Lepospondyli is a diverse taxon of early tetrapods. With the exception of one late-surviving lepospondyl from the

Late Permian Late may refer to: * LATE, an acronym which could stand for: ** Limbic-predominant age-related TDP-43 encephalopathy, a proposed form of dementia ** Local-authority trading enterprise, a New Zealand business law ** Local average treatment effect, ...

Morocco Morocco (),, ) officially the Kingdom of Morocco, is the westernmost country in the Maghreb region of North Africa. It overlooks the Mediterranean Sea to the north and the Atlantic Ocean to the west, and has land borders with Algeria to A ...

('' Diplocaulus minumus''), lepospondyls lived from the

Early Carboniferous Early may refer to: History * The beginning or oldest part of a defined historical period, as opposed to middle or late periods, e.g.: ** Early Christianity ** Early modern Europe Places in the United States * Early, Iowa * Early, Texas * Ear ...

( Mississippian) to the

Early Permian 01 or '01 may refer to: * The year 2001, or any year ending with 01 * The month of January * 1 (number) Music * 01'' (Richard Müller album), 2001 * ''01'' (Son of Dave album), 2000 * ''01'' (Urban Zakapa album), 2011 * ''O1'' (Hiroyuki Sawan ...

and were geographically restricted to what is now

Europe Europe is a large peninsula conventionally considered a continent in its own right because of its great physical size and the weight of its history and traditions. Europe is also considered a Continent#Subcontinents, subcontinent of Eurasia ...

and

North America North America is a continent in the Northern Hemisphere and almost entirely within the Western Hemisphere. It is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South America and th ...

. 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 Lizards are a widespread group of squamate reptiles, with over 7,000 species, ranging across all continents except Antarctica, as well as most oceanic island chains. The group is paraphyletic since it excludes the snakes and Amphisbaenia altho ...

-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 Temnospondyli (from Greek τέμνειν, ''temnein'' 'to cut' and σπόνδυλος, ''spondylos'' 'vertebra') is a diverse order of small to giant tetrapods—often considered primitive amphibians—that flourished worldwide during the Carb ...

amphibians that coexisted with them in the Paleozoic. Lepospondyli was named in 1888 by

Karl Alfred von Zittel Karl Alfred Ritter von Zittel (25 September 1839 – 5 January 1904) was a German palaeontologist best known for his ''Handbuch der Palaeontologie'' (1876–1880). Biography Karl Alfred von Zittel was born in Bahlingen in the Grand Duchy o ...

, who coined the name to include some tetrapods from the

Paleozoic The Paleozoic (or Palaeozoic) Era is the earliest of three geologic eras of the Phanerozoic Eon. The name ''Paleozoic'' ( ;) was coined by the British geologist Adam Sedgwick in 1838 by combining the Greek words ''palaiós'' (, "old") and ...

that shared some specific characteristics in the

notochord In anatomy, the notochord is a flexible rod which is similar in structure to the stiffer cartilage. If a species has a notochord at any stage of its life cycle (along with 4 other features), it is, by definition, a chordate. The notochord consi ...

and

teeth A tooth ( : teeth) is a hard, calcified structure found in the jaws (or mouths) of many vertebrates and used to break down food. Some animals, particularly carnivores and omnivores, also use teeth to help with capturing or wounding prey, ...

. Lepospondyls have sometimes been considered to be either related or ancestral to modern amphibians or to Amniota (the clade containing reptiles and mammals). It has been suggested that the grouping is

polyphyletic A polyphyletic group is an assemblage of organisms or other evolving elements that is of mixed evolutionary origin. The term is often applied to groups that share similar features known as homoplasies, which are explained as a result of conver ...

, with aïstopods being primitive stem-tetrapods, while

recumbirostran 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 o ...

microsaurs are primitive reptiles.Jason D. Pardo, Matt Szostakiwskyj, Per E. Ahlberg & Jason S. Anderson (2017) Hidden morphological diversity among early tetrapods. Nature (advance online publication) doi:10.1038/nature22966>

Description

All lepospondyls are characterised by having simple, spool-shaped

vertebra The spinal column, a defining synapomorphy shared by nearly all vertebrates, Hagfish are believed to have secondarily lost their spinal column is a moderately flexible series of vertebrae (singular vertebra), each constituting a characteristi ...

e that did not

ossify Ossification (also called osteogenesis or bone mineralization) in bone remodeling is the process of laying down new bone material by cells named osteoblasts. It is synonymous with bone tissue formation. There are two processes resulting in th ...

from

cartilage Cartilage is a resilient and smooth type of connective tissue. In tetrapods, it covers and protects the ends of long bones at the joints as articular cartilage, and is a structural component of many body parts including the rib cage, the neck ...

, 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).Colbert 1969

Classification

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

salamander Salamanders are a group of amphibians typically characterized by their 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. All t ...

s (Urodela), but not the other modern amphibians. This view is no longer held and all modern amphibians (frogs, salamanders, and

caecilian Caecilians (; ) are a group of limbless, vermiform or serpentine amphibians. They mostly live hidden in the ground and in stream substrates, making them the least familiar order of amphibians. Caecilians are mostly distributed in the tropics o ...

s) are now grouped within the clade

Lissamphibia The Lissamphibia is a group of tetrapods that includes all modern amphibians. Lissamphibians consist of three living groups: the Salientia (frogs, toads, and their extinct relatives), the Caudata (salamanders, newts, and their extinct relativ ...

