Temporal range: 340–230Ma Descendant taxon Amniota survives to present
|Mounted skeleton of Diadectes, American Museum of Natural History|
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.
The informal variant of the name, "reptiliomorphs", is also occasionally used to refer to stem-amniotes, i.e. a grade of reptile-like labyrinthodont tetrapods that are more closely related to amniotes than they are to lissamphibians, but are not amniotes themselves; the name is used in this meaning e.g. by Ruta, Coates and Quicke (2003). An alternative name, "Anthracosauria" is also commonly used for the group but is confusingly also used for the more primitive grade of reptiliomorphs by Benton.
As the exact phylogenetic position of Lissamphibia within Tetrapoda remains uncertain, it also remains controversial which fossil tetrapods are more closely related to amniotes than to lissamphibians, and thus, which ones of them were reptiliomorphs in any meaning of the word.
The name Reptiliomorpha was coined by Professor Gunnar Säve-Söderbergh in 1934 to designate amniotes and various types of late Paleozoic tetrapods than were more closely related to amniotes than to living amphibians. In his view, the amphibians had evolved from fish twice, with one group composed of the ancestors of modern salamanders and the other, which Säve-Söderbergh referred to as Eutetrapoda, consisting of anurans (frogs), amniotes and their ancestors, with the origin of caecilians being uncertain. Säve-Söderbergh's Eutetrapoda consisted of two sister-groups: Batrachomorpha, containing anurans and their ancestors, and Reptiliomorpha, containing anthracosaurs and amniotes. Säve-Söderbergh subsequently added Seymouriamorpha to his Reptiliomorpha as well.
Alfred Sherwood Romer rejected Säve-Söderbergh's theory of a biphyletic amphibia and used the name Anthracosauria to describe the labyrinthodont lineage from which amniotes evolved. In 1970, the German paleontologist Alec Panchen took up Säve-Söderberghs name for this group as having priority, but Romer's terminology is still in use, e.g. by Carroll (1988 and 2002) and by Hildebrand & Goslow (2001). Some writers preferring phylogenetic nomenclature use Anthracosauria.
In 1956, Friedrich von Huene included both amphibians and anapsid reptiles in the Reptiliomorpha. This included the following orders: Anthracosauria, Seymouriamorpha, Microsauria, Diadectomorpha, Procolophonia, Pareiasauria, Captorhinidia, Testudinata.
Michael Benton (2000, 2004) made it the sister-clade to Lepospondyli, containing "anthracosaurs" (in the strict sense, i.e. Embolomeri), seymouriamorphs, diadectomorphs and amniotes. However, when considered in a Linnean framework, Reptiliomorpha is given the rank of superorder and includes only reptile-like amphibians, not their amniote descendants.
Several phylogenetic studies indicate that amniotes and diadectomorphs share more recent common ancestor with lepospondyls than with seymouriamorphs, Gephyrostegus and Embolomeri (e.g. Laurin and Reisz, 1997, 1999; Ruta, Coates and Quicke, 2003; Vallin and Laurin, 2004; Ruta and Coates, 2007). Lepospondyls are one of the groups of tetrapods suggested to be ancestors of living amphibians; as such, their potential close relationship to amniotes has important implications for the content of Reptiliomorpha. Assuming that lissamphibians aren't descended from lepospondyls but from a different group of tetrapods, e.g. from temnospondyls, it would mean that Lepospondyli belonged to Reptiliomorpha sensu Laurin (2001), as it would make them more closely related to amniotes than to lissamphibians. On the other hand, if lissamphibians are descended from lepospondyls, then not only Lepospondyli would have to be excluded from Reptiliomorpha, but seymouriamorphs, Gephyrostegus and Embolomeri would also have to be excluded from this group, as this would make them more distantly related to amniotes than living amphibians are. In that case, the clade Reptiliomorpha sensu Laurin would contain, apart from Amniota, only diadectomorphs and possibly also Solenodonsaurus.
Seymouriamorphs and diadectomorphs were land-based, reptile-like amphibians, while embolomeres were aquatic amphibians with long body and short limbs. Their anatomy falls between the mainly aquatic Devonian labyrinthodonts and the first reptiles. University of Bristol paleontologist Professor Michael J. Benton gives the following characteristics for the Reptiliomorpha (in which he includes embolomeres, seymouriamorphs and diadectomorphs):
- narrow premaxillae (less than half the skull width)
- vomers taper forward
- phalangeal formulae (number of joints in each toe) of foot 22.214.171.124.4–5
The groups traditionally assigned to Reptiliomorpha, i.e. embolomeres, seymouriamorphs and diadectomorphs, differed from the contemporary non-reptiliomorph labyrinthodonts in having a deeper and taller skull, but retained the primitive kinesis (loose attachment) between the skull roof and the cheek. The deeper skull allowed for laterally placed eyes, contrary to the dorsally placed eyes commonly found in amphibians. The skulls of the group are usually found with fine radiating grooves. The quadrate bone in the back of the skull held a deep otic notch, likely holding a spiracle rather than a tympanum.
