Labyrinthodontia

Labyrinthodontia* Fossil range: 365–100 Ma Descendant taxon Amniotes and Lissamphibians survives to present. PreЄ Є O S D C P T J K Pg N
Proterogyrinus, an Anthracosaur.
Scientific classification
Kingdom:	Animalia
Phylum:	Chordata
Subphylum:	Vertebrata
Class:	Amphibia sensu lato
Orders
Ordo Ichthyostegalia † Ordo Temnospondyli † (unranked) Batrachomorpha (ancestors of mordern amphibians) Superordo Reptiliomorpha † (ancestors of reptiles)

Labyrinthodont (Greek, "maze-toothed") is an obsolete term for any member of the extinct superorder (or subclass) (Labyrinthodontia) of amphibians, which constituted some of the dominant animals of Late Paleozoic and Early Mesozoic times (about 350 to 210 million years ago). The name describes the pattern of infolding of the dentine and enamel of the teeth, which are often the only part of the creatures that fossilize. They are also distinguished by a heavily armoured skull roof (and therefore often named "Stegocephalia"), and complex vertebrae, the structure of which is useful in older classifications of the group.

Anatomy

Cross-section of a laybrintodont tooth

The labyrinthodonts flourished for more than 150 million years. Particularly the early forms exhibited a lot of variation, yet there are still a few basic anatomical traits that make them fairly easy to recognise:

• Strongly folded tooth surface, involving infolding of the dentine and enamel of the teeth, so that a cross section resembles a classical labyrinth (or maze), hence the name of the group.
• Massive skull roof, with openings only for the nostrils, eyes and a parietal eye, similar to the structure of the anapsids. With the exception of the later more reptile-like forms, the skull was rather flat with copious dermal armour, accounting for the older term for the group: Stegocephalia.
• Complex vertebrae made of 4 pieces, an intercentrum, two pleurocentra, and a neural arch/spine. The relative sizes of these pieces distinguishes different groups of labyrinthodonts.

The Labyrinthodonts in life

Labyrinthodont.

The early labyrinthodonts were mostly aquatic, hunting in shallow water or weed filled tidal channels. They were short-legged and large headed, some could be up to four meters long. Their skulls were massive, and their jaws were lined with small, sharp, conical teeth. Also, there was a second row of teeth on the roof of the mouth. In their way of living labyrinthodonts were probably similar to fishes – they laid eggs in the water, where their larvae developed into mature animals.

Characteristically labyrinthodonts have vertebrae made of 4 pieces, an intercentrum, two pleurocentra, and a neural arch/spine. While able to support some weight, the labyrinthodonts would have been slow and clumsy on land. Some of the larger adults may have been confined to water. They also had special sense organs in the skin, that formed a system for perception of water fluctuations. Some of them possessed well developed gills and many seemingly had primitive lungs. They could breath atmospheric air; that was a great advantage for residents of warm shoals with low oxygen levels in the water. The air was inflated into the lungs by contractions of a special throat sac. Primitive members of all labyrinthodont groups were probably true water predators, and only advanced forms that arose independently in different groups and times, gained an amphibious, semi-aquatic mode of living. Their bulky skeleton and their short limbs suggest that the majority of the labyrinthodonts were slow walkers on land.

Evolution

Platyoposaurus

The Labyrinthodontia evolved from a bony fish group: the fleshy-finned rhipidistia. Nowadays only a few living representatives of these fish remains: two species of coelacanth and six species of lungfish.

The most diverse group of the labyrinthodonts was the Batrachomorpha. Though these animals looked more like crocodiles, they most probably gave rise to the order Anura, the amphibians without tails, which include, in particular, the modern frogs. Batrachomorphs appeared in the Late Devonian, but they had worldwide distribution in the continental shallow basins of the Permian (Platyoposaurus, Melosaurus) and Triassic Periods (Thoosuchus, Benthosuchus, Eryosuchus). Some batrachomorphs existed until the end of the Cretaceous.

Classification

Pteroplax, an Anthracosaurian.

