Salientia

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Salientia
Temporal range:
Early TriassicPresent, 250–0  Ma
Hourglass Treefrogs (Hyla ebraccata) 3.jpg
Hourglass Treefrogs Hyla ebraccata
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Class: Amphibia
Superorder: Batrachia
Clade: Salientia
Laurenti, 1768
Subgroups

The Salientia (Latin salire, salio meaning "to jump") are a total group of amphibians that includes the order Anura, the frogs and toads, and various extinct proto-frogs that are more closely related to the frogs than they are to the Urodela, the salamanders and newts. [1] The oldest fossil "proto-frog" appeared in the early Triassic of Madagascar, but molecular clock dating suggests their origins may extend further back to the Permian, 265 million years ago.

Contents

Characteristics

Very few fossils of early salientians have been found, which makes defining the characteristics of the group and their taxonomic relationships difficult. The arrangement of pectoral elements and the number of vertebrae are some guides, but the degree of vertebral articulation and the arrangement of the bones in the leg have not been found to be reliable indicators. The early proto-frogs developed from temnospondyl ancestors in which some of the elements of their vertebrae remained separate. The structure of the salientian pelvis and hind limb was probably developed for swimming rather than jumping. From the structure of the vertebrae, the group appears not to be monophyletic. The evolution of salientians seems to have been rapid and radiative. The essential features of recent groupings seem to have been established during the Mesozoic or early Tertiary. The families Alytidae, Pipidae, and Pelobatidae are ecologically isolated, the harlequin frogs, restricted to a neotropical range in Central and South America, and the Ranidae and Bufonidae probably radiated from tropical regions of Africa and Asia. [2]

Evolution

The origins and evolutionary relationships between the three main groups of amphibians are hotly debated. A molecular phylogeny based on rDNA analysis dating from 2005 suggests that salamanders and caecilians are more closely related to each other than they are to frogs, and the divergence of the three groups took place in the Paleozoic or early Mesozoic before the breakup of the supercontinent Pangaea and soon after their divergence from the lobe-finned fishes. This would help account for the relative scarcity of amphibian fossils from the period before the groups split. [3] Another molecular phylogenetic analysis conducted about the same time concluded the lissamphibians first appeared about 330 million years ago and that the temnospondyl-origin hypothesis is more credible than other theories. The neobatrachians seemed to have originated in Africa/India, the salamanders in East Asia and the caecilians in tropical Pangaea. [4] Other researchers, while agreeing with the main thrust of this study, questioned the choice of calibration points used to synchronise the data. They proposed that the date of lissamphibian diversification be put in the Permian, rather less than 300 million years ago, a date in better agreement with the palaeontological data. [5] A further study in 2011 using both extinct and living taxa sampled for morphological, as well as molecular data, came to the conclusion that the Lissamphibia are monophyletic and should be nested within the Lepospondyli rather than within the Temnospondyli. The study postulated the Lissamphibia originated no earlier than the late Carboniferous, some 290 to 305 million years ago. The split between Anura and Caudata was estimated as taking place 292 million years ago, rather later than most molecular studies suggest, with the caecilians splitting off 239 million years ago. [6]

A fossilized frog from the Czech Republic, possibly Palaeobatrachus gigas Fossilised frog.jpg
A fossilized frog from the Czech Republic, possibly Palaeobatrachus gigas

In 2008, Gerobatrachus hottoni , a temnospondyl with many frog- and salamander-like characteristics, was discovered in Texas. It dated back 290 million years and was hailed as a missing link, a stem batrachian close to the common ancestor of frogs and salamanders, consistent with the widely accepted hypothesis that frogs and salamanders are more closely related to each other (forming a clade called the Batrachia) than they are to caecilians. [7] [8] However, others have suggested that Gerobatrachus hottoni was only a dissorophoid temnospondyl unrelated to extant amphibians. [9] The earliest known salientians (see below), closer to the extant frogs than to the extant salamanders, are Triadobatrachus massinoti, from the Early Triassic of Madagascar (about 250 million years ago), and the fragmentary Czatkobatrachus polonicus from the Early Triassic of Poland (about the same age as Triadobatrachus). [10] The skull of Triadobatrachus is frog-like, being broad with large eye sockets, but the fossil has features diverging from modern frogs. These include a longer body with more vertebrae. The tail has separate vertebrae, unlike the fused urostyle or coccyx found in modern frogs. The tibia and fibula bones are also separate, making it probable that Triadobatrachus was not an efficient leaper. [10] The Salientia (Latin salere (salio), "to jump") are a stem group including modern frogs in the order Anura and their close fossil relatives the "proto-frogs" (e.g., Triadobatrachus and Czatkobatrachus ). The common features possessed by the "proto-frogs" in the Salientia group include 14 presacral vertebrae (modern frogs have eight or nine), a long and forward-sloping ilium in the pelvis, the presence of a frontoparietal bone, and a lower jaw without teeth.

