Lepospondyls Temporal range: | |
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Diplocaulus , a diplocaulid "nectridean" | |
Scientific classification | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Clade: | Sarcopterygii |
Clade: | Tetrapodomorpha |
Clade: | Stegocephali |
Subclass: | † Lepospondyli Zittel, 1888 |
Groups | |
Lissamphibia? Contents |
Lepospondyli is a diverse taxon of early tetrapods. With the exception of one late-surviving lepospondyl from the Late Permian of Morocco ( Diplocaulus minimus ), [4] lepospondyls lived from the Visean stage of the Early Carboniferous to the Early Permian and were geographically restricted to what is now Europe and North America. [5] [2] 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 (the biggest genus, the diplocaulid Diplocaulus , reached a meter in length, but most were much smaller), 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 [6] [7] [8] or to Amniota (the clade containing reptiles and mammals). [9] It has been suggested that the grouping is polyphyletic, with aïstopods being primitive stem-tetrapods, while recumbirostran microsaurs are primitive reptiles. [10]
All lepospondyls are characterised by having simple, spool-shaped vertebrae that did not ossify from cartilage, but rather grew as bony cylinders around the notochord. In addition, the upper portion of the vertebra, the neural arch, is usually fused to the centrum (the main body of the vertebra). [11]
The position of the Lepospondyli within the Tetrapoda is uncertain because the earliest lepospondyls were already highly specialized when they first appeared in the fossil record. Some lepospondyls were once thought to be related or perhaps ancestral to modern salamanders (Urodela), but not the other modern amphibians. This view is no longer held and all modern amphibians (frogs, salamanders, and caecilians) are now grouped within the clade Lissamphibia. For a long time, the Lepospondyli were considered one of the three subclasses of Amphibia, along with the Lissamphibia and the Labyrinthodontia. [11] [12] [13] However, the dissolution of "labyrinthodonts" into separate groups such as temnospondyls and anthracosaurs has cast doubt on these traditional amphibian subclasses.
Much like "Labyrinthodontia", some studies proposed that Lepospondyli is an artificial (polyphyletic) grouping with some members closely related to extinct stem tetrapod groups and others more closely related to modern amphibians or reptiles. [14] Early phylogenetic analyses conducted in the 1980s and 1990s often maintained the idea that lepospondyls were paraphyletic, with nectrideans close to colosteids and microsaurs close to temnospondyls, which were considered to be ancestral to modern amphibians.
However, a 1995 paper by Robert Carroll argued that lepospondyls were actually a monophyletic group closer to reptiles. Carroll considered them closer to reptiles than the seymouriamorphs, but not as close as the diadectomorphs. [15] Many phylogenetic analyses since Carroll (1995) agreed with his interpretation, including Laurin & Reisz (1997), [16] Anderson (2001), [17] and Ruta et al. (2003). [9] A few have still considered lepospondyls ancestral to amphibians, but came to this conclusion without changing the position of lepospondyls compared to seymouriamorphs and diadectomorphs. [8]
Lepospondyl and tetrapod classification is still controversial, and even recent studies have had doubts about lepospondyl monophyly. For example, a 2007 paper has suggested that adelospondyls are stem-tetrapods close to colosteids [3] and a 2017 paper on Lethiscus has Aïstopoda in the tetrapod stem based on their primitive braincase. [18] These studies differ in the internal and external relationships of the remaining lepospondyl taxa. The former places the remaining lepospondyls into a single clade along the amniote stem. The latter does not treat the relationships of nectrideans or adelospondyls, but finds microsaurs to be early amniotes, and places lysorophians within microsaurs.
Five main groups of lepospondyls are often recognized: Microsauria, a superficially lizard- or salamander-like and species-rich group; Lysorophia, a group with elongated bodies and very small limbs; Aïstopoda, a group of limbless, extremely elongated snake-like forms; Adelospondyli, a group of presumably aquatic forms that resemble aïstopods, but have more solidly built skulls; and Nectridea, another diverse group that includes terrestrial and aquatic newt-like forms. Microsauria is generally considered paraphyletic; rather than being a monophyletic group, it has been considered an evolutionary grade of basal ("primitive") lepospondyls, although there is growing consensus that a large subset of fossorially-adapted microsaurs, the Recumbirostra, is monophyletic. Lysorophia may belong within the Recumbirostran clade, distinct from other derived lepospondyls. Nectridea may also be paraphyletic, consisting of a range of more anatomically-specialized lepospondyls. [19] The name Holospondyli has been proposed for a clade including aïstopods, and nectrideans, and possibly adelospondyls, although not all recent phylogenetic analyses support the grouping. The following cladogram, simplified, is after an analysis of tetrapods and stem-tetrapods presented by Ruta et al. in 2003: [9]
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The "lepospondyl hypothesis" of modern amphibian origins proposes that lissamphibians are monophyletic (that is, they form their own clade) and that they evolved from lepospondyl ancestors. Two alternatives are the "temnospondyl hypothesis", in which lissamphibians originated within Temnospondyli, and the "polyphyly hypothesis", in which caecilians originated from lepospondyls while frogs and salamanders (collectively grouped within Batrachia) evolved from temnospondyls. Of the three hypotheses, the temnospondyl hypothesis is currently the most widely accepted among researchers. Strong support for this relationship comes from a suite of anatomical features shared between lissamphibians and a group of Paleozoic temnospondyls called dissorophoids. [20] Under this hypothesis, Lepospondyli either falls outside crown group Tetrapoda (the smallest clade containing all living tetrapods, i.e. the smallest clade containing Lissamphibia and Amniota), or is closer to amniotes and therefore part of Reptiliomorpha. [9] However, some phylogenetic analyses continue to find support for the lepospondyl hypothesis. The analysis by Vallin and Laurin (2004) found lissamphibians to be most closely related to lysorophians, followed by microsaurs. Pawley (2006) also found lysorophians to be the closest relatives of lissamphibians, but found aïstopods and adelogyrinids rather than microsaurs to be the second most closely related groups. Marjanović (2010) found holospondyls to be the most closely related group to lissamphibians, followed by lysorophians. Under this hypothesis, lepospondyls would be crown tetrapods and temnospondyls would be stem tetrapods. [8]
Below is a cladogram from Ruta et al. (2003) that supports the "temnospondyl hypothesis", showing the position of Lepospondyli within crown group Tetrapoda: [9]
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.
