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114-412: See text for extinct groups. Reptiles , as commonly defined, are a group of tetrapods with an ectothermic ('cold-blooded') metabolism and amniotic development . Living reptiles comprise four orders : Testudines ( turtles ), Crocodilia ( crocodilians ), Squamata ( lizards and snakes ), and Rhynchocephalia (the tuatara ). As of May 2023, about 12,000 living species of reptiles are listed in

228-559: A clade ( monophyletic group) including birds, though the precise definition of this clade varies between authors. Others prioritize the clade Sauropsida , which typically refers to all amniotes more closely related to modern reptiles than to mammals . The earliest known proto-reptiles originated from the Carboniferous period, having evolved from advanced reptiliomorph tetrapods which became increasingly adapted to life on dry land. The earliest known eureptile ("true reptile")

342-441: A temnospondyl ). A series of footprints from the fossil strata of Nova Scotia dated to 315  Ma show typical reptilian toes and imprints of scales. These tracks are attributed to Hylonomus , the oldest unquestionable reptile known. It was a small, lizard-like animal, about 20 to 30 centimetres (7.9 to 11.8 in) long, with numerous sharp teeth indicating an insectivorous diet. Other examples include Westlothiana (for

456-619: A full complement of limbs. Similar considerations apply to caecilians and aquatic mammals . Newer taxonomy is frequently based on cladistics instead, giving a variable number of major "branches" ( clades ) of the tetrapod family tree . As is the case throughout evolutionary biology today, there is debate over how to properly classify the groups within Tetrapoda. Traditional biological classification sometimes fails to recognize evolutionary transitions between older groups and descendant groups with markedly different characteristics. For example,

570-576: A membrane ensuring gas exchange out of water and can therefore be laid on land. Amphibians and amniotes were affected by the Carboniferous rainforest collapse (CRC), an extinction event that occurred around 307 million years ago. The sudden collapse of a vital ecosystem shifted the diversity and abundance of major groups. Amniotes and temnospondyls in particular were more suited to the new conditions. They invaded new ecological niches and began diversifying their diets to include plants and other tetrapods, previously having been limited to insects and fish. In

684-453: A more recent common ancestry with living amphibians than with living amniotes (reptiles, birds, and mammals). Reptiliomorphs are all animals sharing a more recent common ancestry with living amniotes than with living amphibians. Gaffney (1979) provided the name Neotetrapoda to the crown group of tetrapods, though few subsequent authors followed this proposal. Tetrapoda includes three living classes: amphibians, reptiles, and mammals. Overall,

798-557: A pair of vestigial spurs that are remnants of the hindlimbs . Tetrapods evolved from a group of primitive semiaquatic animals known as the Tetrapodomorpha which, in turn, evolved from ancient lobe-finned fish ( sarcopterygians ) around 390  million years ago in the Middle Devonian period . Tetrapodomorphs were transitional between lobe-finned fishes and true four-limbed tetrapods, though most still fit

912-488: A remnant of the limbs of their distant ancestors. Others returned to being amphibious or otherwise living partially or fully aquatic lives, the first during the Carboniferous period, others as recently as the Cenozoic . One fundamental subgroup of amniotes, the sauropsids , diverged into the reptiles : lepidosaurs (lizards, snakes, and the tuatara ), archosaurs ( crocodilians and dinosaurs , of which birds are

1026-400: A rigid spine. In conjunction with robust forelimbs and shoulder girdle, both Tiktaalik and Ichthyostega may have had the ability to locomote on land in the manner of a seal, with the forward portion of the torso elevated, the hind part dragging behind. Finally, Tiktaalik fin bones are somewhat similar to the limb bones of tetrapods. However, there are issues with positing Tiktaalik as

1140-494: 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

1254-401: A separate subclass, but they are more closely related to mammals than to living reptiles. Considerations like these have led some authors to argue for a new classification based purely on phylogeny , disregarding the anatomy and physiology. Tetrapods evolved from early bony fishes (Osteichthyes), specifically from the tetrapodomorph branch of lobe-finned fishes ( Sarcopterygii ), living in

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1368-497: A single common ancestor. In this sense, Tetrapoda can also be defined as the "clade of limbed vertebrates", including all vertebrates descended from the first limbed vertebrates. A portion of tetrapod workers, led by French paleontologist Michel Laurin , prefer to restrict the definition of tetrapod to the crown group . A crown group is a subset of a category of animal defined by the most recent common ancestor of living representatives. This cladistic approach defines "tetrapods" as

1482-452: A subset of animals related to, but not within, the crown group. The stem and crown group together are combined into the total group , given the name Tetrapodomorpha , which refers to all animals closer to living tetrapods than to Dipnoi ( lungfishes ), the next closest group of living animals. Many early tetrapodomorphs are clearly fish in ecology and anatomy, but later tetrapodomorphs are much more similar to tetrapods in many regards, such as

