachis was cannibalistic , eating its young like many amphibians do today . If this was the case , the bones of these smaller individuals were originally located in the throat and were pushed into the pharyngeal pouch as the animal fossilized .
 Body impressions of Early Carboniferous temnospondyls from Pennsylvania suggest that some terrestrial temnospondyls mated on land like some modern amphibians . They reproduced through internal fertilization rather than mating in water . The presence of three individuals in one block of sandstone shows that the temnospondyls were gregarious . The head of one individual rests under the tail of another in what may be a courtship display . Internal fertilization and similar courtship behavior are seen in modern salamanders .

 = = = Growth = = =

 While most types of temnospondyls are distinguished on the basis of features in mature specimens , several are known from juvenile and larval specimens . Metamorphosis is seen in dissorophoids , eryopids , and zatrachydids , with aquatic larvae developing into adults capable of living on land . Several types of dissorophoids do not fully metamorphose , but retain features of juveniles such as gills and small body size in what is known as neoteny . Dvinosaurians and the plagiosaurid Gerrothorax were also neotenic because they retained gills , but they are only known from adult specimens .
 Temnospondyl larvae are often distinguished by poorly developed bones and the presence of a hyobranchial apparatus , a series of bones that gills would attach to in life . However , some fully mature temnospondyls also possess hyobranchial bones but did not have external gills . A dense covering of scales is also seen in larvae and adults . Major body changes occur in metamorphosis , including the reshaping and strengthening of skull bones , the thickening of postcranial bones , and an increase in body size .
 Temnospondyls like Sclerocephalus are known from both large adult specimens and small larvae , showing an extreme change in body shape . In these species , the shape and proportions of skull bones change in the early stages of development . The ornamentation on the surface of the skull roof also develops at this time . Small , regularly spaced pits are the first to form , followed by larger ridges . As development continues , the external gills disappear . Small teeth that once covered the palate are lost . The postcranial skeleton does not develop at the same rate as the skull , with ossification ( the replacement of cartilage by bone ) happening more slowly . Vertebrae and limb bones are poorly developed , ribs and fingers are absent in the early stages , and the scapulocoracoid and ischium are entirely absent through most of development . Once maturity is reached , most bones have fully formed and growth rate slows . The bones of some temnospondyls like Dutuitosaurus show growth marks , possibly an indication that growth rate varied with the change in seasons . Fossils of temnospondyls like Metoposaurus and Cheliderpeton show that individuals grew larger past maturity . The oldest individuals usually have more pitting on their skulls with deeper sulci .
 One group of temnospondyls , the Branchiosauridae , is also known from larval specimens . Branchiosaurids like Branchiosaurus and Apateon are represented by many fossils preserving skin and external gills . An entire growth series is exhibited in the wide range of sizes among specimens , but the lack of terrestrially adapted adult forms suggests that these temnospondyls were neotenic . Unlike other temnospondyls , their postcranial skeletons developed quickly but were still partly cartilaginous when fully mature . Adults likely had an aquatic lifestyle similar to juveniles . Recently , large specimens of Apateon gracilis were described with adaptations toward a terrestrial lifestyle , indicating that not all branchiosaurs were neotenic .
 While most temnospondyls are aquatic in early stages of life , most metoposaurids appear to have been terrestrial in their juvenile stage . Like other Mesozoic temnospondyls , adult metoposaurids were adapted to a semiaquatic lifestyle . Their bones are not highly developed for movement on land . The cross @-@ sectional thickness of limb bones in adult metoposaurids shows that they could not withstand the stress of terrestrial locomotion . Juvenile individuals have bones that are thick enough to withstand this stress , and could probably move about on land . To maintain a terrestrial lifestyle , a temnospondyl 's limb bones would have to thicken with positive allometry , meaning that they would grow at a greater rate than the rest of the body . This is not the case in metoposaurids , meaning that as their bodies grew larger they became less adapted toward a terrestrial lifestyle .

