Temporal range: Pennsylvanian–Holocene, 312–0Ma
|A baby tortoise emerges from an amniotic egg|
The amniotes are a group of tetrapods (descendants of four-limbed and backboned animals) that have an egg equipped with an amnios, an adaptation to lay eggs on land rather than in water as anamniotes (including frogs) typically do. They include synapsids (mammals along with their extinct kin) and sauropsids (reptiles including birds), as well as their fossil ancestors. Amniote embryos, whether laid as eggs or carried by the female, are protected and aided by several extensive membranes. In eutherian mammals (such as humans), these membranes include the amniotic sac that surrounds the fetus. These embryonic membranes, and the lack of a larval stage, distinguish amniotes from tetrapod amphibians.
The first amniotes (referred to as "basal amniotes") resembled small lizards and evolved from the amphibian reptiliomorphs about 312 million years ago, in the Carboniferous geologic period. Their eggs could survive out of the water, allowing amniotes to branch out into drier environments. The eggs could also "breathe" and cope with wastes, allowing the eggs and the amniotes themselves to evolve into larger forms.
The amniotic egg represents a critical divergence within the vertebrates, one enabled to reproduce on dry land—free of the need to return to water for reproduction as required of the amphibians. From this point the amniotes spread across the globe, eventually to become the dominant land vertebrates.
Very early in the evolutionary history of amniotes, basal amniotes diverged into two main lines, the synapsids and the sauropsids, both of which persist into the modern era. The oldest known fossil synapsid is Protoclepsydrops from about 312 million years ago, while the oldest known sauropsid is probably Paleothyris, in the order Captorhinida, from the Middle Pennsylvanian epoch (ca. 306–312 million years ago).
Amniotes can be characterized in part by embryonic development that includes the formation of several extensive membranes, the amnion, chorion, and allantois. Amniotes develop directly into a (typically) terrestrial form with limbs and a thick stratified epithelium, rather than first entering a feeding larval tadpole stage followed by metamorphosis as in amphibians. In amniotes, the transition from a two-layered periderm to cornified epithelium is triggered by thyroid hormone during embryonic development, rather than metamorphosis. The unique embryonic features of amniotes may reflect specializations of eggs to survive drier environments; or the massive size and yolk content of eggs may have evolved to allow direct development of the embryo to a larger size.
Adaptions for a terrestrial life
Features of amniotes evolved for survival on land include a sturdy but porous leathery or hard eggshell and an allantois evolved to facilitate respiration while providing a reservoir for disposal of wastes. Their kidneys and large intestines are also well-suited to water retention. Most mammals do not lay eggs, but corresponding structures may be found inside the placenta.
The ancestors of true amniotes, such as Casineria kiddi, which lived about 340 million years ago, evolved from amphibian reptiliomorphs and resembled small lizards. At the late Devonian mass extinction 360 million years ago, all known tetrapods were essentially aquatic and fish-like. Because the reptiliomorphs were already established 20 million years later when all their fishlike relatives were extinct, it appears they separated from the other tetrapods somewhere during Romer's gap, when the adult tetrapods became fully terrestrial (some forms would later become secondary aquatic). The first amniotic eggs were small and covered with a leathery membrane, not a hard shell like those of birds or crocodiles. Although some modern amphibians lay eggs on land, with or without significant protection, they all lack advanced traits like an amnion. This kind of egg became possible only with internal fertilization. The outer membrane, a soft shell, evolved as a protection against the harsher environments on land, as species evolved to lay their eggs on land where they were safer than in the water. The ancestors of the amniotes probably laid their eggs in moist places, as such modest-sized animals would not have difficulty finding depressions under fallen logs or other suitable places in the ancient forests; and dry conditions were probably not the main reason the soft shell emerged. Indeed, many modern day amniotes are dependent on moisture to keep their eggs from desiccating.
The egg membranes
In fish and amphibians there is only one inner membrane, the embryonic membrane. The amniote egg evolved new internal structures to accommodate gas exchange between the embryo and the atmosphere and to deal with wastes. A thicker and tougher shell necessitated new ways to supply the embryo with oxygen, as diffusion alone was not enough. After these structures developed, further adaption allowed amniotes to lay bigger eggs and to lay them in drier habitats. Larger eggs allowed the evolution of larger offspring and consequently larger adults. Further growth, however, was limited by reliance on small invertebrates as the main food source. Amniotes soon expanded their diets to include plants and larger animals, both vertebrate and invertebrate, permitting larger body size. New habits and heavier bodies resulted in further evolution for the amniotes, both in anatomy and in behavior.
While the early amniotes resembled their amphibian ancestors in many respects, a key difference was the lack of an otic notch at the back margin of the skull roof. In their ancestors, this notch held a spiracle, an unnecessary structure in an animal without an aquatic larval stage. There are three main lines of amniotes, which may be distinguished by the structure of the skull and in particular the number of temporal fenestrae (openings) behind each eye. In anapsids, the ancestral condition, there are none, in synapsids (mammals and their extinct relatives) there is one, and in most diapsids (including birds, crocodilians, squamates, and tuataras), there are two. Traditionally, turtles were classified as anapsids because they lack fenestrae, but molecular testing places them in the diapsid line of descent.
Post cranial remains of amniotes can be identified from their Labyrinthodont ancestors by their having at least two pairs of sacral ribs, a sternum in the pectoral girdle (some amniotes have lost it) and an astragalus bone in the ankle.
