Reptiles

Reptiles, or members of the class Reptilia, are air-breathing, cold-blooded amniotes that have skin covered in scales or scutes as opposed to hair or feathers. They are tetrapods (having or having descended from vertebrates with four limbs) and lay amniote eggs, whose embryos are surrounded by the amnion membrane. Modern reptiles inhabit every continent with the exception of Antarctica, and four living orders are currently recognized:

• Crocodilia (crocodiles, gavials, caimans, and alligators): 23 species
• Sphenodontia (tuatara from New Zealand): 2 species
• Squamata (lizards, snakes, and amphisbaenids ("worm-lizards"): approximately 7,900 species
• Testudines (turtles, tortoises, and terrapins): approximately 300 species

The majority of reptile species are oviparous (egg-laying) although certain species of squamates are capable of giving live birth. This is achieved, either through ovoviviparity (egg retention), or viviparity (offspring born without use of calcified eggs). Many of the viviparous species feed their fetuses through various forms of placenta analogous to those of mammals with some providing initial care for their hatchlings. Extant reptiles range in size from a tiny gecko, Sphaerodactylus ariasae, that grows to only 1.6 cm (0.6 in), to the saltwater crocodile that may reach 6 m in length and weigh over 1,000 kg. The science dealing with reptiles is called herpetology.

Classification

History of classification

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 "III - Amphibia" in his Systema Naturae. The terms "reptile" and "amphibian" were largely interchangeable, "reptiles" being preferred by the French. Josephus Nicolaus Laurenti were the first to formally use the term "Reptilia" for an expanded, though basically similar selection of reptiles and amphibians to that of Linnaeus. Not until the turn of the century did it become clear that reptiles and amphibians are in fact quite different animals, and Pierre André 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 Thomas Henry Huxley made Latreille's definition popular, and together with Richard Owen expanded Reptilia to include the various fossil “Antediluvian monster”, including 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 (the latter containing the fishes and amphibians). He subsequently proposed the names of Sauropsida and Ichthyopsida for the two.

Around the end of the 19th century, the class reptilia had come to included all the amniotes except birds and mammals. Thus reptiles were defined as the set of animals that includes crocodiles, alligators, tuatara, lizards, snakes, amphisbaenians, and turtles, grouped together as the class Reptilia (Latin repere, "to creep"). This is still the usual definition of the term. However, in recent years, many taxonomists have begun to insist that taxa should be monophyletic, that is, groups should include all descendants of a particular form. The reptiles as defined above would be paraphyletic, since they exclude both birds and mammals, although these also developed from the original reptile. Colin Tudge writes:

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. It is instead 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 Reptila are 'non-avian, non-mammalian amniotes'.

The terms "Sauropsida" ("Lizard Faces") and "Theropsida" ("Beast Faces") were taken up again in 1916 by E.S. Goodrich to distinguish between lizards, birds, and their relatives on one hand (Sauropsida) and mammals and their extinct relatives (Theropsida) 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, the Protosauria ("First Lizards") which included some Paleozoic amphibians as well as early reptiles.

In 1956 D.M.S. Watson observed that the first two groups diverged very early in reptilian history, and so he divided Goodrich's Protosauria among them. He also reinterpreted the Sauropsida and Theropsida to exclude birds and mammals respectively. Thus his Sauropsida included Procolophonia, Eosuchia, Millerosauria, Chelonia (turtles), Squamata (lizards and snakes), Rhynchocephalia, Crocodilia, "thecodonts" (paraphyletic basal Archosauria), non-avian dinosaurs, pterosaurs, ichthyosaurs, and sauropyterygians.

This classification supplemented, but was never as popular as, the classification of the reptiles (according to Romer's classic Vertebrate Paleontology) into four subclasses according to the positioning of temporal fenestrae, openings in the sides of the skull behind the eyes. Those divisions were:

• Anapsida – no fenestrae
• Synapsida – one low fenestra (no longer considered true reptiles)
• Euryapsida – one high fenestra (now included within Diapsida)
• Diapsida – two fenestrae

All of the above but Synapsida fall under Sauropsida.

Taxonomy

Classification to order level, after Benton, 2004.

• Series Amniota
  • Class Synapsida
    • Order Pelycosauria*
    • Order Therapsida
      • Class Mammalia
  • 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 Sphenodontia (tuatara)
          • Order Squamata (lizards & snakes)
      • Infraclass Archosauromorpha
        • Order Prolacertiformes
        • Division Archosauria
          • Subdivision Crurotarsi
            • Superorder Crocodylomorpha
              • Order Crocodilia
              • Order Phytosauria
              • Order Rauisuchia
              • Order Rynchosauria
          • Subdivision Avemetatarsalia
            • Infradivision Ornithodira
              • Order Pterosauria
              • Superorder Dinosauria
                • Order Saurischia
                  • Class Aves
                • Order Ornithischia

Phylogeny

The cladogram presented here illustrates the "family tree" of reptiles, and follows a simplified version of the relationships found by Laurin and Gauthier (1996), presented as part of the Tree of Life Web Project.

