Reptile
Reptiles, as commonly defined, are tetrapod vertebrate animals with an ectothermic metabolism and amniotic development. Reptiles traditionally comprise four orders: Testudines, Crocodilia, Squamata and Rhynchocephalia, with about 12,000 extant species listed in the Reptile Database. The study of the traditional reptile orders, customarily in combination with the study of modern amphibians, is called herpetology.
Reptiles have been subject to several conflicting taxonomic definitions. In classical evolutionary taxonomy, reptiles are gathered together under the class Reptilia, which corresponds to common usage. Modern cladistic taxonomy regards that group as paraphyletic, since genetic and paleontological evidence has determined that birds are the only surviving group of Dinosauria, a major clade of diapsids that are more closely related to crocodilians than to other living reptiles, and thus birds are nested among reptiles from a phylogenetic perspective. Many cladistic systems therefore redefine Reptilia as a clade including birds, though the precise definition of this clade varies between authors. A similar concept is clade Sauropsida, which refers to all amniotes more closely related to modern reptiles than to mammals.
The earliest known members of the reptile lineage appeared during the late Carboniferous period, having evolved from advanced reptiliomorph tetrapods which became increasingly adapted to life on dry land. Genetic and fossil data argues that the two largest lineages of reptiles, Archosauromorpha and Lepidosauromorpha, diverged during the Permian period. In addition to the living reptiles, there are many diverse groups that are now extinct, in some cases due to mass extinction events. In particular, the Cretaceous–Paleogene extinction event wiped out the pterosaurs, plesiosaurs, and all non-avian dinosaurs alongside many species of crocodyliforms and squamates. Modern non-bird reptiles inhabit all the continents except Antarctica.
Extant reptiles range in size from the tiny Jaragua dwarf gecko, which only grow up to ; to the saltwater crocodile, which can reach over in length and weigh over. As tetrapods, reptiles generally have four limbs or, in the case of snakes and legless lizards, are descended from four-limbed ancestors but have lost the limbs through evolution. Unlike the anamniotic amphibians, reptiles do not rely on waterbodies for reproduction and have no aquatic larval stage. Most reptiles are oviparous with shelled eggs, although several species of squamates are viviparous, as were some extinct marine reptile clades. As amniotes, reptile eggs have extraembryonic membranes that retain water and facilitate biochemical exchange with the external environment, allowing reptiles to reproduce on dry land, even in extreme, arid habitats. Viviparous species have the eggs developing and hatching inside the mother's body, usually by internal incubation, although some species can nourish the eggs as fetuses through various forms of placenta analogs, with some providing initial parental care for their hatchlings.
Classification
Classical taxonomy and research
In the 13th century, the category of reptile was recognized in Europe as consisting of a miscellany of egg-laying creatures, including "snakes, various fantastic monsters, lizards, assorted amphibians, and worms", as recorded by Beauvais in his Mirror of Nature.In the 18th century, the reptiles were, from the outset of classification, grouped with the amphibians. Linnaeus, working from species-poor Sweden, where the common adder and grass snake are often found hunting in water, included all reptiles and amphibians in class Amphibia in the 10th edition of Systema Naturae| in his Systema Naturæ.
The terms reptile and amphibian were largely interchangeable, reptile being preferred by the French. J.N. Laurenti was the first to formally use the term Reptilia for an expanded selection of reptiles and amphibians basically similar to that of Linnaeus. Today, the two groups are still commonly treated under the single heading herpetology.
File:MosasaurDiscovery.jpg|thumb|left|"Antediluvian monster", a Mosasaurus discovered in a Maastricht limestone quarry, 1770
It was not until the beginning of the 19th century that it became clear that reptiles and amphibians are, in fact, quite different animals, and P.A. Latreille erected the class Batracia for the latter, dividing the tetrapods into the four familiar classes of reptiles, amphibians, birds, and mammals. The British anatomist T.H. Huxley made Latreille's definition popular and, together with Richard Owen, expanded Reptilia to include the various fossil "antediluvian monsters", including dinosaurs and the mammal-like Dicynodon he helped describe. This was not the only possible classification scheme: In the Hunterian lectures delivered at the Royal College of Surgeons in 1863, Huxley grouped the vertebrates into mammals, sauroids, and ichthyoids. He subsequently proposed the names of Sauropsida and Ichthyopsida for the latter two groups. In 1866, Haeckel demonstrated that vertebrates could be divided based on their reproductive strategies, and that reptiles, birds, and mammals were united by the amniotic egg.
The terms Sauropsida and Theropsida were used again in 1916 by E.S. Goodrich to distinguish between lizards, birds, and their relatives on the one hand and mammals and their extinct relatives on the other. Goodrich supported this division by the nature of the hearts and blood vessels in each group, and other features, such as the structure of the forebrain. According to Goodrich, both lineages evolved from an earlier stem group, Protosauria in which he included some animals today considered reptile-like amphibians, as well as early reptiles.
