Limnarchia
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Limnarchia is a
clade In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...
of
temnospondyl Temnospondyli (from Greek language, Greek τέμνειν, ''temnein'' 'to cut' and σπόνδυλος, ''spondylos'' 'vertebra') or temnospondyls is a diverse ancient order (biology), order of small to giant tetrapods—often considered Labyrinth ...
s. It includes the mostly
Carboniferous The Carboniferous ( ) is a Geologic time scale, geologic period and System (stratigraphy), system of the Paleozoic era (geology), era that spans 60 million years, from the end of the Devonian Period Ma (million years ago) to the beginning of the ...
-
Permian The Permian ( ) is a geologic period and System (stratigraphy), stratigraphic system which spans 47 million years, from the end of the Carboniferous Period million years ago (Mya), to the beginning of the Triassic Period 251.902 Mya. It is the s ...
age
Dvinosauria Dvinosaurs are one of several new clades of temnospondyls named in the phylogenetic review of the group by Yates and Warren 2000. They represent a group of primitive semi-aquatic to completely aquatic temnospondyls, and are known from the Late C ...
and the mostly
Permian The Permian ( ) is a geologic period and System (stratigraphy), stratigraphic system which spans 47 million years, from the end of the Carboniferous Period million years ago (Mya), to the beginning of the Triassic Period 251.902 Mya. It is the s ...
-
Triassic The Triassic ( ; sometimes symbolized 🝈) is a geologic period and system which spans 50.5 million years from the end of the Permian Period 251.902 million years ago ( Mya), to the beginning of the Jurassic Period 201.4 Mya. The Triassic is t ...
age
Stereospondylomorpha Stereospondylomorpha is a clade of temnospondyls. It includes the superfamily Archegosauroidea and the more diverse group Stereospondyli. Stereospondylomorpha was first proposed by Yates and Warren (2000), who found Archegosauroidea and Stereosp ...
. The clade was named in a 2000
phylogenetic In biology, phylogenetics () is the study of the evolutionary history of life using observable characteristics of organisms (or genes), which is known as phylogenetic inference. It infers the relationship among organisms based on empirical dat ...
analysis of
stereospondyl The Stereospondyli are a group of extinct temnospondyl amphibians that existed primarily during the Mesozoic period. They are known from all seven continents and were common components of many Triassic ecosystems, likely filling a similar ecologi ...
s and their relatives. Limnarchia means "lake rulers" in
Greek Greek may refer to: Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group *Greek language, a branch of the Indo-European language family **Proto-Greek language, the assumed last common ancestor of all kno ...
, in reference to their aquatic lifestyles and long existence over a span of approximately 200 million years from the
Late Carboniferous Late or LATE may refer to: Everyday usage * Tardy, or late, not being on time * Late (or the late) may refer to a person who is dead Music * Late (The 77s album), ''Late'' (The 77s album), 2000 * Late (Alvin Batiste album), 1993 * Late!, a pseudo ...
to the
Early Cretaceous The Early Cretaceous (geochronology, geochronological name) or the Lower Cretaceous (chronostratigraphy, chronostratigraphic name) is the earlier or lower of the two major divisions of the Cretaceous. It is usually considered to stretch from 143.1 ...
. In phylogenetic terms, Limnarchia is a
stem-based taxon Phylogenetic nomenclature is a method of nomenclature for taxon, taxa in biology that uses phylogenetics, phylogenetic definitions for taxon names as explained below. This contrasts with Biological classification, the traditional method, by which ...
including all temnospondyls more closely related to ''
Parotosuchus ''Parotosuchus'' is an extinct genus of capitosaurian temnospondyls within the family Mastodonsauridae. Fossils are known from the Early Triassic of Europe, Africa, Australia, and Antarctica. It was about long and likely lived in aquatic envir ...
'' than to ''
Eryops ''Eryops'' (; from Greek , , 'drawn-out' + , , 'face', because most of its skull was in front of its eyes) is a genus of extinct, amphibious temnospondyls. It contains the single species , the fossils of which are found mainly in early Permian (a ...
''. It is the
sister group In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree. Definition The expression is most easily illustrated by a cladogram: Taxon A and ...
of the clade
Euskelia Euskelia is a proposed clade of extinct temnospondyl amphibians. The naming derives from the ancient Greek ''eu'', meaning "true", and ''skelos'', meaning "limb", in reference to well-ossified limb bones with crests to which muscles were attached ...
, which is all temnospondyls more closely related to ''Eryops'' than to ''Parotosuchus''. Limnarchians represent an
evolutionary radiation An evolutionary radiation is an increase in taxonomic diversity that is caused by elevated rates of speciation, that may or may not be associated with an increase in morphological disparity. A significantly large and diverse radiation within ...
of temnospondyls into aquatic environments, while euskelians represent a radiation into terrestrial environments. While many euskelians were adapted to life on land with strong limbs and bony scutes, most limnarchians were better adapted for the water with poorly developed limbs and
lateral line The lateral line, also called the lateral line organ (LLO), is a system of sensory organs found in fish, used to detect movement, vibration, and pressure gradients in the surrounding water. The sensory ability is achieved via modified epithelia ...
sensory systems in their skulls. ''
Chinlestegophis ''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 relati ...
'', a putative Triassic stereospondyl considered to be related to metoposauroids such as ''
Rileymillerus ''Rileymillerus'' is an extinct genus of temnospondyl amphibian from the Late Triassic Post Quarry in the Dockum Group of Texas that was described by John Bolt and Sankar Chatterjee in 2000. The holotype, a nearly complete skull with articulated ...
'', has been noted to share many features with
caecilian 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 ...
s, a living group of legless burrowing amphibians. If ''Chinlestegophis'' is indeed both an advanced stereospondyl and a relative of caecilians, this means that limnarchians (in the form of caecilians) survived to the present day.