Definition and classification
Amniota was first formally described by embryologist Ernst Haeckel in 1866 on the presence of the amnion, hence the name. A problem with this definition is that the trait (apomorphy) in question does not fossilize, and the status of fossil forms has to be inferred from other traits.
- Class Reptilia (reptiles)
- Class Aves (birds)
- Subclass Archaeornithes (reptile-like birds, progenitors of all other birds)
- Subclass Enantiornithes (early birds with an alternative shoulder joint)
- Subclass Hesperornithes (toothed aquatic flightless birds)
- Subclass Ichthyornithes (toothed, but otherwise modern birds)
- Subclass Neornithes (all living birds)
- Class Mammalia (mammals)
This rather orderly scheme is the one most commonly found in popular and basic scientific works. It has come under critique from cladistics, as the class Reptilia is paraphyletic—it has given rise to two other classes not included in Reptilia.
Classification into monophyletic taxa
- Series Amniota
- Class Synapsida – includes mammal-like reptiles
- Class Sauropsida
- Subclass Anapsida
- Subclass Diapsida
- Order Araeoscelidia †
- Order Younginiformes †
- Infraclass Ichthyosauria †
- Infraclass Lepidosauromorpha
- Infraclass Archosauromorpha
- Order Prolacertiformes †
- Division Archosauria
- Subdivision Crurotarsi
- Subdivision Avemetatarsalia
With the advent of cladistics, other researchers have attempted to establish new classes, based on phylogeny, but disregarding the physiological and anatomical unity of the groups. Unlike Benton, for example, Jacques Gauthier and colleagues forwarded a definition of Amniota in 1988 as "the most recent common ancestor of extant mammals and reptiles, and all its descendants". As Gauthier makes use of a crown group definition, Amniota has a slightly different content than the biological amniotes as defined by an apomorphy.
The cladogram presented here illustrates the phylogeny (family tree) of amniotes, and follows a simplified version of the relationships found by Laurin & Reisz (1995). The cladogram covers the group as defined under Gauthier's definition.
The inclusion of Testudines within Parareptilia is controversial. More recent morphological phylogenetic studies placed turtles firmly within diapsids. All molecular studies have strongly upheld the placement of turtles within diapsids; some place turtles within Archosauria, or, more commonly, as a sister group to extant archosaurs, though an analysis conducted by Lyson et al. (2012) recovered turtles as the sister group of lepidosaurs instead.
- Benton, Michael J. (1997). Vertebrate Palaeontology. London: Chapman & Hall. pp. 105–109. ISBN 0-412-73810-4.
- Benton M.J. and Donoghue P.C.J. 2006. Palaeontological evidence to date the tree of life. Molecular biology and evolution. 24(1): 26–53. 
- Alexander M. Schreiber and Donald D. Brown *. "Tadpole skin dies autonomously in response to thyroid hormone at metamorphosis". Pnas.org. Retrieved 2011-04-25.
- Research project: The Mid-Palaeozoic biotic crisis: Setting the trajectory of Tetrapod evolution
- Stewart J. R. (1997): Morphology and evolution of the egg of oviparous amniotes. In: S. Sumida and K. Martin (ed.) Amniote Origins-Completing the Transition to Land (1): 291–326. London: Academic Press.
- Cunningham, B.; Huene, E. (Jul–Aug 1938). "Further Studies on Water Absorption by Reptile Eggs". The American Naturalist 72 (741): 380–385. doi:10.1086/280791. JSTOR 2457547.
- 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
- Gauthier, J., Kluge, A.G. and Rowe, T. (1988). "The early evolution of the Amniota." Pp. 103–155 in Benton, M.J. (ed.), The phylogeny and classification of the tetrapods, Volume 1: amphibians, reptiles, birds. Oxford: Clarendon Press.
- Romer, A.S. & Parsons, T.S. (1985): The Vertebrate Body. (6th ed.) Saunders, Philadelphia.
- Carroll, R. L. (1988), Vertebrate Paleontology and Evolution, WH Freeman & Co.
- Hildebrand, M. & G. E. Goslow, Jr. Principal ill. Viola Hildebrand. (2001). Analysis of vertebrate structure. New York: Wiley. p. 429. ISBN 0-471-29505-1.
- Colbert, E.H. & Morales, M. (2001): Colbert's Evolution of the Vertebrates: A History of the Backboned Animals Through Time. 4th edition. John Wiley & Sons, Inc, New York — ISBN 978-0-471-38461-8.
- *Hope, S. (2002) The Mesozoic record of Neornithes (modern birds). In: Chiappe, L.M. and Witmer, L.M. (eds.): Mesozoic Birds: Above the Heads of Dinosaurs: 339–388. University of California Press, Berkeley. ISBN 0-520-20094-2
- Benton, M.J. (2004). Vertebrate Paleontology. Blackwell Publishers. xii–452. ISBN 0-632-05614-2.
- Lee, M.S.Y. & Spencer, P.S. (1997): Crown clades, key characters and taxonomic stability: when is an amniote not an amniote? In: Sumida S.S. & Martin K.L.M. (eds.) Amniote Origins: completing the transition to land. Academic Press, pp 61–84. Google books
- Laurin, M. and Reisz, R.R. (1995). "A reevaluation of early amniote phylogeny." Zoological Journal of the Linnean Society, 113: 165–223.
- Rieppel & DeBraga 1996
- Müller 2004
- Mannen & Li 1999
- Zardoya & Meyer 1998
- Iwabe et al. 2004
- Roos, Aggarwal & Janke 2007
- Katsu et al. 2010
- Lyson et al. 2012
|This article needs additional citations for verification. (January 2009)|