Batrachomorpha

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Batrachomorpha
Temporal range: Early Carboniferous–present
Pangerpeton sinensis.jpg
Pangerpeton , a salamander from the Jurassic
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Superclass: Tetrapoda
Clade: Batrachomorpha
Säve-Söderbergh, 1934
Groups

The Batrachomorpha ("frog forms") are a clade containing extant and extinct amphibians that are more closely related to modern amphibians than they are to mammals and reptiles (including birds). According to many analyses they include the extinct Temnospondyli; some show that they include the Lepospondyli instead. The name traditionally indicated a more limited group.

Contents

The first tetrapods were all amphibians in the physiological sense that they laid their eggs in water, and are colloquially sometimes referred to as labyrinthodonts or stegocephalians. The term "amphibians" can be ambiguous, however. In modern phylogeny, the basalmost tetrapods are discerned from two derived branches, one consisting of the crown group of modern amphibians, the Lissamphibia, and its relatives. In this scheme, the name Batrachomorpha is given to this branch of amphibians, while the name Reptiliomorpha denotes the other branch. While the actual phylogeny of the modern amphibians is not well understood, their ancestors are descended from one line of batrachomorphs. [1] All other living tetrapods (reptiles, birds and mammals) are descended from one branch of reptiliomorphs, the amniotes. Amniotes achieved dominance, while all other reptiliomorphs and most batrachomorphs have gone extinct.

Classification

Etymology

The name Batrachomorpha was coined by the Swedish palaeontologist Gunnar Säve-Söderbergh in 1934 to refer to ichthyostegids, temnospondyls, anthracosaurs, and the frogs. Säve-Söderbergh held the view that salamanders and caecilians are not related to the other tetrapods, but had developed independently from a different group of lobe-finned fish, the porolepiformes. [2] In this view amphibians would be a biphyletic group, and Batrachomorpha was erected to form a natural group consisting of the "true amphibians" (i.e. frogs in Säve-Söderberghs view) and their fossil relatives. The salamanders and the Lepospondyli was consigned to "Urodelomorpha".

Friedrich von Huene adopted it as a superorder of his subclass "Eutetrapoda" (the lower tetrapods exclusive of the urodeles) and included the orders Stegocephalia (here including a number of Labyrinthodonts) and Anura. [3] Erik Jarvik, who took over Säve-Söderberghs work and shared his view of the origin of salamanders, used the term more informally, but in a wider sense, to include the ancestral osteolepiform fishes. [4]

Though never a majority view, the notion that tetrapods had evolved twice, together with the usage of the term batrachomorpha, lingered until genetic analysis started confirming the monophyly of living amphibians in the 1990s. [5] Jarviks classification is no longer followed, all living amphibians and their fossil relatives now being classified together in the group Lissamphibia.

Phylogenetics

The short, broad skull of Eryops is typical of the batrachomorphans PSM V27 D633 Skull of eryops megacephalus.jpg
The short, broad skull of Eryops is typical of the batrachomorphans

Michael Benton adopted the term Batrachomorpha to include all living amphibians and extinct relatives more closely related to amphibians than to amniotes. In his scheme, Batrachomorpha is a superorder of amphibious tetrapods containing the following subgroups: [6]

The other groups of tetrapods considered more closely related to amniotes are put in the superorder Reptiliomorpha. [6]

The phylogenetic relationships of Paleozoic tetrapods have not yet been worked out with certainty, and the validity of Batrachomorpha as a clade depends on where other amphibians and early amniotes fit on the evolutionary tree. The actual content of Batrachomorpha as cladistically defined is therefore uncertain, and in some phylogenies the clade is redundant (e.g. Laurin 1996). [7] [8]

Anatomy

Batrachomorphs are distinguished by a number of features in the skeleton, including a flat or shallow skull, a fused skull roof with no cranial kinesis, exoccipital-postparietal contact on the occiput, and four or fewer fingers on the hand. [9]

Benton contrasts Batrachomorphs with Reptiliomorphs; both are stem-based clades; the former constitutes the "amphibian" evolutionary radiation, the latter the contemporary proto-reptilian and early amniote evolution.

In the appendix to Vertebrate Palaeontology, which combines cladistic and linnaean rankings, Benton has given Batrachomorpha the rank of Subclass in his 2001 edition, Class in the 2004 edition, and Superorder in the 2014 edition.