1596-449: A subset), turtles , and various other extinct forms. The remaining group of amniotes, the synapsids , include mammals and their extinct relatives. Amniotes include the only tetrapods that further evolved for flight—such as birds from among the dinosaurs, the extinct pterosaurs from earlier archosaurs, and bats from among the mammals. The precise definition of "tetrapod" is a subject of strong debate among paleontologists who work with

1710-475: A tetrapod ancestor. For example, it had a long spine with far more vertebrae than any known tetrapod or other tetrapodomorph fish. Also the oldest tetrapod trace fossils (tracks and trackways) predate it by a considerable margin. Several hypotheses have been proposed to explain this date discrepancy: 1) The nearest common ancestor of tetrapods and Tiktaalik dates to the Early Devonian. By this hypothesis,

1824-568: A tiny gecko, Sphaerodactylus ariasae , which can grow up to 17 mm (0.7 in) to the saltwater crocodile , Crocodylus porosus , which can reach over 6 m (19.7 ft) in length and weigh over 1,000 kg (2,200 lb). 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

1938-621: A variety of diets. The following table shows summary estimates for each tetrapod class from the IUCN Red List of Threatened Species , 2014.3, for the number of extant species that have been described in the literature, as well as the number of threatened species . The classification of tetrapods has a long history. Traditionally, tetrapods are divided into four classes based on gross anatomical and physiological traits. Snakes and other legless reptiles are considered tetrapods because they are sufficiently like other reptiles that have

2052-575: A variety of marine organisms and was apparently salt water. The average water temperature was 30 degrees C (86 F). The second oldest evidence for tetrapods, also tracks and trackways, date from ca. 385 Mya ( Valentia Island , Ireland). The oldest partial fossils of tetrapods date from the Frasnian beginning ≈380 mya. These include Elginerpeton and Obruchevichthys . Some paleontologists dispute their status as true (digit-bearing) tetrapods. All known forms of Frasnian tetrapods became extinct in

2166-507: Is foetus , except in medical usage, where fetus is preferred. The -oe- spelling is first attested in 1594 and arose in Late Latin by analogy with classical Latin words like amoenus . In humans, the fetal stage starts nine weeks after fertilization. At this time the fetus is typically about 30 millimetres ( 1 + 1 ⁄ 4  in) in length from crown to rump , and weighs about 8 grams. The head makes up nearly half of

2280-433: Is a concern for medical providers due to the tendency of these infants, described as " premature by weight", to have a higher risk of secondary medical problems. X-rays are known to have possible adverse effects on the development of the fetus, and the risks need to be weighed against the benefits. Congenital disorders are acquired before birth. Infants with certain congenital heart defects can survive only as long as

2394-410: Is also still used in some specialist works like Fortuny et al. (2011). The taxonomy down to subclass level shown here is from Hildebrand and Goslow (2001): This classification is the one most commonly encountered in school textbooks and popular works. While orderly and easy to use, it has come under critique from cladistics . The earliest tetrapods are grouped under class Amphibia, although several of

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2508-708: Is any four- limbed vertebrate animal of the superclass Tetrapoda ( / t ɛ ˈ t r æ p ə d ə / ). Tetrapods include all extant and extinct amphibians and amniotes , with the latter in turn evolving into two major clades , the sauropsids ( reptiles , including dinosaurs and therefore birds ) and synapsids (extinct pelycosaurs , therapsids and all extant mammals , including humans ). Hox gene mutations have resulted in some tetrapods becoming limbless ( snakes , legless lizards , and caecilians ) or two-limbed ( cetaceans , moas , and some lizards ). Nevertheless, these limbless groups still qualify as tetrapods through their ancestry, and some retain

2622-476: 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. General classification of extinct and living reptiles, focusing on major groups. The cladogram presented here illustrates

2736-403: Is highly susceptible to anomalies in its growth and metabolism, increasing the risk of birth defects . One area of concern is the lifestyle choices made during pregnancy. Diet is especially important in the early stages of development. Studies show that supplementation of the person's diet with folic acid reduces the risk of spina bifida and other neural tube defects. Another dietary concern

2850-400: Is imminent and occurs around the 38th week after fertilization. The fetus is considered full-term between weeks 37 and 40 when it is sufficiently developed for life outside the uterus . It may be 48 to 53 cm (19 to 21 in) in length when born. Control of movement is limited at birth, and purposeful voluntary movements continue to develop until puberty . There is much variation in

2964-500: Is possible from the second half of pregnancy . Evidence suggests that the perception of pain in the fetus occurs well before late gestation. Whether a fetus has the ability to feel pain and suffering is part of the abortion debate . In the United States, for example, anti-abortion advocates have proposed legislation that would require providers of abortions to inform pregnant women that their fetuses may feel pain during

3078-467: Is rare for a baby weighing less than 500 g (1 lb 2 oz) to survive. When such premature babies are born, the main causes of mortality are that neither the respiratory system nor the central nervous system are completely differentiated. If given expert postnatal care, some preterm babies weighing less than 500 g (1 lb 2 oz) may survive, and are referred to as extremely low birth weight or immature infants . Preterm birth

