Amniota

Amniote Fossil range: Mississippian (Carboniferous)–Recent, 340-0 Ma
Scientific classification
Infraphylum	Gnathostomata
Superclass	Tetrapoda
Family (Unranked)	Amniota Haeckel, 1866
Clades	Synapsida Sauropsida

The amniotes are a group of tetrapod vertebrates that have a terrestrially adapted egg. They include mammals, birds and reptiles, 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 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, which resembled small lizards, probably evolved 340 million years ago (Casineria). Their eggs could survive out of the water, allowing amniotes to branch out into drier environments. The eggs could also "breathe" and cope with waste, allowing the eggs and the amniotes themselves to evolve into larger forms. The amniotes spread across the globe and became the dominant land vertebrates. Very early in their history the amniotes split into two main lines. The oldest known fossil synapsid is Protoclepsydrops from about 320 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).

Description

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 evolved for direct development to a larger size.

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 first amniotes, such as Casineria kiddi, which lived about 334 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 and 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. In order to grow a thicker and tougher shell new ways to supply the embryo with oxygen had to be developed as diffusion alone was not enough. After the egg had developed these structures, further sophistication allowed the amniotes to lay much bigger eggs in much drier habitats. Bigger eggs allowed for bigger offspring and bigger adults could produce bigger eggs, which meant that the amniotes developed the opportunity to grow bigger than their ancestors. Real growth was not possible however, 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 (turtles) there are none, in synapsids (mammals and their extinct relatives) there is one, and in most diapsids (non-anapsid reptiles, dinosaurs, and birds) there are two.

The skeletal remains of amniotes can be identified 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 do not fossilize, and the status of fossil forms has to be interfered from other traits. Thus Jacques Gauthier and colleagues redefined Amniota in 1988, as "the most recent common ancestor of extant mammals and reptiles, and all its descendants". Gauthiers definition being node-based, the group under his definition has a slightly different content than than the group defined as biological amniotes (apomorphy-based clade).

Traditional classification

Classifications of the amniotes have traditionally recognised three classes, based on major traits and physiology:

• Class Reptilia (reptiles)
  • Subclass Anapsida ("proto-reptiles", possibly including turtles)
  • Subclass Synapsida (mammal-like reptiles, progenitors of mammals)
  • Subclass Diapsida (majority of reptiles, progenitors of birds)
• Class Aves (birds)
  • Subclass Neornithes (all modern birds, several extinct subclasses recognised)
• Class Mammalia (mammals)
  • Subclass Monotremata (egg-laying mammals)
  • Subclass Theria (marsupials and placental 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, that is, it has given rise to two other classes not included in Reptilia.

Phylogenetic classification

With the advent of cladistics, some researchers have attempted to establish new classes, based on phylogeny, but disregarding the physiological and anatomical unity of the groups. One such classification, by Michael Benton, is presented in simplified form below.

• Series Amniota ("reptiles")
  • Class Synapsida - includes non-mammalian synapsids (formerly mammal-like reptiles)
    • *Order Pelycosauria †
    • Order Therapsida
      • Class Mammalia - Mammals
  • Class Sauropsida
    • Subclass Anapsida
      • Order Testudines - Turtles
    • Subclass Diapsida
      • Order Araeoscelidia
      • Order Younginiformes
      • Infraclass Ichthyosauria †
      • Infraclass Lepidosauromorpha
        • Superorder Sauropterygia
          • Order Placodontia †
          • Order Nothosauroidea †
          • Order Plesiosauria †
        • Superorder Lepidosauria
          • Order Sphenodontida - Tuatara
          • Order Squamata - Lizards & snakes
      • Infraclass Archosauromorpha
        • • • Order Prolacertiformes †
            • Division Archosauria
              • Subdivision Crurotarsi
                • Order Crocodylia - Crocodilians
              • Subdivision Avemetatarsalia
                • Order Pterosauria †
                • Superorder Dinosauria
                  • Order Ornithischia †
                  • Order Saurischia
                    • Class Aves - Birds

Cladogram of Amniotes

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 defines under Gauthier's definition.

Amniota	Synapsida Sauropsida Mesosauridae Reptilia Parareptilia Millerettidae unnamed Pareiasauria unnamed Procolophonoidea Testudines (turtles, tortoises, and terrapins) Eureptilia Captorhinidae Romeriida Protorothyrididae* Diapsida (lizards, snakes, crocodiles, birds, etc.)

External links

• Out of the Swamps