 = = = Hearing = = =

 Temnospondyls and other early tetrapods have rounded otic notches in the back of the skull that project into the cheek region . In life , the otic notch would have been covered by a membrane called the tympanum , which is seen as a disk @-@ like area in living frogs . The tympanum is involved in hearing , and is similar to the ear drum of more advanced tetrapods . It was traditionally thought that the tympanum developed very early in tetrapod evolution as a hearing organ and progressed to form the ear drum of amniotes . Thus , temnospondyls possessed a hearing system supposedly ancestral to that of living amphibians and reptiles .
 Frogs and all other living tetrapods have a rod @-@ like bone called the stapes that aids in hearing by transferring vibrations from the ear drum — or homologous tympanum — to the inner ear . Temnospondyls also have a stapes , which projects into the otic cavity . The stapes likely evolved from the hyomandibula of lobe @-@ finned fishes . The positioning of the stapes and the shape of the otic region suggests that the tympani of temnospondyls and frogs are homologous , but the tympani of these amphibians are no longer considered homologous with the hearing systems of reptiles , birds , and mammals . Therefore , ear structures in temnospondyls were not ancestral to those of all other tetrapods .
 The ability of the tympanum and stapes to effectively transmit vibrations is called impedance matching . Early tetrapods like temnospondyls have thick stapes with poor impedance matching , so it is now thought that they were not used for hearing . Instead , these thick stapes may have functioned to support the tissue that covers the otic notch . Early temnospondyls like Dendrerpeton could not hear airborne sound but would have been able to detect vibration in the ground . Later temnospondyls like Doleserpeton had otic regions adapted to hearing . Doleserpeton has a structure in the inner ear called the perilymphatic duct , which is also seen in frogs and is associated with hearing . Its stapes is also a better transmitter of sound . The hearing system of Doleserpeton and related temnospondyls was able to detect airborne sound and may have been ancestral to that of living amphibians .


 = Osbert de Bayeux =

 Osbert de Bayeux ( floruit 1121 to 1184 ) was a medieval English cleric and archdeacon in the Diocese of York . A relative of Thurstan , the Archbishop of York , Osbert probably owed his ecclesiastical positions to this relative . After Thurstan 's death , Osbert was opposed to one of the candidates for the archbishopric , William fitzHerbert , and worked to secure fitzHerbert 's deposition and replacement by Henry Murdac . After Murdac 's death in 1153 , Osbert tried to prevent the return of fitzHerbert , but these attempts were unsuccessful . When fitzHerbert died suddenly in 1154 , Osbert was accused of murdering the newly returned archbishop . Although he was never convicted of the murder in either a secular or an ecclesiastical court , he was stripped of his clerical status and became a layman before 1158 . He died after 1184 , perhaps even after 1194 .

 = = Early life = =

 Osbert was first mentioned in the historical record between 1121 and 1128 when he appears in a charter , which although likely a forgery , probably contains an authentic witness list . This document lists him as " Osbert archdeacon " , which means that he probably held the archdeaconry of Richmond . He was the nephew of Thurstan , who was Archbishop of York from 1114 to 1140 . Presumably he owed his position as archdeacon to his uncle and was probably appointed at a young age . A charter of Thurstan 's , dating to around 1138 , names Osbert explicitly as Thurstan 's nephew .

 = = Opposition to William fitzHerbert = =

 Osbert was opposed to the election of William fitzHerbert as Archbishop of York and supported William 's rival and successor Henry Murdac . Although he remained a supporter of Murdac after 1147 , he did oppose Murdac 's interventions in Selby Abbey , where Murdac had deposed one abbot and appointed another . In 1153 , Osbert deposed Murdac 's choice as abbot of Selby and appointed another abbot . Originally , Osbert had supported Elias Paynel , Murdac 's choice for abbot , but then changed his stance and helped with the deposition .
 After Murdac 's death in 1153 , Osbert was opposed to William 's return as archbishop , but was unsuccessful in his attempts to prevent William 's reappointment . William died a week after his return to York , however , and Osbert , along with Robert of Ghent , the Dean of York , secured the quick election of the new archbishop , Roger de Pont L 'Évêque .