Species

The earliest salientian yet discovered is Triadobatrachus massinoti, known from a single fossil specimen found in Madagascar. It dates back to the Early Triassic, about 250 million years ago. It had many frog-like features, but had 14 presacral vertebrae, while modern frogs have nine or 10. Previous fossil amphibians had many more presacral vertebrae than this and T. massinoti provides a missing link between salamanders and frogs. Other characteristics that distinguish it from modern frogs include the possession of a short tail with unfused vertebrae, a separate radius and ulna in the fore limb, and separate tibia and fibula in the hind limb. The features it shares with modern frogs include a forward-sloping ilium, the fusion of the frontal and parietal bones into a single structure known as the frontoparietal, and a lower jaw bone with no teeth. [11]

Czatkobatrachus is another proto-frog with some characteristics similar to Triadobatrachus. It is from the early Triassic in Poland and has a shortened vertebral column, reduced tail, and elongated ilium. [12]

Another early proto-frog was Prosalirus bitis , several fossil specimens of which have been found in Arizona. It dates back to the Early Jurassic, 190 million years ago. It has primitive features, but has a urostyle and an elongated, forward-directed ilium in its pelvis. These adaptations made it better able to absorb the impact of landing after a jump. [13]

Dating back to a similar date is Vieraella herbsti , a single specimen of which has been found in Santa Cruz Province, Argentina. It had 10 presacral vertebrae, but is considered to be more basal than Notobatrachus and living frogs.

Several specimens of Notobatrachus degiustoi have been found in Patagonia, Argentina. They date back to the Middle Jurassic, 160 million years ago. Whether it should be considered the first modern frog or be placed in a sister group to Anura is uncertain. [14]

Phylogeny

Amphibia
Batrachia

Salientia

Caudata – Salamanders and newts

Albanerpetontidae – Extinct

Gymnophiona – Caecilians

Cladogram from Tree of Life Web Project. [15]

Related Research Articles

<span class="mw-page-title-main">Amphibian</span> Class of ectothermic tetrapods

Amphibians are ectothermic, anamniotic, four-limbed vertebrate animals that constitute the class Amphibia. In its broad sense, it is paraphyletic group encompassing all tetrapods excluding the amniotes. All extant (living) amphibians belong to the monophyletic subclass Lissamphibia, with three living orders: Anura (frogs), Urodela (salamanders), and Gymnophiona (caecilians). Evolved to be mostly semiaquatic, amphibians have adapted to inhabit a wide variety of habitats, with most species living in freshwater, wetland or terrestrial ecosystems. Their life cycle typically starts out as aquatic larvae with gills known as tadpoles, but some species have developed behavioural adaptations to bypass this.

<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">Caudata</span> Clade of amphibians

The Caudata are a group of amphibians containing the extant salamanders and all extinct species of amphibians more closely related to salamanders than to frogs. They are typically characterized by a superficially lizard-like appearance, with slender bodies, blunt snouts, short limbs projecting at right angles to the body, and the presence of a tail in both larvae and adults.

<span class="mw-page-title-main">Caecilian</span> Order of amphibians

Caecilians are a group of limbless, vermiform (worm-shaped) or serpentine (snake-shaped) amphibians with small or sometimes nonexistent eyes. They mostly live hidden in soil or in streambeds, and this cryptic lifestyle renders caecilians among the least familiar amphibians. Modern caecilians live in the tropics of South and Central America, Africa, and southern Asia. Caecilians feed on small subterranean creatures such as earthworms. The body is cylindrical and often darkly coloured, and the skull is bullet-shaped and strongly built. Caecilian heads have several unique adaptations, including fused cranial and jaw bones, a two-part system of jaw muscles, and a chemosensory tentacle in front of the eye. The skin is slimy and bears ringlike markings or grooves and may contain scales.

<span class="mw-page-title-main">Batrachia</span> Clade of amphibians

The Batrachia are a clade of amphibians that includes frogs and salamanders, but not caecilians nor the extinct allocaudates. The name Batrachia was first used by French zoologist Pierre André Latreille in 1800 to refer to frogs, but has more recently been defined in a phylogenetic sense as a node-based taxon that includes the last common ancestor of frogs and salamanders and all of its descendants. The idea that frogs and salamanders are more closely related to each other than either is to caecilians is strongly supported by morphological and molecular evidence; they are, for instance, the only vertebrates able to raise and lower their eyes. However, an alternative hypothesis exists in which salamanders and caecilians are each other's closest relatives as part of a clade called the Procera, with frogs positioned as the sister taxon of this group.

