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Reptiles, or members of the class Reptilia, are air-breathing, generally "cold-blooded" (poikilothermic) amniotes whose skin is usually covered in scales or scutes. They are tetrapods (either having four limbs or being descended from four-limbed ancestors) and lay amniotic eggs, in which the embryo is surrounded by a membrane called the amnion. Modern reptiles inhabit every continent with the exception of Antarcticamarker, and four living orders are currently recognized:

The majority of reptile species are oviparous (egg-laying), although certain species of squamates are capable of giving live birth. This is achieved by either ovoviviparity (egg retention) or viviparity (birth of offspring without the development 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, Crocodylus porosus, 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 Swedenmarker, 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, "reptile" (from Latin repere, "to creep") being preferred by the French.Josephus Nicolaus Laurenti was 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 (1825) 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 monsters”, 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 Surgeonsmarker 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 include all the amniotes except birds and mammals. Thus reptiles were defined as the set of animals that includes the extant crocodiles, alligators, tuatara, lizards, snakes, amphisbaenians, and turtles, as well as fossil groups like dinosaurs, synapsids and the primitive pareiasaurs. 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 evolved 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 Reptilia 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 the 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 between 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 sauropterygians.

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 number and position of temporal fenestrae, openings in the sides of the skull behind the eyes. Those subclasses 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 except Synapsida are included within Sauropsida.

Taxonomy

Classification to order level, after Benton, 2004.

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 lies about 320–310 million years ago, in the steaming swamps of the late Carboniferous, when the first reptiles evolved from advanced reptiliomorph labyrinthodonts.The oldest trace of reptiles is a series of footprints from the fossil strata of Nova Scotiamarker, dated to 315 million years ago.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 in) long, with numerous sharp teeth indicating an insectivorous diet.Other examples include Westlothiana (for the moment considered to be more closely related to amphibians than to amniotes) and Paleothyris, both of similar build and presumably similar habit. One of the best known early reptiles is Mesosaurus, a genus from the early Permian that had returned to water, feeding on 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

A = Anapsid, B = Synapsid, C = Diapsid
The first reptiles are categorized as anapsids, having a solid skull with holes for only nose, eyes, spinal cord, etc.Turtles are believed by some to be surviving anapsids, since they 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 cite strong evidence, and the conflict has yet to be resolved.

Very soon after the first reptiles appeared, they split into two branches.One branch, the Synapsida ("mammal-like reptiles" or "stem mammals"), had two openings in the skull roof behind the eyes; the other branch, the 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 both 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 reptiliomorph 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 change of fauna: The primitive pelycosaurs were replaced by the more advanced therapsids.

The anapsid reptiles, whose massive skulls had no 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 diapsid reptiles split into two main lineages, the archosaurs (forefathers of crocodiles and dinosaurs) and the lepidosaur (predecessors of modern snakes, lizards, and tuataras). Both groups remained lizard-like and relatively small and inconspicuous during the Permian.

The 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, being replaced by the archosauromorph diapsids. The archosaurs were 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 developed smaller forms, including the feather-bearing smaller theropods. In the mid-Jurassic, these gave rise to the first birds.

The lepidosauromorph diapsids may have been ancestral to the sea reptiles.These reptiles developed into the sauropterygians in the early Triassic and the ichthyosaurs during the Middle Triassic. The mosasaurs also evolved in the Mesozoic Era, emerging during the Cretaceous period.

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

Demise of the dinosaurs

The close of the Cretaceous 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” led into the “Age of Mammals”. Despite this, reptiles are still a major faunal component, particularly in tropical climates. There are about 8,200 extant species of reptiles (of which almost half are snakes), compared with 5,400 species of mammals (of which two-thirds are rodents and bats). The most numerous modern group with reptilian roots are the birds, with over 9,000 species.

Systems

Circulatory

Most reptiles have a three-chambered heart consisting of two atria, one variably-partitioned ventricle, and two aortas that lead to the systemic circulation. The degree of mixing of oxygenated and deoxygenated blood in the three-chambered heart varies depending on the species and physiological state. Under different conditions, deoxygenated blood can be shunted back to the body or oxygenated blood can be shunted back to the lungs. This variation in blood flow has been hypothesized to allow more effective thermoregulation and longer diving times for aquatic species, but has not been shown to be a fitness advantage.