Related Research Articles

<span class="mw-page-title-main">Tetrapod</span> Superclass of the first four-limbed vertebrates and their descendants

A tetrapod is any four-limbed vertebrate animal of the superclass Tetrapoda. Tetrapods include all extant and extinct amphibians and amniotes, with the latter in turn evolving into two major clades, the sauropsids and synapsids. Some tetrapods such as snakes, legless lizards, and caecilians had evolved to become limbless via mutations of the Hox gene, although some do still have a pair of vestigial spurs that are remnants of the hindlimbs.

<span class="mw-page-title-main">Lissamphibia</span> Subclass of amphibians

The Lissamphibia is a group of tetrapods that includes all modern amphibians. Lissamphibians consist of three living groups: the Salientia, the Caudata, and the Gymnophiona.

<span class="mw-page-title-main">Amniote</span> Clade of tetrapods including reptiles, birds and mammals

Amniotes are tetrapod vertebrate animals belonging to the clade Amniota, a large group that comprises the vast majority of living terrestrial and semiaquatic vertebrates. Amniotes evolved from amphibian ancestors during the Carboniferous period and further diverged into two groups, namely the sauropsids and synapsids. They are distinguished from the other living tetrapod clade — the non-amniote lissamphibians — by the development of three extraembryonic membranes, thicker and keratinized skin, and costal respiration.

<span class="mw-page-title-main">Vertebrate paleontology</span> Scientific study of prehistoric vertebrates

Vertebrate paleontology is the subfield of paleontology that seeks to discover, through the study of fossilized remains, the behavior, reproduction and appearance of extinct vertebrates. It also tries to connect, by using the evolutionary timeline, the animals of the past and their modern-day relatives.

<i>Ichthyostega</i> Extinct genus of tetrapodomorphs

Ichthyostega is an extinct genus of limbed tetrapodomorphs from the Late Devonian of what is now Greenland. It was among the earliest four-limbed vertebrates ever in the fossil record and was one of the first with weight-bearing adaptations for terrestrial locomotion. Ichthyostega possessed lungs and limbs that helped it navigate through shallow water in swamps. Although Ichthyostega is often labelled a 'tetrapod' because of its limbs and fingers, it evolved long before true crown group tetrapods and could more accurately be referred to as a stegocephalian or stem tetrapod. Likewise, while undoubtedly of amphibian build and habit, it is not a true member of the group in the narrow sense, as the first modern amphibians appeared in the Triassic Period. Until finds of other early stegocephalians and closely related fishes in the late 20th century, Ichthyostega stood alone as a transitional fossil between fish and tetrapods, combining fish and tetrapod features. Newer research has shown that it had an unusual anatomy, functioning more akin to a seal than a salamander, as previously assumed.

<span class="mw-page-title-main">Erik Jarvik</span> Swedish paleontologist

Anders Erik Vilhelm Jarvik was a Swedish paleontologist who worked extensively on the sarcopterygian fish Eusthenopteron. In a career that spanned some 60 years, Jarvik produced some of the most detailed anatomical work on this fish, making it arguably the best known fossil vertebrate.

<span class="mw-page-title-main">Labyrinthodontia</span> Paraphyletic group of tetrapodomorphs

"Labyrinthodontia" is an informal grouping of extinct predatory amphibians which were major components of ecosystems in the late Paleozoic and early Mesozoic eras. Traditionally considered a subclass of the class Amphibia, modern classification systems recognize that labyrinthodonts are not a formal natural group (clade) exclusive of other tetrapods. Instead, they consistute an evolutionary grade, ancestral to living tetrapods such as lissamphibians and amniotes. "Labyrinthodont"-grade vertebrates evolved from lobe-finned fishes in the Devonian, though a formal boundary between fish and amphibian is difficult to define at this point in time.

<span class="mw-page-title-main">Lepospondyli</span> Polyphyletic group of tetrapodomorphs

Lepospondyli is a diverse taxon of early tetrapods. With the exception of one late-surviving lepospondyl from the Late Permian of Morocco, lepospondyls lived from the Early Carboniferous (Mississippian) to the Early Permian and were geographically restricted to what is now Europe and North America. Five major groups of lepospondyls are known: Adelospondyli; Aïstopoda; Lysorophia; Microsauria; and Nectridea. Lepospondyls have a diverse range of body forms and include species with newt-like, eel- or snake-like, and lizard-like forms. Various species were aquatic, semiaquatic, or terrestrial. None were large, and they are assumed to have lived in specialized ecological niches not taken by the more numerous temnospondyl amphibians that coexisted with them in the Paleozoic. Lepospondyli was named in 1888 by Karl Alfred von Zittel, who coined the name to include some tetrapods from the Paleozoic that shared some specific characteristics in the notochord and teeth. Lepospondyls have sometimes been considered to be either related or ancestral to modern amphibians or to Amniota. It has been suggested that the grouping is polyphyletic, with aïstopods being primitive stem-tetrapods, while recumbirostran microsaurs are primitive reptiles.