The vertebrae showed the typical multi-element construction seen in labyrinthodonts. According to Benton, in the vertebrae of "anthracosaurs" (i.e. Embolomeri) the intercentrum and pleurocentrum may be of equal size, while in the vertebrae of seymouriamorphs the pleurocentrum is the dominant element and the intercentrum is reduced to a small wedge. The intercentrum gets further reduced in the vertebrae of amniotes, where it becomes a thin plate or disappears altogether. Unlike most labyrinthodonts, the body was moderately deep rather than flat, and the limbs were well-developed and ossified, indicating a predominantly terrestrial lifestyle except in secondarily aquatic groups. Each foot held 5 digits, the pattern seen in their amniote descendants. They did however lack the reptilian type of ankle bone that would have allowed the use of the feet as levers for propulsion rather than as holdfasts.
The general build was heavy in all forms, though otherwise very similar to that of early reptiles. The skin, at least in the more advanced forms probably had a water-tight epidermal horny overlay, similar to the one seen in today's reptiles, though they lacked horny claws. In chroniosuchians and some seymouriamorphans, like Discosauriscus, dermal scales are found in post-metamorphic specimens, indicating they may have had a "knobbly" if not scaly appearance.
Seymouriamorphs reproduced in amphibian fashion with aquatic eggs that hatched into larvae (tadpoles) with external gills; it is unknown how other tetrapods traditionally assigned to Reptiliomorpha reproduced.
During the Carboniferous and Permian periods, some tetrapods started to evolve towards a reptilian condition. Some of these tetrapods (e.g. Archeria, Eogyrinus) were elongate, eel-like aquatic forms with diminutive limbs, while others (e.g. Seymouria, Solenodonsaurus, Diadectes, Limnoscelis) were so reptile-like that until quite recently they actually had been considered to be true reptiles, and it is likely that to a modern observer they would have appeared as large to medium-sized, heavy-set lizards. Several groups however remained aquatic or semiaquatic. Some of the chroniosuchians show the build and presumably habits of modern crocodiles and were probably also similar to crocodylians in that they were river-side predators. While some other Chroniosuchians possessed elongated newt- or eel-like bodies. The two most terrestrially adapted groups were the medium sized insectivorous or carnivorous Seymouriamorpha and the mainly herbivorous Diadectomorpha, with many large forms. The latter group is in most analysis the closest relatives of the Amniotes.
From aquatic to terrestrial eggs
Their terrestrial life style combined with the need to return to the water to lay eggs hatching to larvae (tadpoles) led to a drive to abandon the larval stage and aquatic eggs. A possible reason may have been competition for breeding ponds, to exploit drier environments with less access to open water, or to avoid predation on tadpoles by fish, a problem still plaguing modern amphibians. Whatever the reason, the drive led to internal fertilization and direct development (completing the tadpole stage within the egg). A striking parallel can be seen in the frog family Leptodactylidae, which has a very diverse reproductive system, including foam nests, non-feeding terrestrial tadpoles and direct development. The Diadectomorphans generally being large animals would have had correspondingly large eggs, unable to survive on land.
Fully terrestrial life was achieved with the development of the amniote egg, where a number of membranous sacks protect the embryo and facilitate gas exchange between the egg and the atmosphere. The first to evolve was probably the allantois, a sack that develops from the gut/yolk-sack. This sack contains the embryo's nitrogenous waste (urea) during development, stopping it from poisoning the embryo. A very small allantois is found in modern amphibians. Later came the amnion surrounding the fetus proper, and the chorion, encompassing the amnion, allantois, and yolk-sack.
Origin of amniotes
Exactly where the border between reptile-like amphibians (non-amniote reptiliomorphs) and amniotes lies will probably never be known, as the reproductive structures involved fossilize poorly, but various small, advanced reptiliomorphs have been suggested as the first true amniotes, including Solenodonsaurus, Casineria and Westlothiana. Such small animals lay small eggs, 1 cm in diameter or less. Small eggs will have a small enough volume to surface ratio to be able to develop on land without the amnion and chorin actively effecting gas exchange, setting the stage for the evolution of true amniotic eggs. Although the first amniote probably appeared as early as the Middle Mississippian sub-epoch, non-amniote (or amphibian) reptiliomorphs continued to flourish alongside their amniote descendants for many millions of years. By the middle Permian the non-amniote terrestrial forms had died out, but several aquatic non-amniote groups continued to the end of the Permian, and in the case of the Chroniosuchids survived the end Permian mass extinction, only to die out at the end of the Early Triassic. Meanwhile, the single most successful daughter-clade of the reptiliomorphs, the amniotes, continued to flourish and to inherit the Earth.
- Laurin, M. (2001). "L'utilisation de la taxonomie phylogénétique en paléontologie: avantages et inconvénients". Biosystema 19: 197–211.
- Vallin, Grégoire; Laurin, Michel (2004). "Cranial morphology and affinities of Microbrachis, and a reappraisal of the phylogeny and lifestyle of the first amphibians". Journal of Vertebrate Paleontology 24 (1): 56–72. doi:10.1671/5.1.