There are some interesting exceptions to the general physiology. For instance, crocodilians have an anatomically four-chambered heart, but also have two systemic aortas and are therefore capable of bypassing only their pulmonary circulation. Also, some snake and lizard species (e.g., pythons and monitor lizards) have three-chambered hearts that become functionally four-chambered hearts during contraction. This is made possible by a muscular ridge that subdivides the ventricle during ventricular diastole and completely divides it during ventricular systole. Because of this ridge, some of these squamates are capable of producing ventricular pressure differentials that are equivalent to those seen in mammalian and avian hearts.

Respiratory

Reptilian lungs

All reptiles breathe using lungs. Aquatic turtles have developed more permeable skin, and some species have modified their cloaca to increase the area for gas exchange (Orenstein, 2001). Even with these adaptations, breathing is never fully accomplished without lungs. Lung ventilation is accomplished differently in each main reptile group. In squamates, the lungs are ventilated almost exclusively by the axial musculature. This is also the same musculature that is used during locomotion. Because of this constraint, most squamates are forced to hold their breath during intense runs. Some, however, have found a way around it. Varanids, and a few other lizard species, employ buccal pumping as a complement to their normal "axial breathing." This allows the animals to completely fill their lungs during intense locomotion, and thus remain aerobically active for a long time. Tegu lizards are known to possess a proto-diaphragm, which separates the pulmonary cavity from the visceral cavity. While not actually capable of movement, it does allow for greater lung inflation, by taking the weight of the viscera off the lungs (Klein et al., 2003). Crocodilians actually have a muscular diaphragm that is analogous to the mammalian diaphragm. The difference is that the muscles for the crocodilian diaphragm pull the pubis (part of the pelvis, which is movable in crocodilians) back, which brings the liver down, thus freeing space for the lungs to expand. This type of diaphragmatic setup has been referred to as the "hepatic piston."

Turtles and tortoises

How turtles and tortoises breathe has been the subject of much study. To date, only a few species have been studied thoroughly enough to get an idea of how turtles do it. The results indicate that turtles and tortoises have found a variety of solutions to this problem. The difficulty is that most turtle shells are rigid and do not allow for the type of expansion and contraction that other amniotes use to ventilate their lungs. Some turtles such as the Indian flapshell (Lissemys punctata) have a sheet of muscle that envelops the lungs. When it contracts, the turtle can exhale. When at rest, the turtle can retract the limbs into the body cavity and force air out of the lungs. When the turtle protracts its limbs, the pressure inside the lungs is reduced, and the turtle can suck air in. Turtle lungs are attached to the inside of the top of the shell (carapace), with the bottom of the lungs attached (via connective tissue) to the rest of the viscera. By using a series of special muscles (roughly equivalent to a diaphragm), turtles are capable of pushing their viscera up and down, resulting in effective respiration, since many of these muscles have attachment points in conjunction with their forelimbs (indeed, many of the muscles expand into the limb pockets during contraction). Breathing during locomotion has been studied in three species, and they show different patterns. Adult female green sea turtles do not breathe as they crutch along their nesting beaches. They hold their breath during terrestrial locomotion and breathe in bouts as they rest. North American box turtles breathe continuously during locomotion, and the ventilation cycle is not coordinated with the limb movements (Landberg et al., 2003). They are probably using their abdominal muscles to breathe during locomotion. The last species to have been studied is the red-eared slider, which also breathes during locomotion, but takes smaller breaths during locomotion than during small pauses between locomotor bouts, indicating that there may be mechanical interference between the limb movements and the breathing apparatus. Box turtles have also been observed to breathe while completely sealed up inside their shells (ibid.).

Palate

Most reptiles lack a secondary palate, meaning that they must hold their breath while swallowing. Crocodilians have evolved a bony secondary palate that allows them to continue breathing while remaining submerged (and protect their brains against damage by struggling prey). Skinks (family Scincidae) also have evolved a bony secondary palate, to varying degrees. Snakes took a different approach and extended their trachea instead. Their tracheal extension sticks out like a fleshy straw, and allows these animals to swallow large prey without suffering from asphyxiation.