3192-465: Is separated from the skull, connected to the torso by muscle and other soft-tissue connections. The result is the appearance of the neck. This feature appears only in tetrapods and Tiktaalik , not other tetrapodomorph fishes. Tiktaalik also had a pattern of bones in the skull roof (upper half of the skull) that is similar to the end-Devonian tetrapod Ichthyostega . The two also shared a semi-rigid ribcage of overlapping ribs, which may have substituted for

3306-477: Is the amnion , which enables the eggs to retain their aqueous contents on land, rather than needing to stay in water. (Some amniotes later evolved internal fertilization , although many aquatic species outside the tetrapod tree had evolved such before the tetrapods appeared, e.g. Materpiscis .) Some tetrapods, such as snakes and caecilians , have lost some or all of their limbs through further speciation and evolution; some have only concealed vestigial bones as

3420-402: Is the earliest known tetrapod that may have had the ability to pull itself onto land and drag itself forward with its forelimbs. There is no evidence that it did so, only that it may have been anatomically capable of doing so. The publication in 2018 of Tutusius umlambo and Umzantsia amazana from high latitude Gondwana setting indicate that the tetrapods enjoyed a global distribution by

3534-404: Is the most common cause of infant mortality, causing almost 30 percent of neonatal deaths. At an occurrence rate of 5% to 18% of all deliveries, it is also more common than postmature birth , which occurs in 3% to 12% of pregnancies. The heart and blood vessels of the circulatory system form relatively early during embryonic development , but continue to grow and develop in complexity in

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3648-543: Is the unborn mammalian offspring that develops from an embryo . Following the embryonic stage , the fetal stage of development takes place. Prenatal development is a continuum, with no clear defining feature distinguishing an embryo from a fetus. However, in general a fetus is characterized by the presence of all the major body organs , though they will not yet be fully developed and functional, and some may not yet be situated in their final anatomical location . In human prenatal development, fetal development begins from

3762-497: Is whether breakfast is eaten. Skipping breakfast could lead to extended periods of lower than normal nutrients in the maternal blood, leading to a higher risk of prematurity , or birth defects. Alcohol consumption may increase the risk of the development of fetal alcohol syndrome , a condition leading to intellectual disability in some infants. Smoking during pregnancy may also lead to miscarriages and low birth weight (2,500 grams (5 pounds 8 ounces). Low birth weight

3876-400: The ductus arteriosus , which directs most of this blood away from the lungs (which are not being used for respiration at this point as the fetus is suspended in amniotic fluid ). With the first breath after birth, the system changes suddenly. Pulmonary resistance is reduced dramatically, prompting more blood to move into the pulmonary arteries from the right atrium and ventricle of

3990-526: The Eifelian stage of the Middle Devonian, 390  million years ago , although these traces have also been interpreted as the ichnogenus Piscichnus (fish nests/feeding traces). The adult tetrapods had an estimated length of 2.5 m (8 feet), and lived in a lagoon with an average depth of 1–2 m, although it is not known at what depth the underwater tracks were made. The lagoon was inhabited by

4104-652: The Late Devonian extinction , also known as the end-Frasnian extinction. This marked the beginning of a gap in the tetrapod fossil record known as the Famennian gap, occupying roughly the first half of the Famennian stage. The oldest near-complete tetrapod fossils, Acanthostega and Ichthyostega , date from the second half of the Fammennian. Although both were essentially four-footed fish, Ichthyostega

4218-405: The Late Devonian extinctions , also known as the end-Frasnian and end-Fammenian extinctions. These extinction events led to the disappearance of stem-tetrapods with fish-like features. When stem-tetrapods reappear in the fossil record in early Carboniferous deposits, some 10 million years later, the adult forms of some are somewhat adapted to a terrestrial existence. Why they went to land in

4332-465: The Permian period, amniotes became particularly well-established, and two important clades filled in most terrestrial niches: the sauropsids and the synapsids . The latter were the most important and successful Permian land animals, establishing complex terrestrial ecosystems of predators and prey while acquiring various adaptations retained by their modern descendants, the mammals. Sauropsid diversity

4446-859: 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 Linnaean taxonomy , reptiles are gathered together under the class Reptilia ( / r ɛ p ˈ t ɪ l i ə / rep- TIL -ee-ə ), which corresponds to common usage. Modern cladistic taxonomy regards that group as paraphyletic , since genetic and paleontological evidence has determined that birds (class Aves), as members of Dinosauria , are more closely related to living crocodilians than to other reptiles, and are thus nested among reptiles from an evolutionary perspective. Many cladistic systems therefore redefine Reptilia as

4560-1166: The Visean age of the Early Carboniferous . The specific aquatic ancestors of the tetrapods and the process by which they colonized Earth's land after emerging from water remains unclear. The transition from a body plan for gill -based aquatic respiration and tail -propelled aquatic locomotion to one that enables the animal to survive out of water and move around on land is one of the most profound evolutionary changes known. Tetrapods have numerous anatomical and physiological features that are distinct from their aquatic fish ancestors. These include distinct head and neck structures for feeding and movements, appendicular skeletons ( shoulder and pelvic girdles in particular) for weight bearing and locomotion, more versatile eyes for seeing, middle ears for hearing, and more efficient heart and lungs for oxygen circulation and exchange outside water. Stem-tetrapods and "fish-a-pods" were primarily aquatic . Modern amphibians , which evolved from earlier groups , are generally semiaquatic ;