 = = Poisoning accusations = =

 Osbert was accused of murdering William , specifically by poisoning him through the communion chalice . A fellow cleric , Symphorian , who had been a chaplain of the deceased archbishop , brought murder charges against Osbert . Symphorian obtained a hearing on the charges at a royal council presided over by King Stephen of England at Michaelmas in 1154 , but Stephen 's subsequent death prevented a resolution . Osbert attempted to have the trial switched to an ecclesiastical court and was supported in his efforts by Archbishop Theobald of Canterbury . A trial was finally held in 1156 and Osbert 's accuser did not produce any witnesses , but Osbert was unable to prove his innocence , prompting the transfer of the case to a papal court . No record of any judgment exists , but Osbert apparently appeared before two popes , Adrian IV and Alexander III . A further appeal to the papal court was referred to papal judges @-@ delegate between 1175 and 1180 .
 The case attracted commentary by two contemporary writers . John of Salisbury , who was a secretary for Theobald , added information about Osbert in a letter to Alexander III on unrelated business . In the section of the letter , John pointed out to the pope that no matter what others might say about Osbert , he had failed to secure other clergy willing to swear that he was innocent . Another contemporary , Gilbert Foliot , who was Bishop of Hereford , wrote to the pope to remind him that although Osbert 's accuser had offered to prove his accusations by undergoing a trial by ordeal , this was essentially meaningless since canon law forbade the clergy from the ordeal .

 = = Later life and death = =

 Osbert was no longer archdeacon by 1158 , as his successor is attested by that point . Osbert , however , continued to call himself " archdeacon " even though he held land as a secular lord , including lands in Lacy and Skipton . He also acted as a steward for Hugh de Tilly . Osbert was still alive in 1184 , as he was a witness to a document at York then , and may have been alive as late as 1194 , when Hugh Bardulf was responsible for the farm of Osbert 's lands , as the record of that transaction in the escheat roll is unclear if Osbert was alive at that time or dead .
 Osbert had two sons , William de Bayeux and Turstin de Baius . Osbert was a benefactor to a number of monasteries , including Drax Priory , Pontefract Priory and Gisborough Priory . He also gave land to a hospital in York and to the Templars and Hospitallers .


 = Dvorak technique =

 The Dvorak technique ( developed between 1969 and 1984 by Vernon Dvorak ) is a widely used system to estimate tropical cyclone intensity ( which includes tropical depression , tropical storm , and hurricane / typhoon / intense tropical cyclone intensities ) based solely on visible and infrared satellite images . Within the Dvorak satellite strength estimate for tropical cyclones , there are several visual patterns that a cyclone may take on which define the upper and lower bounds on its intensity . The primary patterns used are curved band pattern ( T1.0 @-@ T4.5 ) , shear pattern ( T1.5 @-@ T3.5 ) , central dense overcast ( CDO ) pattern ( T2.5 @-@ T5.0 ) , central cold cover ( CCC ) pattern , banding eye pattern ( T4.0 @-@ T4.5 ) , and eye pattern ( T4.5 - T8.0 ) .
 Both the central dense overcast and embedded eye pattern use the size of the CDO . The CDO pattern intensities start at T2.5 , equivalent to minimal tropical storm intensity ( 40 mph , 65 km / h ) . The shape of the central dense overcast is also considered . The eye pattern utilizes the coldness of the cloud tops within the surrounding mass of thunderstorms and contrasts it with the temperature within the eye itself . The larger the temperature difference is , the stronger the tropical cyclone . Once a pattern is identified , the storm features ( such as length and curvature of banding features ) are further analyzed to arrive at a particular T @-@ number . The CCC pattern indicates little development is occurring , despite the cold cloud tops associated with the quickly evolving feature .
 Several agencies issue Dvorak intensity numbers for tropical cyclones and their precursors , including the National Hurricane Center 's Tropical Analysis and Forecast Branch ( TAFB ) , the NOAA / NESDIS Satellite Analysis Branch ( SAB ) , and the Joint Typhoon Warning Center at the Naval Meteorology and Oceanography Command in Pearl Harbor , Hawaii .