<span class="mw-page-title-main">Reptiliomorpha</span> Clade of reptile-like animals

Reptiliomorpha 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 defined by Michel Laurin (2001) and Vallin and Laurin (2004) as the largest clade that includes Homo sapiens, but not Ascaphus truei. Laurin and Reisz (2020) defined Pan-Amniota as the largest total clade containing Homo sapiens, but not Pipa pipa, Caecilia tentaculata, and Siren lacertina.

<span class="mw-page-title-main">Anthracosauria</span> Paraphyletic group of tetrapodomorphs

Anthracosauria is an order of extinct reptile-like amphibians that flourished during the Carboniferous and early Permian periods, although precisely which species are included depends on one's definition of the taxon. "Anthracosauria" is sometimes used to refer to all tetrapods more closely related to amniotes such as reptiles, mammals, and birds, than to lissamphibians such as frogs and salamanders. An equivalent term to this definition would be Reptiliomorpha. Anthracosauria has also been used to refer to a smaller group of large, crocodilian-like aquatic tetrapods also known as embolomeres.

<span class="mw-page-title-main">Diadectomorpha</span> Extinct clade of tetrapods

Diadectomorpha is a clade of large tetrapods that lived in Euramerica during the Carboniferous and Early Permian periods and in Asia during Late Permian (Wuchiapingian), They have typically been classified as advanced reptiliomorphs positioned close to, but outside of the clade Amniota, though some recent research has recovered them as the sister group to the traditional Synapsida within Amniota, based on inner ear anatomy and cladistic analyses. They include both large carnivorous and even larger herbivorous forms, some semi-aquatic and others fully terrestrial. The diadectomorphs seem to have originated during late Mississippian times, although they only became common after the Carboniferous rainforest collapse and flourished during the Late Pennsylvanian and Early Permian periods.

<i>Westlothiana</i> Extinct genus of tetrapods

Westlothiana is a genus of reptile-like tetrapod that lived about 338 million years ago during the latest part of the Viséan age of the Carboniferous. Members of the genus bore a superficial resemblance to modern-day lizards. The genus is known from a single species, Westlothiana lizziae. The type specimen was discovered in the East Kirkton Limestone at the East Kirkton Quarry, West Lothian, Scotland in 1984. This specimen was nicknamed "Lizzie the lizard" by fossil hunter Stan Wood, and this name was quickly adopted by other paleontologists and the press. When the specimen was formally named in 1990, it was given the specific name "lizziae" in homage to this nickname. However, despite its similar body shape, Westlothiana is not considered a true lizard. Westlothiana's anatomy contained a mixture of both "labyrinthodont" and reptilian features, and was originally regarded as the oldest known reptile or amniote. However, updated studies have shown that this identification is not entirely accurate. Instead of being one of the first amniotes, Westlothiana was rather a close relative of Amniota. As a result, most paleontologists since the original description place the genus within the group Reptiliomorpha, among other amniote relatives such as diadectomorphs and seymouriamorphs. Later analyses usually place the genus as the earliest diverging member of Lepospondyli, a collection of unusual tetrapods which may be close to amniotes or lissamphibians, or potentially both at the same time.

<i>Tulerpeton</i> Extinct genus of tetrapodomorphs

Tulerpeton is an extinct genus of Devonian four-limbed vertebrate, known from a fossil that was found in the Tula Region of Russia at a site named Andreyevka. This genus and the closely related Acanthostega and Ichthyostega represent the earliest tetrapods.

<span class="mw-page-title-main">Embolomeri</span> Extinct order of tetrapods

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 Carboniferous (Pennsylvanian) Period. They were specialized semiaquatic predators with long bodies for eel-like undulatory swimming. Embolomeres are characterized by their vertebral centra, which are formed by two cylindrical segments, the pleurocentrum at the rear and intercentrum at the front. These segments are equal in size. Most other tetrapods have pleurocentra and intercentra which are drastically different in size and shape.

<i>Solenodonsaurus</i> Extinct genus of reptiles

Solenodonsaurus is an extinct genus of reptiliomorphs that lived in what is now Czech Republic, during the Westphalian stage.