4674-418: The amniotic egg . The terms Sauropsida ("lizard faces") and Theropsida ("beast faces") were used 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

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4788-418: The foramen ovale is known as the fossa ovalis . The ductus arteriosus normally closes within one or two days of birth, leaving the ligamentum arteriosum , while the umbilical vein and ductus venosus usually closes within two to five days after birth, leaving, respectively, the liver's ligamentum teres and ligamentum venosus . The placenta functions as a maternal-fetal barrier against

4902-401: The heart . In the fetus, there is an opening between the right and left atrium (the foramen ovale ), and most of the blood flows from the right into the left atrium, thus bypassing pulmonary circulation . The majority of blood flow is into the left ventricle from where it is pumped through the aorta into the body. Some of the blood moves from the aorta through the internal iliac arteries to

5016-415: The tristichopterids (notably Eusthenopteron ), and more recently the elpistostegalians (also known as Panderichthyida) notably the genus Tiktaalik . A notable feature of Tiktaalik is the absence of bones covering the gills. These bones would otherwise connect the shoulder girdle with skull, making the shoulder girdle part of the skull. With the loss of the gill-covering bones, the shoulder girdle

5130-1732: The "family tree" of reptiles, and follows a simplified version of the relationships found by M.S. Lee, in 2013. All genetic studies have supported the hypothesis that turtles are diapsids; some have placed turtles within Archosauromorpha, though a few have recovered turtles as Lepidosauromorpha instead. The cladogram below used a combination of genetic (molecular) and fossil (morphological) data to obtain its results. Synapsida ( mammals and their extinct relatives) [REDACTED] † Millerettidae [REDACTED] † Eunotosaurus † Lanthanosuchidae [REDACTED] † Pareiasauromorpha [REDACTED] † Procolophonoidea [REDACTED] † Captorhinidae [REDACTED] † Paleothyris † Araeoscelidia [REDACTED] † Claudiosaurus [REDACTED] † Younginiformes [REDACTED] † Kuehneosauridae [REDACTED] Rhynchocephalia ( tuatara and their extinct relatives) [REDACTED] Squamata ( lizards and snakes ) [REDACTED] [REDACTED] † Eosauropterygia [REDACTED] † Placodontia [REDACTED] † Sinosaurosphargis † Odontochelys † Proganochelys Testudines ( turtles ) [REDACTED] † Choristodera [REDACTED] † Prolacertiformes [REDACTED] † Rhynchosauria [REDACTED] † Trilophosaurus [REDACTED] Archosauriformes ( crocodiles , birds , dinosaurs and extinct relatives) [REDACTED] [REDACTED] The placement of turtles has historically been highly variable. Classically, turtles were considered to be related to

5244-605: The 13.9-million year Tournaisian, the first stage of the Carboniferous period. Tetrapod-like vertebrates first appeared in the Early Devonian period, and species with limbs and digits were around by the Late Devonian. These early "stem-tetrapods" included animals such as Ichthyostega , with legs and lungs as well as gills, but still primarily aquatic and poorly adapted for life on land. The Devonian stem-tetrapods went through two major population bottlenecks during

5358-452: 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 "III – Amphibia" in his Systema Naturæ . The terms reptile and amphibian were largely interchangeable, reptile (from Latin repere , 'to creep') being preferred by

5472-632: The Cenozoic, similar to mammals. Following the great extinction event at the end of the Mesozoic, representatives of seven major groups of tetrapods persisted into the Cenozoic era. One of them, a group of semiaquatic reptiles known as the Choristodera , became extinct 11 million years ago for unclear reasons. The seven Cenozoic tetrapods groups are: Stem tetrapods are all animals more closely related to tetrapods than to lungfish, but excluding

5586-507: The French zoologist Pierre André Latreille recognized the large physiological differences at the beginning of the 19th century and split the herptiles into two classes, giving the four familiar classes of tetrapods: amphibians, reptiles, birds and mammals. With the basic classification of tetrapods settled, a half a century followed where the classification of living and fossil groups was predominantly done by experts working within classes. In

5700-429: 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 . 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

5814-689: The Permian saw a major turnover in fauna during the Permian–Triassic extinction event . There was a protracted loss of species, due to multiple extinction pulses. Many of the once large and diverse groups died out or were greatly reduced. The diapsid reptiles (a subgroup of the sauropsids) strongly diversified during the Triassic , giving rise to the turtles , pseudosuchians (crocodilian ancestors), dinosaurs , pterosaurs , and lepidosaurs , along with many other reptile groups on land and sea. Some of

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5928-655: The anapsid condition has been found to occur so variably among unrelated groups that it is not now considered a useful distinction. 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