 = = Evolution of the method = =

 The initial development of this technique occurred in 1969 by Vernon Dvorak , using satellite pictures of tropical cyclones within the northwest Pacific ocean . The system as it was initially conceived involved pattern matching of cloud features with a development and decay model . As the technique matured through the 1970s and 1980s , measurement of cloud features became dominant in defining tropical cyclone intensity and central pressure of the tropical cyclone 's low @-@ pressure area . Use of infrared satellite imagery led to a more objective assessment of the strength of tropical cyclones with eyes , using the cloud top temperatures within the eyewall and contrasting them with the warm temperatures within the eye itself . Constraints on short term intensity change are used less frequently than they were back in the 1970s and 1980s . The central pressures assigned to tropical cyclones have required modification , as the original estimates were 5 @-@ 10 hPa ( 0 @.@ 15 @-@ 0 @.@ 29 inHg ) too low in the Atlantic and up to 20 hPa ( 0 @.@ 59 inHg ) too high in the northwest Pacific . This led to the development of a separate wind @-@ pressure relationship for the northwest Pacific , devised by Atkinson and Holliday in 1975 , then modified in 1977 .
 As human analysts using the technique lead to subjective biases , efforts have been made to make more objective estimates using computer programs , which have been aided by higher @-@ resolution satellite imagery and more powerful computers . Since tropical cyclone satellite patterns can fluctuate over time , automated techniques use a six @-@ hour averaging period to lead to more reliable intensity estimates . Development of the objective Dvorak technique began in 1998 , which performed best with tropical cyclones that had eyes ( of hurricane or typhoon strength ) . It still required a manual center placement , keeping some subjectivity within the process . By 2004 , an advanced objective Dvorak technique was developed which utilized banding features for systems below hurricane intensity and to objectively determine the tropical cyclone 's center . A central pressure bias was uncovered in 2004 relating to the slope of the tropopause and cloud top temperatures which change with latitude that helped improve central pressure estimates within the objective technique .

 = = Details of the method = =

 In a developing cyclone , the technique takes advantage of the fact that cyclones of similar intensity tend to have certain characteristic features , and as they strengthen , they tend to change in appearance in a predictable manner . The structure and organization of the tropical cyclone are tracked over 24 hours to determine if the storm has weakened , maintained its intensity , or strengthened . Various central cloud and banding features are compared with templates that show typical storm patterns and their associated intensity . If infrared satellite imagery is available for a cyclone with a visible eye pattern , then the technique utilizes the difference between the temperature of the warm eye and the surrounding cold cloud tops to determine intensity ( colder cloud tops generally indicate a more intense storm ) . In each case a " T @-@ number " ( an abbreviation for Tropical Number ) and a Current Intensity ( CI ) value are assigned to the storm . These measurements range between 1 ( minimum intensity ) and 8 ( maximum intensity ) . The T @-@ number and CI value are the same except for weakening storms , in which case the CI is higher . For weakening systems , the CI is held as the tropical cyclone intensity for 12 hours , though research from the National Hurricane Center indicates that six hours is more reasonable . The table at right shows the approximate surface wind speed and sea level pressure that corresponds to a given T @-@ number . The amount a tropical cyclone can change in strength per 24 ‑ hour period is limited to 2 @.@ 5 T @-@ numbers per day .