- Ruta, M.; Coates, M.I.; and Quicke, D.L.J. (2003). "Early tetrapod relationships revisited". Biological Reviews 78 (2): 251–345. doi:10.1017/S1464793102006103. PMID 12803423.
- Benton, M. J. (2000), Vertebrate Paleontology, 2nd Ed. Blackwell Science Ltd 3rd ed. 2004 – see also taxonomic hierarchy of the vertebrates, according to Benton 2004
- Säve-Söderbergh, G. (1934). "Some points of view concerning the evolution of the vertebrates and the classification of this group". Arkiv för Zoologi 26A: 1–20.
- Säve-Söderbergh, G. (1935). "On the dermal bones of the head in labyrinthodont stegocephalians and primitive Reptilia with special reference to Eotriassic stegocephalians from East Greenland". Meddelelser Om Grønland 98 (3): 1–211.
- Panchen, A. L. (1970). Handbuch der Paläoherpetologie - Encyclopedia of Paleoherpetology Part 5a - Batrachosauria (Anthracosauria), Gustav Fischer Verlag - Stuttgart & Portland, 83 pp., ISBN 3-89937-021-X.
- Carroll, R. L., (1988): Vertebrate paleontology and evolution. W. H. Freeman and company, New York.
- Carroll, R. (2002): ""Palaeontology: Early land vertebrates". Nature 418: 35-36.
- Hildebrand, M.; Goslow, G. E., Jr (2001). Analysis of vertebrate structure. Principal ill. Viola Hildebrand. New York: Wiley. p. 429. ISBN 0-471-29505-1.
- Gauthier, J., Kluge, A.G., & Rowe, T. (1988): "The early evolution of the Amniota". In The phylogeny and classification of the tetrapods, no 1: amphibians, reptiles, birds. Edited by M.J. Benton. Clarendon Press, Oxford, pp. 103–155.
- Von Huene, F., (1956), Paläontologie und Phylogenie der niederen Tetrapoden, G. Fischer, Jena.
- Systema Naturae (2000) / Classification Superorder Reptiliomorpha
- Laurin, M.; and Reisz, R.R. (1997). "A new perspective on tetrapod phylogeny". In Sumida, S.S. and Martin, K.L.M. (eds.). Amniote Origins: Completing the Transition to Land. Academic Press. pp. 9–60. ISBN 0-12-676460-3, 9780126764604 Check
- Laurin, M.; and Reisz, R.R. (1999). "A new study of Solenodonsaurus janenschi, and a reconsideration of amniote origins and stegocephalian evolution". Canadian Journal of Earth Sciences 36 (8): 1239–1255. doi:10.1139/e99-036.
- Ruta, M.; and Coates, M.I. (2007). "Dates, nodes and character conflict: addressing the lissamphibian origin problem". Journal of Systematic Palaeontology 5 (1): 69–122. doi:10.1017/S1477201906002008.
- Hillary C. Maddin, Farish A. Jenkins Jr and Jason S. Anderson (2012). "The Braincase of Eocaecilia micropodia (Lissamphibia, Gymnophiona) and the Origin of Caecilians". PLoS ONE 7 (12): e50743. doi:10.1371/journal.pone.0050743.
- Palaeos Reptilomorpha
- Lombard, R. E. & Bolt, J. R. (1979): Evolution of the tetrapod ear: an analysis and reinterpretation. Biological Journal of the Linnean Society No 11: pp 19–76 Abstract
- Chapter 4: "The early tetrapods and amphibians." In: Benton, M. J. (2004), Vertebrate Paleontology, 3rd ed. Blackwell Science Ltd.
- Romer, A.S. & T.S. Parsons. 1977. The Vertebrate Body. 5th ed. Saunders, Philadelphia. (6th ed. 1985)
- Palaeos Reptilomorpha: Cotylosauria
- Romer, A.S. (1946). "The primitive reptile Limnoscelis restudied". American Journal of Science 244: 149–188.
- R. L. Paton, T. R. Smithson and J. A. Clack, "An amniote-like skeleton from the Early Carboniferous of Scotland", (abstract), Nature 398, 508-513 (8 April 1999)
- Maddin & al. (2009): The anatomy and development of the claws ofXenopus laevis (Lissamphibia: Anura) reveal alternate pathways of structural evolution in the integument of tetrapods. Journal of Anatomy, no 214 (4): pp 607-19 Abstract
- Laurin, M. (1996). "A Reevaluation of Ariekanerpeton, a Lower Permian Seymouriamorph (Vertebrata: Seymouriamorpha) from Tadzhikistan". Journal of Vertebrate Paleontology 16 (4): 653–665.
- Špinar, Z. V. (1952): Revision of some Morovian Discosauriscidae. Rozpravy ustrededniho Uštavu Geologickeho no 15, pp 1–160
- Laurin, M. (1996): Phylogeny of Stegocephalians, from the Tree of Life Web Project
- Duellman, W.E. & Trueb, L. (1994): Biology of amphibians. The Johns Hopkins University Press
- Carroll R.L. (1991): The origin of reptiles. In: Schultze H.-P., Trueb L., (ed) Origins of the higher groups of tetrapods — controversy and consensus. Ithaca: Cornell University Press, pp 331-353.