6042-441: The apomorphy-based definition is often supported by an equivalent cladistic definition. Cladistics is a modern branch of taxonomy which classifies organisms through evolutionary relationships, as reconstructed by phylogenetic analyses . A cladistic definition would define a group based on how closely related its constituents are. Tetrapoda is widely considered a monophyletic clade , a group with all of its component taxa sharing

6156-506: The best understood animals since earliest times. By Aristotle 's time, the basic division between mammals, birds and egg-laying tetrapods (the " herptiles ") was well known, and the inclusion of the legless snakes into this group was likewise recognized. With the birth of modern biological classification in the 18th century, Linnaeus used the same division, with the tetrapods occupying the first three of his six classes of animals. While reptiles and amphibians can be quite similar externally,

6270-507: The biodiversity of lissamphibians , as well as of tetrapods generally, has grown exponentially over time; the more than 30,000 species living today are descended from a single amphibian group in the Early to Middle Devonian. However, that diversification process was interrupted at least a few times by major biological crises, such as the Permian–Triassic extinction event , which at least affected amniotes. The overall composition of biodiversity

6384-482: The birds, which evolved from the dinosaurs, are defined as a separate group from them, because they represent a distinct new type of physical form and functionality. In phylogenetic nomenclature , in contrast, the newer group is always included in the old. For this school of taxonomy, dinosaurs and birds are not groups in contrast to each other, but rather birds are a sub-type of dinosaurs. The tetrapods, including all large- and medium-sized land animals, have been among

6498-517: The body plan expected of other lobe-finned fishes. The oldest fossils of four-limbed vertebrates (tetrapods in the broad sense of the word) are trackways from the Middle Devonian , and body fossils became common near the end of the Late Devonian , around 370–360 million years ago. These Devonian species all belonged to the tetrapod stem group , meaning that they were not directly related to any modern tetrapod group. Broad anatomical descriptors like "tetrapod" and "amphibian" can approximate some members of

6612-629: The characteristic Paleozoic non-amniote tetrapods, few survived into the Mesozoic. Temnospondyls briefly recovered in the Triassic, spawning the large aquatic stereospondyls and the small terrestrial lissamphibians (the earliest frogs, salamanders, and caecilians). However, stereospondyl diversity would crash at the end of the Triassic. By the Late Cretaceous, the only surviving amphibians were lissamphibians. Many groups of synapsids, such as anomodonts and therocephalians , that once comprised

6726-420: The class Batracia (1825) 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 ( synapsid ) Dicynodon he helped describe. This

6840-460: The death of the fetus and where this is rarely not resolved it can lead to its formation into a lithopedion . The existence and implications of fetal pain are debated politically and academically. According to the conclusions of a review published in 2005, "Evidence regarding the capacity for fetal pain is limited but indicates that fetal perception of pain is unlikely before the third trimester." However, developmental neurobiologists argue that

6954-527: The dominant terrestrial fauna of the Permian, also became extinct during the Triassic. During the Jurassic, one synapsid group ( Cynodontia ) gave rise to the modern mammals , which survived through the rest of the Mesozoic to later diversify during the Cenozoic. The Cretaceous-Paleogene extinction event at the end of the Mesozoic killed off many organisms, including all the non-avian dinosaurs and nearly all marine reptiles. Birds survived and diversified during

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7068-555: The ductus remains open: in such cases the closure of the ductus can be delayed by the administration of prostaglandins to permit sufficient time for the surgical correction of the anomalies. Conversely, in cases of patent ductus arteriosus , where the ductus does not properly close, drugs that inhibit prostaglandin synthesis can be used to encourage its closure, so that surgery can be avoided. Other heart birth defects include ventricular septal defect , pulmonary atresia , and tetralogy of Fallot . An abdominal pregnancy can result in

7182-405: The earliest members of the group. A majority of paleontologists use the term "tetrapod" to refer to all vertebrates with four limbs and distinct digits (fingers and toes), as well as legless vertebrates with limbed ancestors. Limbs and digits are major apomorphies (newly evolved traits) which define tetrapods, though they are far from the only skeletal or biological innovations inherent to

7296-428: The early 1930s, American vertebrate palaeontologist Alfred Romer (1894–1973) produced an overview, drawing together taxonomic work from the various subfields to create an orderly taxonomy in his Vertebrate Paleontology . This classical scheme with minor variations is still used in works where systematic overview is essential, e.g. Benton (1998) and Knobill and Neill (2006). While mostly seen in general works, it

7410-553: 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. 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

7524-730: The early to middle Devonian period . The first tetrapods probably evolved in the Emsian stage of the Early Devonian from Tetrapodomorph fish living in shallow water environments. The very earliest tetrapods would have been animals similar to Acanthostega , with legs and lungs as well as gills, but still primarily aquatic and unsuited to life on land. The earliest tetrapods inhabited saltwater, brackish-water, and freshwater environments, as well as environments of highly variable salinity. These traits were shared with many early lobed-finned fishes. As early tetrapods are found on two Devonian continents, Laurussia ( Euramerica ) and Gondwana , as well as