Description

Several characteristics unique to Limnarchia were identified when the clade was named in 2000. Most of these characteristics, called
synapomorphies In phylogenetics, an apomorphy (or derived trait) is a novel character or character state that has evolved from its ancestral form (or plesiomorphy). A synapomorphy is an apomorphy shared by two or more taxa and is therefore hypothesized to ...
, relate to the shapes of bones on the underside and back of the skull. Limnarchian synapomorphies include the presence of a row of teeth stemming from the ectopterygoid bone on the palate, the
maxilla In vertebrates, the maxilla (: maxillae ) is the upper fixed (not fixed in Neopterygii) bone of the jaw formed from the fusion of two maxillary bones. In humans, the upper jaw includes the hard palate in the front of the mouth. The two maxil ...
bone touching the
vomer The vomer (; ) is one of the unpaired facial bones of the skull. It is located in the midsagittal line, and articulates with the sphenoid, the ethmoid, the left and right palatine bones, and the left and right maxillary bones. The vomer forms ...
near the edge of the palate (these bones are separated by an opening in euskelian skulls), and the lack of bony projections called denticles on the vomer. Other characteristic features of limnarchians include the extension of the lower jaw behind the jaw joint, and a small hole called the paraquadrate foramen at the back of the skull. The only limnarchian
synapomorphy In phylogenetics, an apomorphy (or derived trait) is a novel Phenotypic trait, character or character state that has evolution, evolved from its ancestral form (or Plesiomorphy and symplesiomorphy, plesiomorphy). A synapomorphy is an apomorphy sh ...
not found in the skull is the elongated shape of the
interclavicle An interclavicle is a bone which, in most tetrapods, is located between the clavicles. Therian mammals ( marsupials and placentals) are the only tetrapods which never have an interclavicle, although some members of other groups also lack one. In ...
bone in the
pectoral girdle The shoulder girdle or pectoral girdle is the set of bones in the appendicular skeleton which connects to the arm on each side. In humans, it consists of the clavicle and scapula; in those species with three bones in the shoulder, it consists o ...
.


Phylogeny

Yates and Warren (2000) erected Limnarchia in their phylogenetic study of stereospondyls and their relatives. They found support for two major divisions of temnospondyls, which they named Euskelia and Limnarchia. Limnarchia included a group of small-bodied aquatic temnospondyls called
Dvinosauria Dvinosaurs are one of several new clades of temnospondyls named in the phylogenetic review of the group by Yates and Warren 2000. They represent a group of primitive semi-aquatic to completely aquatic temnospondyls, and are known from the Late C ...
as the most basal members of the clade, followed by the large-bodied
Archegosauroidea Archegosauroidea is an extinct superfamily of Permian temnospondyls. The superfamily is assigned to the clade Stereospondylomorpha and is the sister taxon to the suborder Stereospondyli. It includes the families Actinodontidae and Archegosauri ...
and the diverse
Stereospondyli The Stereospondyli are a group of extinct temnospondyl amphibians that existed primarily during the Mesozoic period. They are known from all seven continents and were common components of many Triassic ecosystems, likely filling a similar ecologi ...
, which together formed the clade
Stereospondylomorpha Stereospondylomorpha is a clade of temnospondyls. It includes the superfamily Archegosauroidea and the more diverse group Stereospondyli. Stereospondylomorpha was first proposed by Yates and Warren (2000), who found Archegosauroidea and Stereosp ...
. Below is a
cladogram A cladogram (from Greek language, Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an Phylogenetic tree, evolutionary tree because it does not s ...
from Yates and Warren (2000) showing this phylogeny:Sigurdsen, T.; Bolt, J.R. (2010). "The Lower Permian amphibamid Doleserpeton (Temnospondyli: Dissorophoidea), the interrelationships of amphibamids, and the origin of modern amphibians". Journal of Vertebrate Paleontology. 30 (5): 1360–1377. . Some more recent phylogenetic analyses such as Ruta ''et al.'' (2007) place Dvinosauria outside the Euskelia-Limnarchia split as much more basal temnospondyls. If this is the case, Limnarchia would include the same taxa as Stereospondylomorpha.


References

{{Taxonbar, from=Q6549732 Temnospondyli Pennsylvanian first appearances Early Cretaceous extinctions