 = = = Pattern types = = =

 Within the Dvorak satellite strength estimate for tropical cyclones , there are several visual patterns that a cyclone may take on which define the upper and lower bounds on its intensity . The primary patterns used are curved band pattern ( T1.0 @-@ T4.5 ) , shear pattern ( T1.5 @-@ T3.5 ) , central dense overcast ( CDO ) pattern ( T2.5 @-@ T5.0 ) , banding eye pattern ( T4.0 @-@ T4.5 ) , eye pattern ( T4.5 - T8.0 ) , and central cold cover ( CCC ) pattern . Both the central dense overcast and embedded eye pattern utilize the size of the CDO . The CDO pattern intensities start at T2.5 , equivalent to minimal tropical storm intensity ( 40 miles per hour ( 64 km / h ) ) . The shape of the central dense overcast is also considered . The farther the center is tucked into the CDO , the stronger it is deemed . Tropical cyclones with maximum sustained winds between 65 miles per hour ( 105 km / h ) and 100 miles per hour ( 160 km / h ) can have their center of circulations obscured by cloudiness within visible and infrared satellite imagery , which makes diagnosis of their intensity a challenge .
 The CCC pattern , with its large and quickly developing mass of thick cirrus clouds spreading out from an area of convection near a tropical cyclone center within a short time frame , indicates little development . When it develops , rainbands and cloud lines around the tropical cyclone weaken and the thick cloud shield obscures the circulation center . While it resembles a CDO pattern , it is rarely seen .
 The eye pattern utilizes the coldness of the cloud tops within the surrounding mass of thunderstorms and contrasts it with the temperature within the eye itself . The larger the temperature difference is , the stronger the tropical cyclone . Winds within tropical cyclones can also be estimated by tracking features within the CDO using rapid scan geostationary satellite imagery , whose pictures are taken minutes apart rather than every half @-@ hour .
 Once a pattern is identified , the storm features ( such as length and curvature of banding features ) are further analyzed to arrive at a particular T @-@ number .

 = = Usage = =

 Several agencies issue Dvorak intensity numbers for tropical cyclones and their precursors . These include the National Hurricane Center 's Tropical Analysis and Forecast Branch ( TAFB ) , the National Oceanic and Atmospheric Administration 's Satellite Analysis Branch ( SAB ) , and the Joint Typhoon Warning Center at the Naval Pacific Meteorology and Oceanography Center in Pearl Harbor , Hawaii .
 The National Hurricane Center will often quote Dvorak T @-@ numbers in their tropical cyclone products . The following example is from discussion number 3 of Tropical Depression 24 ( eventually Hurricane Wilma ) of the 2005 Atlantic hurricane season :
 BOTH TAFB AND SAB CAME IN WITH A DVORAK SATELLITE INTENSITY ESTIMATE OF T2.5 / 35 KT . HOWEVER ... OFTENTIMES THE SURFACE WIND FIELD OF LARGE DEVELOPING LOW PRESSURE SYSTEMS LIKE THIS ONE WILL LAG ABOUT 12 HOURS BEHIND THE SATELLITE SIGNATURE . THEREFORE ... THE INITIAL INTENSITY HAS ONLY BEEN INCREASED TO 30 KT .
 Note that in this case the Dvorak T @-@ number ( in this case T2.5 ) was simply used as a guide but other factors determined how the NHC decided to set the system 's intensity .
 The Cooperative Institute for Meteorological Satellite Studies ( CIMSS ) at the University of Wisconsin – Madison has developed the Objective Dvorak Technique ( ODT ) . This is a modified version of the Dvorak technique which uses computer algorithms rather than subjective human interpretation to arrive at a CI number . This is generally not implemented for tropical depressions or weak tropical storms . The China Meteorological Agency ( CMA ) is expected to start using the standard 1984 version of Dvorak in the near future . The Indian Meteorological Department ( IMD ) prefers using visible satellite imagery over infrared imagery due to a perceived high bias in estimates derived from infrared imagery during the early morning hours of convective maximum . The Japan Meteorological Agency ( JMA ) uses the infrared version of Dvorak over the visible imagery version . Hong Kong Observatory and JMA continue to utilize Dvorak after tropical cyclone landfall . Various centers hold on to the maximum current intensity for 6 – 12 hours , though this rule is broken when rapid weakening is obvious .
 Citizen science site Cyclone Center uses a modified version of the Dvorak technique to categorize post @-@ 1970 tropical weather .
 Satellite Images of Selected Tropical Storms and Associated T @-@ Number