<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">Batrachomorpha</span> Clade of amphibians

The Batrachomorpha are a clade containing recent and extinct amphibians that are more closely related to modern amphibians than they are to mammals and reptiles. 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.

<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">Dissorophoidea</span> Extinct superfamily of amphibians

Dissorophoidea is a clade of medium-sized, temnospondyl amphibians that appeared during the Moscovian in Euramerica, and continued through to the Late Permian and the Early Triassic of Gondwana. They are distinguished by various details of the skull, and many species seem to have been well adapted for life on land.

<span class="mw-page-title-main">Temnospondyli</span> Ancestors of modern amphibians adapted to life on land

Temnospondyli or temnospondyls is a diverse ancient order of small to giant tetrapods—often considered primitive amphibians—that flourished worldwide during the Carboniferous, Permian and Triassic periods, with fossils being found on every continent. A few species continued into the Jurassic and Early Cretaceous periods, but all had gone extinct by the Late Cretaceous. During about 210 million years of evolutionary history, they adapted to a wide range of habitats, including freshwater, 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 amphibians, many had characteristics such as scales and armour-like bony plates that distinguish them from the modern soft-bodied lissamphibians.

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

Triadobatrachus is an extinct genus of salientian frog-like amphibians, including only one known species, Triadobatrachus massinoti. It is the oldest member of the frog lineage known, and an excellent example of a transitional fossil. It lived during the Early Triassic about 250 million years ago, in what is now Madagascar.

<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">Albanerpetontidae</span> Family of amphibians

The Albanerpetontidae are an extinct family of small amphibians, native to the Northern Hemisphere during the Mesozoic and Cenozoic. The only members of the order Allocaudata, they are thought to be allied with living amphibians belonging to Lissamphibia. Despite a superficially salamander-like bodyform, their anatomy is strongly divergent from modern amphibians in numerous aspects. The fossil record of albanerpetontids spans over 160 million years from the Middle Jurassic to the beginning of the Pleistocene, about 2.13–2 million years ago.

Czatkobatrachus is an extinct genus of Early Triassic (Olenekian) salientian amphibians. It was first described in 1998 based on fossils found in the Czatkowice 1 quarry in Poland. It is, with Triadobatrachus, one of the two oldest known lissamphibians. More precisely, it is a member of Salientia; it is related to, but outside Anura, the taxon that includes all extant frogs. It is known only from the early Triassic of Poland. Its vertebral column may have been short as in other salientians, but the exact count is unknown. It had a short tail, and an elongated ilium.

Doleserpeton is an extinct, monospecific genus of dissorophoidean temnospondyl within the family Amphibamidae that lived during the Upper Permian, 285 million years ago. Doleserpeton is represented by a single species, Doleserpeton annectens, which was first described by John R. Bolt in 1969. Fossil evidence of Doleserpeton was recovered from the Dolese Brothers Limestone Quarry in Fort Sill, Oklahoma. The genus name Doleserpeton is derived from the initial discovery site in Dolese quarry of Oklahoma and the Greek root "herp-", meaning "low or close to the ground". This transitional fossil displays primitive traits of amphibians that allowed for successful adaptation from aquatic to terrestrial environments. In many phylogenies, lissamphibians appear as the sister group of Doleserpeton.

<span class="mw-page-title-main">Amphibamidae</span> Ancient family of amphibians

The Amphibamidae are an ancient family of dissorophoid temnospondyls known from Late Carboniferous-Early Permian strata in the United States.

Chinlestegophis is a diminutive Late Triassic stereospondyl that has been interpreted as a putative stem caecilian, a living group of legless burrowing amphibians. If Chinlestegophis is indeed both an advanced stereospondyl and a relative of caecilians, this means that stereospondyls survived to the present day; historically the group was thought to have gone extinct by the early Cretaceous. Chinlestegophis jenkinsi, the type and only species, is known from two partial skulls discovered in the Chinle Formation in Colorado.

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

Triassurus is an extinct genus of amphibian, and the oldest member of Caudata. It is known from the Middle to Upper Triassic (Ladinian-Carnian) aged Madygen Formation in Kyrgyzstan. The type species is T. sixtelae.

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

Funcusvermis is an extinct genus of stem-caecilian from the Late Triassic of Arizona. It is based on a large sample of jaws and other skull and postcranial fragments, discovered in an approximately 220 million years old layer of rock in the Blue Mesa Member of the Chinle Formation at Petrified Forest National Park. There is a single species, called Funcusvermis gilmorei.

References

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  15. David Cannatella (2007). "Living amphibians". Tree of Life Web Project. Retrieved 2012-08-14.