The term Labyrinthodont was coined by Hermann Burmeister in reference to the tooth structure.^[1] Labyrinthodontia was first used as a systematic term by Richard Owen in 1860, and assigned to Amphibia the following year.^[2] It was ranked as superorder by Romer, A. S. in 1947.^[3] The traditional classification (e.g. Romer 1966, also repeated in Colbert 1969, Daly 1973 and Carroll 1988) has three orders:^{[4][5] [6] [7]}

• Ichthyostegalia (primitive ancestral forms (e.g. Ichthyostega) – Late Devonian only). While undoubtly amphibians on anatomy and habit, the Ichthyostegalia are ancestral to all tetrapods, and are not amphibians in the cladistic sense.
• Temnospondyli (common, small to large, flat-headed forms with either strong or secondarily weak vertebrae and limbs – mainly Carboniferous to Triassic e.g. Eryops from the early Permian is a well-known genus, More recently fossil Jurassic and Cretaceous temnospondyls have been found. Originally considered ancestral to Anura (frogs), may or may not be ancestral to all modern amphibians. Temnospondyls are the only "Labyrinthodonts" currently considered to be "true amphibians" in that they are more closely related to modern Lissamphibia than to other tetrapodes.
• Anthracosauria (deep skulls, strong vertebrae but weak limbs, evolving towards and ancestral to reptiles – Carboniferous and Permian – e.g. Seymouria). The Anthracosauria are thought the direct ancestral to the early reptiles, and thus separate from modern ("true") amphibians.

Systematic overview of Labyrinthodontia (Stegocephalia)

Suggested evolutionary tree of Labyrinthodontia, from Colbert 1969 and Caroll 1997.^[8][9]

From Crossopterygii (lobe-fined fish)
   `- Eusthenopteron (advanced lobe-fined fish)
      `- Panderichthys (lobe-fined fish with limb-like fins)
         `- Tiktaalik (transitional fish/amphibian)
            `- Acanthostega (early amphibian with fishlike gills)
               `- Ichthyostega (early amphibian)
                  `- Crassigyrinus (early amphibian)
                     |- Loxommatidae (eel-like primitive temnospondyles)
                     |  `-Temnospondyls (large, flat-headed stegocephalians)
                     `- Anthracosaurs (reptile-like amphibians)
                         |- Seymouriamorphs (advanced reptile-like amphibians)
                         |  |- Westlothiana (small amphibian or possibly early reptile)
                         |  `+ Diadectomorphs (earliest reptiles or sister groups of reptiles)
                         |    `- amniotes (i.e. first reptiles)
                         `- Batrachomorpha (modern amphibians and their extinct relatives)
                             `- Lepospondyls (small stegocephalians)
                                 `?- Lissamphibia (modern amphibians)

A good summary (with diagram) of characteristics and main evolutionary trends of the above three orders is given in Colbert 1969 pp.102–103, but see Kent & Miller (1997) for an alternative tree.^[10]

While Labyrinthodontia is a traditional designation and a name commonly found in textbooks, the name has fallen out of favor in recent taxonomies as paraphyletic: the group does not include all the descendants of the cenancestor (most recent common ancestor). The groups that have usually been placed within Labyrinthodontia are currently variously classified as basal tetrapods, non-amniote Reptiliomorpha and as a monophyletic or paraphyletic Temnospondyli, according to cladistic analysis. This reflects on the importance of ascertaining lineage and ancestral-descendant relatedness in modern-day cladistics. The name does linger as a handy reference for the early amphibian tetrapods, and as a anatomical description.

See also

• Tetrapod
• Temnospondyli
• Batrachosauria
• Reptiliomorpha
• Amphibian
• Prehistoric amphibian
• Ichthyostega
• Eryops
• Prehistoric life

References

External links

Wikimedia Commons has media related to: Labyrinthodontia

• Palaeos.com Cladogram of Reptiliomorpha
• Subclass Labyrinthodontia
• Comparative Vertebrate Anatomy