 = = Benefits and disadvantages = =

 The most significant benefit of the use of the technique is that it has provided a more complete history of tropical cyclone intensity in areas where aircraft reconnaissance is neither possible nor routinely available . Intensity estimates of maximum sustained wind are currently within 5 miles per hour ( 8 @.@ 0 km / h ) of what aircraft are able to measure half of the time , though the assignment of intensity of systems with strengths between moderate tropical @-@ storm force ( 60 miles per hour ( 97 km / h ) ) and weak hurricane- or typhoon @-@ force ( 100 miles per hour ( 160 km / h ) ) is the least certain . Its overall precision has not always been true , as refinements in the technique led to intensity changes between 1972 and 1977 of up to 20 miles per hour ( 32 km / h ) . The method is internally consistent in that it constrains rapid increases or decreases in tropical cyclone intensity . Some tropical cyclones fluctuate in strength more than the 2 @.@ 5 T numbers per day limit allowed by the rule , which can work to the technique 's disadvantage and has led to occasional abandonment of the constraints since the 1980s . Systems with small eyes near the limb , or edge , of a satellite image can be biased too weakly using the technique , which can be resolved through use of polar @-@ orbiting satellite imagery . Subtropical cyclone intensity cannot be determined using Dvorak , which led to the development of the Hebert @-@ Poteat technique in 1975 . Cyclones undergoing extratropical transition , losing their thunderstorm activity , see their intensities underestimated using the Dvorak technique . This led to the development of the Miller and Lander extratropical transition technique which can be used under these circumstances .


 = New York State Route 31B =

 New York State Route 31B ( NY 31B ) was a state highway in central New York in the United States . It served as a connector between NY 31 , its parent route , in the Cayuga County village of Weedsport and NY 5 in the Onondaga County town of Elbridge . NY 31B was assigned c . 1933 , replacing New York State Route 293 , a route assigned as part of the 1930 renumbering of state highways in New York . The NY 31B designation was removed in 1980 and replaced by County Route 31B ( CR 31B ) in Cayuga County and CR 99 in Onondaga County .

 = = Route description = =

 NY 31B began at an intersection with its parent route , NY 31 , in the Cayuga County village of Weedsport . The highway went eastward , intersecting with NY 34 less than 0 @.@ 1 miles ( 0 @.@ 2 km ) later . Much of Weedsport was urbanized , with the highway passing residential homes and businesses as it progressed eastward through the village . The highway intersected with CR 12B before leaving Weedsport and entering the town of Brutus as Brutus Road .
 In Brutus , NY 31B continued to the east through the rural town , intersecting CR 136A and passing the Weedsport Rural Cemetery before turning to the southeast . After a short distance , the highway went through an isolated area of homes , where it intersected with CR 14 and CR 15A . NY 31B continued on , intersecting several local highways before crossing into Onondaga County and terminating at an intersection with NY 5 in the town of Elbridge .

 = = History = =

 What became NY 31B was originally designated as NY 293 as part of the 1930 renumbering of state highways in New York . NY 293 was renumbered to NY 31B c . 1933 , allowing the NY 293 designation to be transferred to another highway in Orange County . NY 31B remained unchanged until January 2 , 1980 , when the NY 31B designation was removed .
 Ownership and maintenance of NY 31B 's former routing was gradually transferred to the counties it went through , namely Cayuga and Onondaga . The first section to be transferred was the portion within Onondaga County , which was given to the county on April 1 , 1980 , and designated as CR 99 . The Cayuga County section of former NY 31B was transferred to the county one year later on April 1 , 1981 , and designated as CR 31B for the state route that preceded it .