7638-584: The end of the Devonian and even extend into the high latitudes. The end-Fammenian marked another extinction, known as the end-Fammenian extinction or the Hangenberg event , which is followed by another gap in the tetrapod fossil record, Romer's gap , also known as the Tournaisian gap. This gap, which was initially 30 million years, but has been gradually reduced over time, currently occupies much of

7752-405: The establishment of thalamocortical connections (at about 6 + 1 ⁄ 2 months) is an essential event with regard to fetal perception of pain. Nevertheless, the perception of pain involves sensory, emotional and cognitive factors and it is "impossible to know" when pain is experienced, even if it is known when thalamocortical connections are established. Some authors argue that fetal pain

7866-451: The exception of humans. The duration of gestation in placental mammals varies from 18 days in jumping mice to 23 months in elephants . Generally speaking, fetuses of larger land mammals require longer gestation periods. The benefits of a fetal stage means that young are more developed when they are born. Therefore, they may need less parental care and may be better able to fend for themselves. However, carrying fetuses exerts costs on

7980-453: The fetus develops within the mother, using a (non-mammalian) placenta rather than contained in an eggshell . As amniotes, reptile eggs are surrounded by membranes for protection and transport, which adapt them to reproduction on dry land. 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

8094-430: The first evidence of their function does not occur until around 30 weeks. Bones are fully developed but are still soft and pliable. Iron , calcium , and phosphorus become more abundant. Fingernails reach the end of the fingertips. The lanugo , or fine hair, begins to disappear until it is gone except on the upper arms and shoulders. Small breast buds are present in both sexes. Head hair becomes coarse and thicker. Birth

8208-446: The first place is still debated. During the early Carboniferous, the number of digits on hands and feet of stem-tetrapods became standardized at no more than five, as lineages with more digits died out (exceptions within crown-group tetrapods arose among some secondarily aquatic members). By mid-Carboniferous times, the stem-tetrapods had radiated into two branches of true ("crown group") tetrapods, one ancestral to modern amphibians and

8322-489: The first stages of their lives are as waterborne eggs and fish-like larvae known as tadpoles , and later undergo metamorphosis to grow limbs and become partly terrestrial and partly aquatic. However, most tetrapod species today are amniotes , most of which are terrestrial tetrapods whose branch evolved from earlier tetrapods early in the Late Carboniferous . The key innovation in amniotes over amphibians

8436-437: The first time ( nulliparous ) typically feels fetal movements at about 21 weeks, whereas a woman who has given birth before will typically feel movements by 20 weeks. By the end of the fifth month, the fetus is about 20 cm (8 in) long. The amount of body fat rapidly increases. Lungs are not fully mature. Neural connections between the sensory cortex and thalamus develop as early as 24 weeks of gestational age, but

8550-491: The group. The first vertebrates with limbs and digits evolved in the Devonian , including the Late Devonian -age Ichthyostega and Acanthostega , as well as the trackmakers of the Middle Devonian -age Zachelmie trackways . Defining tetrapods based on one or two apomorphies can present a problem if these apomorphies were acquired by more than one lineage through convergent evolution . To resolve this potential concern,

8664-419: The groups are more closely related to amniotes than to modern day amphibians . Traditionally, birds are not considered a type of reptile, but crocodiles are more closely related to birds than they are to other reptiles, such as lizards. Birds themselves are thought to be descendants of theropod dinosaurs . Basal non-mammalian synapsids ("mammal-like reptiles") traditionally also sort under class Reptilia as

8778-415: The growing fetus. A functional circulatory system is a biological necessity since mammalian tissues can not grow more than a few cell layers thick without an active blood supply. The prenatal circulation of blood is different from postnatal circulation, mainly because the lungs are not in use. The fetus obtains oxygen and nutrients from the mother through the placenta and the umbilical cord . Blood from

8892-539: The growth of the human fetus. When the fetal size is less than expected, the condition is known as intrauterine growth restriction also called fetal growth restriction; factors affecting fetal growth can be maternal , placental , or fetal . Fetal growth is often classified as follows: small for gestational age (SGA), appropriate for gestational age (AGA), and large for gestational age (LGA). SGA can result in low birth weight , although premature birth can also result in low birth weight. Low birth weight increases

9006-412: The heart and less to flow through the foramen ovale into the left atrium . The blood from the lungs travels through the pulmonary veins to the left atrium, producing an increase in pressure that pushes the septum primum against the septum secundum , closing the foramen ovale and completing the separation of the newborn's circulatory system into the standard left and right sides. Thereafter,

9120-750: The human fetus within their mothers. However, the anatomy of the area surrounding a fetus is different in litter-bearing animals compared to humans: each fetus of a litter-bearing animal is surrounded by placental tissue and is lodged along one of two long uteri instead of the single uterus found in a human female. Development at birth varies considerably among animals, and even among mammals. Altricial species are relatively helpless at birth and require considerable parental care and protection. In contrast, precocial animals are born with open eyes, have hair or down, have large brains, and are immediately mobile and somewhat able to flee from, or defend themselves against, predators . Primates are precocial at birth, with