Skin

Reptilian skin is covered in a horny epidermis, making it watertight and enabling reptiles to live on dry land, in contrast to amphibians. Compared to mammalian skin, that of reptiles is rather thin and lacks the thick dermal layer that produces leather in mammals.Exposed parts of reptiles are protected by scales or scutes, sometimes with a bony base, forming armor. In lepidosaurians like lizards and snakes, the whole skin is covered in epidermal scales. Such scales were once thought to be typical of the class Reptilia as a whole, but are now known to occur only in lepidosaurians. The scales found in turtles and crocodiles are of dermal, rather than epidermal, origin and are properly termed scutes. In turtles, the body is hidden inside a hard shell composed of fused scutes.

Excretory

Excretion is performed mainly by two small kidneys. In diapsids, uric acid is the main nitrogenous waste product; turtles, like mammals, excrete mainly urea. Unlike the kidneys of mammals and birds, reptile kidneys are unable to produce liquid urine more concentrated than their body fluid. This is because they lack a specialized structure called a loop of Henle, which is present in the nephrons of birds and mammals,. Because of this, many reptiles use the colon to aid in the reabsorption of water. Some are also able to take up water stored in the bladder. Excess salts are also excreted by nasal and lingual salt glands in some reptiles.

Digestive systems

Most reptiles are carnivorous and have rather simple and comparatively short guts, meat being fairly simple to break down and digest. Digestion is slower than in mammals, reflecting their lower metabolism and their inability to divide and masticate their food. Being poikilotherms (with varying body temperature regulated by their environment), their energy requirement is about a fifth to a tenth of that of a mammal of the same size. Large reptiles like crocodiles and the large constrictors can live from a single large meal for months, digesting it slowly.

While modern reptiles are predominately carnivorous, during the early history of reptiles several groups produced a herbivorous megafauna: in the Paleozoic the pareiasaurs and the synapsid dicynodonts, and in the Mesozoic several lines of dinosaurs. Today the turtles are the only predominantly herbivorous reptile group, but several lines of agams and iguanas have evolved to live wholly or partly on plants.

Herbivorous reptiles face the same problems of mastication as herbivorous mammals, but, lacking the complex teeth of mammals, many species swallow rocks and pebbles (so called gastroliths) to aid in digestion: The rocks are washed around in the stomach, helping to grind up plant matter. Fossil gastroliths have been found associated with sauropods. Sea turtles, crocodiles, and marine iguanas also use gastroliths as ballast, helping them to dive.

Nervous system

The reptilian nervous system contains the same basic part of the amphibian brain, but the reptile cerebrum and cerebellum are slightly larger. Most typical sense organs are well developed with certain exceptions, most notably the snake's lack of external ears (middle and inner ears are present). There are twelve pairs of cranial nerves.[4198]

Reptiles are generally considered less intelligent than mammals and birds.The size of their brain relative to their body is much less than that of mammals, the encephalization quotient being about one tenth of that of mammals.Crocodiles have relatively larger brains and show a fairly complex social structure. Larger lizards like the monitors are known to exhibit complex behavior, including cooperation.The Komodo dragon is known to engage in play.

Vision

Most reptiles are diurnal animals. The vision is typically adapted to daylight conditions, with color vision and more advanced visual depth perception than in amphibians and most mammals. In some species, such as blind snakes, vision is reduced.Some snakes have extra sets of visual organs (in the loosest sense of the word) in the form of pits sensitive to infrared radiation (heat). Such heat-sensitive pits are particularly well developed in the pit vipers, but are also found in boas and pythons. These pits allow the snakes to sense the body heat of birds and mammals, enabling pit vipers to hunt rodents in the dark.

Reproductive

Most reptiles reproduce sexually, though some are capable of asexual reproduction. All reproductive activity occurs through the cloaca, the single exit/entrance at the base of the tail where waste is also eliminated. Most reptiles have copulatory organs, which are usually retracted or inverted and stored inside the body. In turtles and crocodilians, the male has a single median penis, while squamates, including snakes and lizards, possess a pair of hemipenes. Tuataras, however, lack copulatory organs, and so the male and female simply press their cloacas together as the male excretes sperm.