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.
"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.
Adelospondyli is an order of elongated, presumably aquatic, Carboniferous amphibians. They have a robust skull roofed with solid bone, and orbits located towards the front of the skull. The limbs were almost certainly absent, although some historical sources reported them to be present. Despite the likely absence of limbs, adelospondyls retained a large part of the bony shoulder girdle. Adelospondyls have been assigned to a variety of groups in the past. They have traditionally been seen as members of the subclass Lepospondyli, related to other unusual early tetrapods such as "microsaurs", "nectrideans", and aïstopods. Analyses such as Ruta & Coates (2007) have offered an alternate classification scheme, arguing that adelospondyls were actually far removed from other lepospondyls, instead being stem-tetrapod stegocephalians closely related to the family Colosteidae.
Aistopoda is an order of highly specialised snake-like stegocephalians known from the Carboniferous and Early Permian of Europe and North America, ranging from tiny forms only 5 centimetres (2 in), to nearly 1 metre (3.3 ft) in length. They first appear in the fossil record in the Mississippian period and continue through to the Early Permian.
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.
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. The genus is known from a single species, Westlothiana lizziae. It is the oldest known uncontroversial tetrapod, closely related to but not an amniote.
Nectridea is the name of an extinct order of lepospondyl tetrapods from the Carboniferous and Permian periods, including animals such as Diplocaulus. In appearance, they would have resembled modern newts or aquatic salamanders, although they are not close relatives of modern amphibians. They were characterized by long, flattened tails to aid in swimming, as well as numerous features of the vertebrae.
Microsauria is an extinct, possibly polyphyletic order of tetrapods from the late Carboniferous and early Permian periods. It is the most diverse and species-rich group of lepospondyls. Recently, Microsauria has been considered paraphyletic, as several other non-microsaur lepospondyl groups such as Lysorophia seem to be nested in it. Microsauria is now commonly used as a collective term for the grade of lepospondyls that were originally classified as members of Microsauria.
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.
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.
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.
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.
Odonterpeton is an extinct genus of "microsaur" from the Late Carboniferous of Ohio, containing the lone species Odonterpeton triangulare. It is known from a single partial skeleton preserving the skull, forelimbs, and the front part of the torso. The specimen was found in the abandoned Diamond Coal Mine of Linton, Ohio, a fossiliferous coal deposit dated to the late Moscovian stage, about 310 million years ago.
Rhynchonkos is an extinct genus of rhynchonkid microsaur. Originally known as Goniorhynchus, it was renamed in 1981 because the name had already been given to another genus; the family, likewise, was originally named Goniorhynchidae but renamed in 1988. The type and only known species is R. stovalli, found from the Early Permian Fairmont Shale in Cleveland County, Oklahoma. Rhynchonkos shares many similarities with Eocaecilia, an early caecilian from the Early Jurassic of Arizona. Similarities between Rhynchonkos and Eocaecilia have been taken as evidence that caecilians are descendants of microsaurs. However, such a relationship is no longer widely accepted.
Utaherpeton is an extinct genus of lepospondyl amphibian from the Carboniferous of Utah. It is one of the oldest and possibly one of the most basal ("primitive") known lepospondyls. The genus is monotypic, including only the type species Utaherpeton franklini. Utaherpeton was named in 1991 from the Manning Canyon Shale Formation, which dates to the Mississippian-Pennsylvanian boundary. It was originally classified within Microsauria, a group of superficially lizard- and salamander-like lepospondyls that is now no longer considered to be a valid clade or evolutionary grouping, but rather an evolutionary grade consisting of the most basal lepospondyls. Utaherpeton has been proposed as both the most basal lepospondyl and the oldest "microsaur", although more derived lepospondyls are known from earlier in the Carboniferous. However, its position within Lepospondyli remains uncertain due to the incomplete preservation of the only known specimen. The inclusion of Utaherpeton in various phylogenetic analyses has resulted in multiple phylogenies that are very different from one another, making it a significant taxon in terms of understanding the interrelationships of lepospondyls.
Scincosaurus is an extinct genus of nectridean tetrapodomorphs within the family Scincosauridae.
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.
Recumbirostra is a clade of tetrapods which lived during the Carboniferous and Permian periods. They are thought to have had a fossorial (burrowing) lifestyle and the group includes both short-bodied and long-bodied snake-like forms. At least one species, the long-bodied molgophid Nagini mazonense, lost its forelimbs entirely. Recumbirostra includes the families Pantylidae, Gymnarthridae, Ostodolepidae, Rhynchonkidae and Brachystelechidae, with additional families such as Microbrachidae and Molgophidae being included by some authors. Brachystelechidae and Molgophidae have also been grouped together in the suggested clade Chthonosauria.
The Urocordylidae are an extinct family of nectridean lepospondyl amphibians. Urocordylids lived during the Late Carboniferous and Early Permian in what is now Europe and North America and are characterized by their very long, paddle-like tails. In life, they were probably newt-like and aquatic.