9234-463: The hypothesis that turtles belong to a separate clade within Sauropsida , outside the saurian clade altogether. The origin of the reptiles lies about 310–320 million years ago, in the steaming swamps of the late Carboniferous period, when the first reptiles evolved from advanced reptiliomorphs . The oldest known animal that may have been an amniote is Casineria (though it may have been

9348-408: The island of North China , it is widely supposed that early tetrapods were capable of swimming across the shallow (and relatively narrow) continental-shelf seas that separated these landmasses. Since the early 20th century, several families of tetrapodomorph fishes have been proposed as the nearest relatives of tetrapods, among them the rhizodonts (notably Sauripterus ), the osteolepidids ,

9462-506: 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. The synapsid/sauropsid division supplemented another approach, one that split

9576-508: 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

9690-579: The lineage is the closest to tetrapods, but Tiktaalik itself was a late-surviving relic. 2) Tiktaalik represents a case of parallel evolution. 3) Tetrapods evolved more than once. [REDACTED] Coelacanthiformes (coelacanths) [REDACTED] Dipnoi (lungfish) [REDACTED] †Tetrapodomorph fishes [REDACTED] Tetrapoda [REDACTED] The oldest evidence for the existence of tetrapods comes from trace fossils : tracks (footprints) and trackways found in Zachełmie , Poland, dated to

9804-795: 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 (e.g., mosasaurs ). Modern non-bird reptiles inhabit all the continents except Antarctica. Reptiles are tetrapod vertebrates , creatures that either have four limbs or, like snakes, are descended from four-limbed ancestors. Unlike amphibians , reptiles do not have an aquatic larval stage. Most reptiles are oviparous , although several species of squamates are viviparous , as were some extinct aquatic clades  –

9918-401: The moment considered a reptiliomorph rather than a true amniote ) and Paleothyris , both of similar build and presumably similar habit. However, microsaurs have been at times considered true reptiles, so an earlier origin is possible. Tetrapod A tetrapod ( / ˈ t ɛ t r ə ˌ p ɒ d / ; from Ancient Greek τετρα- (tetra-) 'four' and πούς (poús) 'foot')

10032-412: The nearest common ancestor of all living amphibians (the lissamphibians) and all living amniotes (reptiles, birds, and mammals), along with all of the descendants of that ancestor. In effect, "tetrapod" is a name reserved solely for animals which lie among living tetrapods, so-called crown tetrapods. This is a node-based clade , a group with a common ancestry descended from a single "node" (the node being

10146-408: The nearest common ancestor of living species). Defining tetrapods based on the crown group would exclude many four-limbed vertebrates which would otherwise be defined as tetrapods. Devonian "tetrapods", such as Ichthyostega and Acanthostega , certainly evolved prior to the split between lissamphibians and amniotes, and thus lie outside the crown group. They would instead lie along the stem group ,

10260-802: The new Triassic reptiles would not survive into the Jurassic , but others would flourish during the Jurassic. Lizards , turtles, dinosaurs, pterosaurs, crocodylomorphs , and plesiosaurs were particular beneficiaries of the Triassic-Jurassic transition. Birds , a particular subset of theropod dinosaurs capable of flight via feathered wings, evolved in the Late Jurassic. In the Cretaceous , snakes developed from lizards, rhynchocephalians (tuataras and kin) declined, and modern birds and crocodilians started to establish themselves. Among

10374-558: The ninth week after fertilization (which is the eleventh week of gestational age ) and continues until the birth of a newborn . The word fetus (plural fetuses or rarely feti ) comes from Latin fētus 'offspring, bringing forth, hatching of young'. The Latin plural fetūs is not used in English ; occasionally the plural feti is used in English by analogy with second-declension Latin nouns ending in -us . The predominant British, Irish, and Commonwealth spelling

10488-407: The other ancestral to amniotes. Modern amphibians are most likely derived from the temnospondyls , a particularly diverse and long-lasting group of tetrapods. A less popular proposal draws comparisons to the " lepospondyls ", an eclectic mixture of various small tetrapods, including burrowing, limbless, and other bizarrely-shaped forms. The reptiliomorphs (sometimes known as " anthracosaurs ") were

10602-422: The other euryapsids, and given the older name Parapsida. Parapsida was later discarded as a group for the most part (ichthyosaurs being classified as incertae sedis or with Euryapsida). However, four (or three if Euryapsida is merged into Diapsida) subclasses remained more or less universal for non-specialist work throughout the 20th century. It has largely been abandoned by recent researchers: In particular,

10716-401: The placenta is carried to the fetus by the umbilical vein . About half of this enters the fetal ductus venosus and is carried to the inferior vena cava , while the other half enters the liver proper from the inferior border of the liver. The branch of the umbilical vein that supplies the right lobe of the liver first joins with the portal vein . The blood then moves to the right atrium of

10830-454: The presence of limbs and digits. Laurin's approach to the definition of tetrapods is rooted in the belief that the term has more relevance for neontologists (zoologists specializing in living animals) than paleontologists (who primarily use the apomorphy-based definition). In 1998, he re-established the defunct historical term Stegocephali to replace the apomorphy-based definition of tetrapod used by many authors. Other paleontologists use