Most reptiles lay amniotic eggs covered with leathery or calcareous shells. An amnion, chorion, and allantois are present during embryonic life. There are no larval stages of development. Viviparity and ovoviviparity have evolved only in squamates, and many species, including all boas and most vipers, utilize this mode of reproduction. The degree of viviparity varies: some species simply retain the eggs until just before hatching, others provide maternal nourishment to supplement the yolk, and yet others lack any yolk and provide all nutrients via a structure similar to the mammalian placenta.

Asexual reproduction has been identified in squamate in six families of lizards and one snake. In some species of squamates, a population of females is able to produce a unisexual diploid clone of the mother. This form of asexual reproduction, called parthenogenesis, occurs in several species of gecko, and is particularly widespread in the teiids (especially Aspidocelis) and lacertids (Lacerta). In captivity, Komodo dragons (Varanidae) have reproduced by parthenogenesis.

Parthenogenetic species are suspected to occur among chameleons, agamids, xantusiids, and typhlopids.

Some reptiles exhibit temperature-dependent sex determination (TDSD), in which the incubation temperature determines whether a particular egg hatches as male or female. TDSD is most common in turtles and crocodiles, but also occurs in lizards and tuataras. To date, there has been no confirmation of whether TDSD occurs in snakes.

See also





References

  1. Encyclopaedia Britannica, 9th ed. (1878). original text
  2. Laurenti, J.N. (1768): Specimen Medicum, Exhibens Synopsin Reptilium Emendatam cum Experimentis circa Venena. Facsimile, showing the mixed composition of his Reptilia
  3. Latreielle, P.A. (1804): Nouveau Dictionnaire à Histoire Naturelle, xxiv., cited in Latreille's Familles naturelles du règne animal, exposés succinctement et dans un ordre analytique, 1825
  4. Huxley, T.H. (1863): The Structure and Classification of the Mammalia. Hunterian lectures, presented in Medical Times and Gazette, 1863. original text
  5. , 3rd ed., 1966.
  6. Laurin, M. and Gauthier, J.A. (1996). "Amniota. Mammals, reptiles (turtles, lizards, Sphenodon, crocodiles, birds) and their extinct relatives." Version 01 January 1996. http://tolweb.org/Amniota/14990/1996.01.01 in The Tree of Life Web Project, http://tolweb.org/
  7. ( abstract)
  8. Falcon-Lang, H.J., Benton, M.J. & Stimson, M. (2007): Ecology of early reptiles inferred from Lower Pennsylvanian trackways. Journal of the Geological Society, London, 164; no. 6; pp 1113-1118. article
  9. Earliest Evidence For Reptiles
  10. Romer, A.S. & T.S. Parsons. 1977. The Vertebrate Body. 5th ed. Saunders, Philadelphia. (6th ed. 1985)
  11. , 3rd ed. 2004 ISBN 0632056371
  12. van Tuninen, M. & Hadly, E.A. (2004): Error in Estimation of Rate and Time Inferred from the Early Amniote Fossil Record and Avian Molecular Clocks. Journal of Mulecular Biology, no 59: pp 267-276 PDF
  13. 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 9780471384618.
  14. 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 9780471384618.
  15. 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 9780471384618.
  16. Gauthier J. A. (1994): The diversification of the amniotes. In: D. R. Prothero and R. M. Schoch (ed.) Major Features of Vertebrate Evolution: 129-159. Knoxville, Tennessee: The Paleontological Society.
  17. Hildebran, M. & Goslow, G. (2001): Analysis of Vertebrate Structure. 5th edition. John Wiley & sons inc, New York. 635 pages ISBN 0-471-29505-1
  18. Figure of relative brain size in vertebrates
  19. King, Dennis & Green, Brian. 1999. Goannas: The Biology of Varanid Lizards. University of New South Wales Press. ISBN 0-86840-456-X, p. 43.
  20. [1]
  21. Lutz, Dick (2005), Tuatara: A Living Fossil, Salem, Oregon: DIMI PRESS, ISBN 0-931625-43-2
  22. http://books.google.com/books?id=lc5Bg-hmsBYC&lpg=PA101&ots=Pwea4SV09u&dq=temperature%20dependent%20sex%20determination%20snake&pg=PA101#v=onepage&q=temperature%20dependent%20sex%20determination%20snake&f=false


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