Evolutionary history

Rise of the reptiles

The origin of the reptiles lays about 320-310 million years back, in the steaming swamps of the late Carboniferous, when the first reptiles evolved from advanced reptilomorph labyrinthodonts. The oldest traces of reptiles is a series of footprints from the fossil strata of Nova Scotia, dated to 315 million years old. The tracks are attributed to Hylonomus, the oldest known reptile in the biological sense of the word. It was a small, lizard-like animal, about 20 to 30 cm (8-12 inche) long, with numerous sharp teeth indicating an insectivorous diet. Other examples include Westlothiana (for the moment considered to be more related to amphibians than amniotes) and Paleothyris, both of similar build and presumably habit. One of the best known early reptiles is Mesosaurus, a genus of early reptiles from the early Permian that had returned to water, living off fish. The earliest reptiles were largely overshadowed by bigger labyrinthodont amphibians such as Cochleosaurus, and remained a small, inconspicuous part of the fauna until after the small ice age at the end of the Carboniferous.

Anapsids, synapsids and sauropsids

The first reptiles are categorized as Anapsids, having a solid skull with holes only for nose, eyes, spinal cord, etc. Turtles are believed by some to be surviving Anapsids, as they also share this skull structure, but this point has become contentious lately, with some arguing that turtles reverted to this primitive state in order to improve their armor (see Parareptilia). Both sides have strong evidence, and the conflict has yet to be resolved.

Very early after the first reptiles appeared, two branches split off. One lead to the Synapsida (the "mammal-like reptiles" or "stem mammals"), having two openings in the skull roof behind the eyes high , the other group, Diapsida, possessed a pair of holes in their skulls behind the eyes, along with a second pair located higher on the skull. The function of the holes in bout groups was to lighten the skull and give room for the jaw muscles to move, allowing for a more powerful bite. The diapsids and later anapsids are classed as the "true reptiles", the Sauropsida..

Permian reptiles

With the close of the Carboniferous, reptiles became the dominant tetrapod fauna. While the terrestrial reptilomorph labyrinthodonts still existed, the mammal-like reptiles evolved the first terrestrial megafauna in the form of pelycosaurs like Edaphosaurus and the carnivorous Dimetrodon. In the mid-Permian the climate turned dryer, resulting in a faunal turnover. The primitive pelycosaurs where replaced by the more advanced therapsids.

The anapsid reptiles, with their massive skulls without postorbital holes, continued and flourished throughout the Permian. The pareiasaurs reached giant proportions in the late Permian, eventually disappearing at the close of the period (the turtles being possible survivors).

Early in the period, the diapside reptiles split into two lineages, the lepidosaurs (forefathers of modern snakes, lizards, and tuataras). The group remained lizard-like and relatively small and inconspicuous during the whole periode.

Mesozoic, the "Age of Reptiles"

The close of the Permian saw the greatest mass extinction known (see the Permian–Triassic extinction event). Most of the earlier anapsid/synapsid megafauna disappeared, making room for the archosauromorph diapsids. The archosaurs was characterized by elongated hind-legs and an erect pose, the early forms looking somewhat like long legged crocodiles. The archosaurs became the dominant group during the Triassic, developing into the well known dinosaurs and pterosaurs, as well as crocodiles and phytosaurs. Some of the dinosaurs developed into the largest land animals ever to have lived, making the Mesozoic popularly known as the "Age of Reptiles". The dinosaurs also deveoped smaller forms, including the feather-bearing smaller theropds. In the mid Jurassic, these gave rise to the first birds.

The lepidosauromorph diapsids may have been ancestral to the sea reptiles. Developing into the ichthyosaurs and sauropterygians, they came to dominate the Mesozoic seas.

The Therpasids came under increasing pressure from the archosaurs the early Mesozoic and developed into increasingly smaller and more nocturnal forms, the first mammals being the only survivors of the line by late Jurassic.

Demise of the dinosaurs

The close of the Cretacious saw the demise of the Mesozoic reptilian megafauna (see the Cretaceous–Tertiary extinction event). Of the large marine reptiles, only the sea turtles are left, and of the dinosaurs, only the small feathered theropods survived in the form of birds. The major surviving reptilian line is the lepidosaurs, of which the snakes are currently the most numerous and widespread representatives. The end of the "Age of Reptiles", opened up for the "Age of Mammals". Despite this, reptiles are still a major fauna component, particularly in tropical climates. There are about 8200 extant species of reptiles (whereof almost half are snakes), compared to 5400 species of mammals (of which 2/3 are rodents and bats). The most numerous modern group with reptilian roots are the birds, with over 9000 species.

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