<i>Gerobatrachus</i> Extinct genus of amphibians

Gerobatrachus is an extinct genus of amphibamid temnospondyl that lived in the Early Permian, approximately 290 million years ago (Ma), in the area that is now Baylor County, Texas. When it was first described in 2008, Gerobatrachus was announced to be the closest relative of Batrachia, the group that includes modern frogs and salamanders. It possesses a mixture of characteristics from both groups, including a large frog-like head and a salamander-like tail. These features have led to it being dubbed a frogamander by the press. Some more recent studies place Gerobatrachus as the closest relative of Lissamphibia, the group that contains all modern amphibians including frogs, salamanders, and caecilians, or place modern amphibians far from Gerobatrachus within a group called Lepospondyli.

<span class="mw-page-title-main">Stegocephali</span> Clade of tetrapodomorphs

Stegocephali is a clade of vertebrate animals containing all fully limbed tetrapodomorphs. It is equivalent to a broad definition of the superclass Tetrapoda: under this broad definition, the term "tetrapod" applies to any animal descended from the first vertebrate with four limbs each with five digits in the extremity (pentadactyly), rather than fins of their sarcopterygian relatives.

<i>Acherontiscus</i> Extinct genus of amphibians

Acherontiscus is an extinct genus of stegocephalians that lived in the Early Carboniferous of Scotland. The type and only species is Acherontiscus caledoniae, named by paleontologist Robert Carroll in 1969. Members of this genus have an unusual combination of features which makes their placement within amphibian-grade tetrapods uncertain. They possess multi-bone vertebrae similar to those of embolomeres, but also a skull similar to lepospondyls. The only known specimen of Acherontiscus possessed an elongated body similar to that of a snake or eel. No limbs were preserved, and evidence for their presence in close relatives of Acherontiscus is dubious at best. Phylogenetic analyses created by Marcello Ruta and other paleontologists in the 2000s indicate that Acherontiscus is part of Adelospondyli, closely related to other snake-like animals such as Adelogyrinus and Dolichopareias. Adelospondyls are traditionally placed within the group Lepospondyli due to their fused vertebrae. Some analyses published since 2007 have argued that adelospondyls such as Acherontiscus may not actually be lepospondyls, instead being close relatives or members of the family Colosteidae. This would indicate that they evolved prior to the split between the tetrapod lineage that leads to reptiles (Reptiliomorpha) and the one that leads to modern amphibians (Batrachomorpha). Members of this genus were probably aquatic animals that were able to swim using snake-like movements.

<span class="mw-page-title-main">Stem tetrapoda</span>

The Stem Tetrapoda are a cladistically defined group, consisting of all animals more closely related to extant four-legged vertebrates than to their closest extant relatives, but excluding the crown group Tetrapoda. They are thus paraphyletic, though acceptable in phylogenetic nomenclature as the group is defined by strict reference to phylogeny rather than to traits as in traditional systematics. Thus, some finned sarcopterygians are considered to be stem tetrapods.

References

  1. Laurin, M. (1996): Phylogeny of Stegocephalians, from the Tree of Life Web Project
  2. Säve-Söderbergh, G. (1934). Some points of view concerning the evolution of the vertebrates and the classification of this group. Arkiv för Zoologi 26A: pp 1-20
  3. von Huene, F. (1956): Paläontologie und Phylogenie der niederen Tetrapoden, G. Fischer, Jena
  4. Jarvik, E. (1980). Basic structure and evolution of vertebrates. Vol. 2. London: Academic Press.
  5. San Mauro & al (2005): Initial diversification of living amphibians predated the breakup of Pangaea. American Naturalist no 165: pp 590–599
  6. 1 2 Benton, M. J. (Michael J.) (20 October 2014). Vertebrate palaeontology (Fourth ed.). Chichester, West Sussex. ISBN   978-1-118-40755-4. OCLC   867425553.{{cite book}}: CS1 maint: location missing publisher (link)
  7. Anderson & al. (2008): A stem batrachian from the Early Permian of Texas and the origin of frogs and salamanders. Nature 453: 515-518. doi : 10.1038/nature06865
  8. Laurin, M. (1996): Terrestrial Vertebrates - Stegocephalians: Tetrapods and other digit-bearing vertebrates
  9. Benton, M. J. (2000), Vertebrate Paleontology, 2nd Ed. Blackwell Science Ltd 3rd ed, pp.98-99

General references