Fossil range: Carboniferous (Middle Mississippian) to Recent
A baby tortoise emerges from an amniotic egg.
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Subphylum: Vertebrata
Infraphylum: Gnathostomata
Superclass: Tetrapoda
(unranked) Amniota
Haeckel, 1866
Living subgroups

Class Synapsida
   Class Mammalia (mammals)
Class Sauropsida
       Testudines (turtles)
          Squamata (lizards & snakes)
          Sphenodontida (tuatara)
          Crocodilia (crocodiles)
          Class Aves (birds)

The amniotes are a taxon of tetrapod vertebrates that include the Synapsida (mammals) and Sauropsida (reptiles and dinosaurs, including birds). They are defined 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 [1]. The unique embryonic features of amniotes may reflect specializations of eggs to survive drier environments, or the massive size and yolk content of eggs designed for direct development to a larger size.

File:Anatomy of an amiotic egg.svg

Features of amniotes designed for survival on land include a sturdy but porous leathery or hard eggshell, and an allantois designed 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 first amniotes, such as Casineria kiddi, which lived about 340 million years ago, resembled small lizards. Their eggs were small and covered with a membrane, not a hard shell like most modern amniote eggs. 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 only became possible 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. One can assume the ancestors of the amniotes laid their eggs in moist places, as such modest-sized animals wouldn't have too many difficulties in finding depressions under fallen logs or other suitable places in the ancient forests, and dry conditions were probably not the main reason why the soft shell emerged.

In fish and amphibians there is only one inner membrane, also called an embryonic membrane. In amniotes the inner anatomy of the egg has evolved further, new structures have developed to take care of the gas exchanges between the embryo and the atmosphere, as well as dealing with the waste problems. To grow a thicker and tougher shell there were no other alternatives than finding new ways to supply the embryo with oxygen, as diffusion alone wouldn't be enough any more. After the egg had gotten these structures, further sophistication of them allowed the amniotes to lay much bigger eggs in much drier habitats. Bigger eggs meant bigger offspring, and bigger adults meant bigger eggs, which meant the amniotes had gotten the opportunity to grow bigger than their ancestors. But real growth was not possible until they stopped relying on small invertebrates as their main food source, and started to eat plants or other vertebrates, or returned to the water. New habits and heavier bodies meant further evolution for the amniotes, both in behavior and anatomy.

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 the eye. In anapsids there are none, in synapsids there is one, and in most diapsids there are two.

The skeletal remains of amniotes can be identified by their having at least two pairs of sacral ribs and an astragalus bone in the ankle.


(see sub-pages for in depth classification for each group).

External links

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