In 1956, D.M.S. Watson observed that the first two groups diverged very early in reptilian history, so he divided Goodrich's Protosauria between them. He also reinterpreted Sauropsida and Theropsida to exclude birds and mammals, respectively. Thus his Sauropsida included Procolophonia, Eosuchia, Millerosauria, Chelonia, Squamata, Rhynchocephalia, Crocodilia, "thecodonts", non-avian dinosaurs, pterosaurs, ichthyosaurs, and sauropterygians.
In the late 19th century, a number of definitions of Reptilia were offered. The biological traits listed by Lydekker in 1896, for example, include a single occipital condyle, a jaw joint formed by the quadrate and articular bones, and certain characteristics of the vertebrae. The animals singled out by these formulations, the amniotes other than the mammals and the birds, are still those considered reptiles today.
File:Captorhinus aguti p.jpg|thumb|right|The first reptiles had an anapsid type of skull roof, as seen in the Permian genus Captorhinus
The synapsid/sauropsid division supplemented another approach, one that split the reptiles into four subclasses based on the number and position of temporal fenestrae, openings in the sides of the skull behind the eyes. This classification was initiated by Henry Fairfield Osborn and elaborated and made popular by Romer's classic Vertebrate Paleontology. Those four subclasses were:
- Anapsida – no fenestrae – cotylosaurs and chelonia
- Synapsida – one low fenestra – pelycosaurs and therapsids
- Euryapsida – one high fenestra – protorosaurs and the marine sauropterygians and ichthyosaurs, the latter called Parapsida in Osborn's work.
- Diapsida – two fenestrae – most reptiles, including lizards, snakes, crocodilians, dinosaurs and pterosaurs.
Phylogenetics and modern definition
By the early 21st century, vertebrate paleontologists were beginning to adopt phylogenetic taxonomy, in which all groups are defined in such a way as to be monophyletic; that is, groups which include all descendants of a particular ancestor. The reptiles as historically defined are paraphyletic, since they exclude both birds and mammals. These respectively evolved from dinosaurs and from early therapsids, both of which were traditionally called "reptiles". Birds are more closely related to crocodilians than the latter are to the rest of extant reptiles. Colin Tudge wrote:Mammals are a clade, and therefore the cladists are happy to acknowledge the traditional taxon Mammalia; and birds, too, are a clade, universally ascribed to the formal taxon Aves. Mammalia and Aves are, in fact, subclades within the grand clade of the Amniota. But the traditional class Reptilia is not a clade. It is just a section of the clade Amniota: The section that is left after the Mammalia and Aves have been hived off. It cannot be defined by synapomorphies, as is the proper way. Instead, it is defined by a combination of the features it has and the features it lacks: reptiles are the amniotes that lack fur or feathers. At best, the cladists suggest, we could say that the traditional Reptilia are 'non-avian, non-mammalian amniotes'.
Despite the early proposals for replacing the paraphyletic Reptilia with a monophyletic Sauropsida, which includes birds, that term was never adopted widely or, when it was, was not applied consistently.
File:Bearded Dragon Skeleton.jpg|thumb|Bearded dragon skeleton on display at the Museum of Osteology
When Sauropsida was used, it often had the same content or even the same definition as Reptilia. In 1988, Jacques Gauthier proposed a cladistic definition of Reptilia as a monophyletic node-based crown group containing turtles, lizards and snakes, crocodilians, and birds, their common ancestor and all its descendants. While Gauthier's definition was close to the modern consensus, nonetheless, it became considered inadequate because the actual relationship of turtles to other reptiles was not yet well understood at this time. Major revisions since have included the reassignment of synapsids as non-reptiles, and classification of turtles as diapsids. Gauthier 1994 and Laurin and Reisz 1995's definition of Sauropsida defined the scope of the group as distinct and broader than that of Reptilia, encompassing Mesosauridae as well as Reptilia sensu stricto.
A variety of other definitions were proposed by other scientists in the years following Gauthier's paper. The first such new definition, which attempted to adhere to the standards of the PhyloCode, was published by Modesto and Anderson in 2004. Modesto and Anderson reviewed the many previous definitions and proposed a modified definition, which they intended to retain most traditional content of the group while keeping it stable and monophyletic. They defined Reptilia as all amniotes closer to Lacerta agilis and Crocodylus niloticus than to Homo sapiens. This stem-based definition is equivalent to the more common definition of Sauropsida, which Modesto and Anderson synonymized with Reptilia, since the latter is better known and more frequently used. Unlike most previous definitions of Reptilia, however, Modesto and Anderson's definition includes birds, as they are within the clade that includes both lizards and crocodiles.