 = = Major intersections = =


 = Ben Amos =

 Benjamin Paul " Ben " Amos ( born 10 April 1990 ) is an English professional footballer who plays as a goalkeeper for Bolton Wanderers . Born in Macclesfield , Cheshire , Amos began his career with Crewe Alexandra 's youth academy , but joined Manchester United at the age of 11 . He has spent time on loan at Peterborough United , Molde , and Oldham Athletic . Additionally , Amos is an England youth international , having represented his country at every level from Under @-@ 16 to Under @-@ 21 .

 = = Club career = =

 = = = Crewe Alexandra = = =

 Amos was born in Macclesfield , Cheshire and was a member of the Crewe Alexandra academy until he was released at the age of 10 . While at Crewe , he also played for another local team , Bollington United , as a centre midfielder . One year later , Amos was playing for his local team against the team at the top of the table , and they needed to win the match to win the league . Amos ' team 's goalkeeper was injured during the game , and as the tallest on the team , Amos was put in goal ; however , he had also been the team 's regular penalty taker all season , so when they were awarded a penalty , he went all the way up the pitch to take the kick . He scored , and his team went on to win the game 3 – 2 , together with the league title . After the game , his parents told him that a Manchester United scout had been watching him and that he had been invited for trials . Amos joined Manchester United at the age of 11 .

 = = = Manchester United = = =

 In his first season at Manchester United , Amos became a regular starter for the club 's Under @-@ 13 team , playing in 19 out of 27 matches in the 2001 – 02 season . Amos was named as an unused substitute for the Under @-@ 18 side for the first time on 8 January 2005 , for a league game against Manchester City . His first appearance for the Under @-@ 18s came exactly nine months later , on 8 October 2005 , coming on as a substitute for Danny Rose after starting goalkeeper Ron @-@ Robert Zieler was sent off in a 2 – 0 defeat to Bolton Wanderers . He was regularly named as an unused substitute during the 2005 – 06 season – including for two reserve team matches – but became a frequent starter for the Under @-@ 18s in 2006 – 07 after signing a trainee contract in July 2006 . However , he missed the final of the 2006 – 07 FA Youth Cup with a dislocated shoulder .
 He retained his place in the Under @-@ 18 team for 2007 – 08 , in addition to making his debut for the reserve team against Wigan Athletic on 7 November 2007 , and during the season he impressed enough to be selected to go on the first @-@ team 's 2008 summer tour of South Africa . Amos was named as a substitute for all three matches of the tour , but did not play . En route back from South Africa , United stopped off in Nigeria to play against Portsmouth on 27 July 2008 , with Amos replacing Tomasz Kuszczak after 76 minutes . He made his competitive first @-@ team debut on 23 September 2008 in a 3 – 1 win at home to Middlesbrough in the third round of the League Cup . On 14 December 2008 , Amos travelled to Japan with the Manchester United squad for the 2008 FIFA Club World Cup , having been called up as a late replacement for Ben Foster , who had suffered a hand injury while training .

 = = = Loans to Peterborough and Molde = = =

 On 29 October 2009 , Amos signed for the Championship team Peterborough United on a month 's loan as cover for Peterborough 's suspended first @-@ choice goalkeeper Joe Lewis . He made his only appearance on 31 October in a 2 – 1 defeat against Barnsley . After returning to Manchester United , Amos was again sent out on loan in March 2010 , this time to Norwegian side Molde FK , where he remained on loan until 30 June 2010 .

 = = = Return to Manchester United = = =

 Following the departure of Ben Foster from Manchester United to Birmingham City , United manager Alex Ferguson declared that Amos would be Manchester United 's third @-@ choice goalkeeper for the 2010 – 11 season behind Edwin van der Sar and Tomasz Kuszczak . He made his first appearance of the season on 26 October 2010 , starting in goal for United 's 3 – 2 win over Wolverhampton Wanderers in the fourth round of the League Cup .
 In United 's final Champions League group match on 7 December 2010 , Amos was picked to start against Valencia at Old Trafford . Pablo Hernández scored Valencia 's only goal past him after 32 minutes of the match – the first goal United had conceded in the Champions League that season – as the two sides played out a 1 – 1 draw .