. For a long time, the Lepospondyli were considered one of the three subclasses of

Amphibia Amphibians are four-limbed and ectothermic vertebrates of the class Amphibia. All living amphibians belong to the group Lissamphibia. They inhabit a wide variety of habitats, with most species living within terrestrial, fossorial, arbo ...

, along with the Lissamphibia and the

Labyrinthodontia "Labyrinthodontia" ( Greek, 'maze-toothed') is an informal grouping of extinct predatory amphibians which were major components of ecosystems in the late Paleozoic and early Mesozoic eras (about 390 to 150 million years ago). Traditionally con ...

.Carroll 1988 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 (

) grouping with some members closely related to extinct stem tetrapod groups and others more closely related to modern amphibians or reptiles. 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. Many phylogenetic analyses since Carroll (1995) agreed with his interpretation, including Laurin & Reisz (1997), Anderson (2001), and Ruta ''et al.'' (2003). 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. 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 and a 2017 paper on ''

Lethiscus ''Lethiscus'' is the earliest known representative of the Aistopoda, a group of very specialised snake-like amphibians known from the early Carboniferous ( Mississippian). ''Lethiscus'' is known from only a single specimen from the Holkerian S ...

'' has Aïstopoda in the tetrapod stem based on their primitive braincase. 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.

Interrelationships

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 In taxonomy, a group is paraphyletic if it consists of the group's last common ancestor and most of its descendants, excluding a few monophyletic subgroups. The group is said to be paraphyletic ''with respect to'' the excluded subgroups. In ...

; rather than being a

monophyletic In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic gr ...

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 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 o ...

, 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. 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 A cladogram (from Greek ''clados'' "branch" and ''gramma'' "character") is a diagram used in cladistics to show relations among organisms. A cladogram is not, however, an evolutionary tree because it does not show how ancestors are related to ...

, simplified, is after an analysis of tetrapods and stem-tetrapods presented by Ruta ''et al.'' in 2003:

Position within Tetrapoda

The "lepospondyl hypothesis" of modern amphibian origins proposes that lissamphibians are

(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 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 ...

) 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. 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. However, some

phylogenetic In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups ...

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. Below is a cladogram from Ruta ''et al.'' (2003) that supports the "temnospondyl hypothesis", showing the position of Lepospondyli within crown group Tetrapoda:

Gallery

File:Tuditanus1DB.jpg, ''

Tuditanus ''Tuditanus'' is an extinct genus of tuditanid microsaur from the Carboniferous, ~ 306 Ma ago. It was of small size, reaching a length of about 24 cm. See also * Prehistoric amphibian * List of prehistoric amphibians This list of preh ...

'', a tuditanid "

microsaur Microsauria ("small lizards") 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 ...

" File:Pelodosotis1DB.jpg, '' Pelodosotis'', an ostodolepid "

" File:Pantylus1DB.jpg, '' Pantylus'', a pantylid "

" File:Hyloplesion.jpg, ''

Hyloplesion ''Hyloplesion'' is an extinct genus of microbrachomorph microsaur. It is the type and only genus within the family Hyloplesiontidae. Fossils have been found from the Czech Republic near the towns of Plzeň, Nýřany, and Třemošná, and ...

'', a "

" File:Brachydectes NT small.jpg, ''

Brachydectes ''Brachydectes'' is an extinct genus of lysorophian amphibian that lived from the Carboniferous. It had a very small head A head is the part of an organism which usually includes the ears, brain, forehead, cheeks, chin, eyes, nose, and mout ...

'', a lysorophian File:Adelospondylus2.jpg, ''

Adelospondylus ''Adelospondylus'' is an extinct lepospondyl amphibian Amphibians are four-limbed and ectothermic vertebrates of the class Amphibia. All living amphibians belong to the group Lissamphibia. They inhabit a wide variety of habitats, with ...

'', an adelospondyl File:Oestocephalus.jpg, ''

Oestocephalus ''Oestocephalus'' is an extinct genus of aïstopod Stegocephalian that lived during the Carboniferous period. Fossils have been found in the Czech Republic, and in Ohio and Illinois in the United States. It is the type genus of the family Oesto ...

'', an early

aïstopod Aistopoda (Greek for " avingnot-visible feet") 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 , to nearly in length. They fi ...

File:Phlegethontia NT small.jpg, '' Phlegethontia'', an

File:Diploceraspis12DB.jpg, '' Diploceraspis'', a diplocaulid " nectridean" File:Sauropleura.jpg, '' Sauropleura'', a urocordylid " nectridean"

References

* Benton, M. J. (2000), ''Vertebrate Paleontology'', 2nd ed. Blackwell Science Ltd * Benton, M. J. (2004), ''Vertebrate Paleontology'', 3rd ed. Blackwell Science Lt
Taxonomic hierarchy
* Colbert, E H. (1969), ''Evolution of the Vertebrates'', John Wiley & Sons Inc (2nd ed.) * Laurin, Michel (1996
Terrestrial Vertebrates - Stegocephalians: Tetrapods and other digit-bearing vertebrates
* Romer, A.S., (1966) ''Vertebrate Paleontology'' University of Chicago Press, 1933; 3rd ed. 1966.

External links

{{Taxonbar, from=Q133023 Mississippian first appearances Lopingian extinctions Fossil taxa described in 1888 Polyphyletic groups