 = = = Loan to Oldham Athletic = = =

 With the signing of Danish goalkeeper Anders Lindegaard , Manchester United allowed Amos to join Oldham Athletic on loan for the remainder of the season on 7 January 2011 , although he would continue to train with Manchester United once a week . He made his debut against Swindon Town the next day and kept a clean sheet . However , three days later , he conceded all six goals in a 6 – 0 defeat at home to Southampton ; he allowed Adam Lallana 's 20 @-@ yard shot underneath his body for the second goal , and he was rounded by Lee Barnard for the sixth .
 On 15 March 2011 , Lindegaard was ruled out for five weeks following a knee injury , so Amos was recalled from Oldham to cover for Edwin van der Sar and Tomasz Kuszczak .

 = = = Return to Manchester United = = =

 Amos made his first start of the 2011 – 12 season in a third round League Cup tie at Elland Road against Leeds United . He kept a clean sheet as United cruised through to the next round winning 3 – 0 . He played again in the following round away at Aldershot Town , a game which United also won 3 – 0 and advanced to the quarter finals . He was in goal again for United 's League Cup game in a defeat against Championship club Crystal Palace on 30 November . It also seemed that he had moved above last season 's second choice Tomasz Kuszczak in the pecking order , but was now third choice behind Anders Lindegaard and David de Gea . He made his first Premier League start in a 2 – 0 home win against Stoke City on 31 January 2012 , keeping a clean sheet on his debut .
 On 30 May 2012 , Amos signed a three @-@ year contract extension with Manchester United , which will keep him at the club until at least 2015 .

 = = = Loan to Hull City = = =

 On 31 July 2012 , Amos joined Championship team Hull City on a season @-@ long loan from Manchester United . Before the loan deal was completed , he had joined Hull on their pre @-@ season training camp in Portugal . He made his debut on 11 August 2012 in 7 – 6 penalty shoot @-@ out victory over Rotherham United in the first round of the 2012 – 13 Football League Cup . However , after 19 appearances , including two in the League Cup , Amos returned to Manchester United on 3 January 2013 .

 = = = Loan to Carlisle United = = =

 On 15 November 2013 , Amos joined League One team Carlisle United on a month @-@ long loan from Manchester United .

 = = = Loan to Bolton Wanderers = = =

 On 30 January 2015 he joined Championship club Bolton Wanderers on a month 's loan to provide competition for Andy Lonergan after Ádám Bogdán was ruled out with an ankle injury picked up in training . On 21 February 2015 he made his Bolton debut as a substitute for the injured Lonergan as Wanderers lost 4 – 1 to Nottingham Forest at the City Ground . He played nine Championship games for Bolton before his loan expired on 4 April .

 = = = Return to Manchester United = = =

 Ben Amos returned to Manchester United after his loan spell at Bolton Wanderers , but on 10 June 2015 , it was announced on Manchester United website that Ben Amos has been released from the club along with four other players .

 = = = Bolton Wanderers = = =

 On 1 July 2015 , Amos returned to Bolton Wanderers following his release from Manchester United , signing a four @-@ year contract with the club .

 = = International career = =

 Amos is an England youth international , having played for his country at the Under @-@ 16 , Under @-@ 17 , Under @-@ 18 , Under @-@ 19 , Under @-@ 20 and Under @-@ 21 levels . He was called up to the Under @-@ 21 squad in 2011 for their friendly match against Iceland , but he was an unused substitute .

 = = Career statistics = =

 As of 1 December 2015

 = = Honours = =

 = = = Club = = =

 Manchester United
 FIFA Club World Cup ( 1 ) : 2008

 = = Personal life = =

 Amos attended Fallibroome High School , where he earned 11 GCSEs at grade C or above .