10944-412: The primitive anapsid reptiles. Molecular work has usually placed turtles within the diapsids. As of 2013, three turtle genomes have been sequenced. The results place turtles as a sister clade to the archosaurs , the group that includes crocodiles, non-avian dinosaurs, and birds. However, in their comparative analysis of the timing of organogenesis , Werneburg and Sánchez-Villagra (2009) found support for

11058-637: The procedure and that would require each person to accept or decline anesthesia for the fetus. Abortion of a human pregnancy is legal and/or tolerated in most countries, although with gestational time limits that normally prohibit late-term abortions . A fetus is a stage in the prenatal development of viviparous organisms. This stage lies between embryogenesis and birth. Many vertebrates have fetal stages, ranging from most mammals to many fish. In addition, some invertebrates bear live young, including some species of onychophora and many arthropods . The fetuses of most mammals are situated similarly to

11172-430: The relatives and ancestors of the amniotes (reptiles, mammals, and kin). The first amniotes are known from the early part of the Late Carboniferous . All basal amniotes had a small body size, like many of their contemporaries, though some Carboniferous tetrapods evolved into large crocodile-like predators, informally known as " labyrinthodonts ". Amphibians must return to water to lay eggs; in contrast, amniote eggs have

11286-436: 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: The composition of Euryapsida was uncertain. Ichthyosaurs were, at times, considered to have arisen independently of

11400-414: The risk for perinatal mortality ( death shortly after birth), asphyxia , hypothermia , polycythemia , hypocalcemia , immune dysfunction , neurologic abnormalities, and other long-term health problems. SGA may be associated with growth delay, or it may instead be associated with absolute stunting of growth. Fetal viability refers to a point in fetal development at which the fetus may survive outside

11514-413: The size of the fetus. Breathing-like movements of the fetus are necessary for the stimulation of lung development , rather than for obtaining oxygen. The heart, hands, feet, brain, and other organs are present, but are only at the beginning of development and have minimal operation. Uncontrolled movements and twitches occur as muscles , the brain , and pathways begin to develop. A woman pregnant for

11628-502: The stem group, but a few paleontologists opt for more specific terms such as Stegocephali . Limbs evolved prior to terrestrial locomotion , but by the start of the Carboniferous Period, 360 million years ago, a few stem-tetrapods were experimenting with a semiaquatic lifestyle to exploit food and shelter on land. The first crown -tetrapods (those descended from the last common ancestors of extant tetrapods) appeared by

11742-787: The structure of the forebrain. According to Goodrich, both lineages evolved from an earlier stem group, Protosauria ("first lizards") 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 (turtles), Squamata (lizards and snakes), Rhynchocephalia , Crocodilia , " thecodonts " ( paraphyletic basal Archosauria ), non- avian dinosaurs , pterosaurs , ichthyosaurs , and sauropterygians . In

11856-521: The term stem-tetrapod to refer to those tetrapod-like vertebrates that are not members of the crown group, including both early limbed "tetrapods" and tetrapodomorph fishes. The term "fishapod" was popularized after the discovery and 2006 publication of Tiktaalik , an advanced tetrapodomorph fish which was closely related to limbed vertebrates and showed many apparently transitional traits. The two subclades of crown tetrapods are Batrachomorpha and Reptiliomorpha . Batrachomorphs are all animals sharing

11970-483: The tetrapod crown group. The cladogram below illustrates the relationships of stem-tetrapods. All these lineages are extinct except for Dipnomorpha and Tetrapoda; from Swartz, 2012: Dipnomorpha (lungfishes and relatives) [REDACTED] Kenichthys Rhizodontidae [REDACTED] Marsdenichthys Canowindra Koharalepis Fetus A fetus or foetus ( / ˈ f iː t ə s / ; pl. : fetuses , foetuses , rarely feti or foeti )

12084-414: The transmission of microbes . When this is insufficient, mother-to-child transmission of infectious diseases can occur. Maternal IgG antibodies cross the placenta, giving the fetus passive immunity against those diseases for which the mother has antibodies. This transfer of antibodies in humans begins as early as the fifth month (gestational age) and certainly by the sixth month. A developing fetus

12198-406: The umbilical arteries and re-enters the placenta, where carbon dioxide and other waste products from the fetus are taken up and enter the mother's circulation. Some of the blood from the right atrium does not enter the left atrium, but enters the right ventricle and is pumped into the pulmonary artery . In the fetus, there is a special connection between the pulmonary artery and the aorta, called

12312-520: The womb. The lower limit of viability is approximately 5 + 3 ⁄ 4 months gestational age and is usually later. There is no sharp limit of development, age, or weight at which a fetus automatically becomes viable. According to data from 2003 to 2005, survival rates are 20–35% for babies born at 23 weeks of gestation ( 5 + 3 ⁄ 4 months); 50–70% at 24–25 weeks (6 – 6 + 1 ⁄ 4 months); and >90% at 26–27 weeks ( 6 + 1 ⁄ 2 – 6 + 3 ⁄ 4 months) and over. It

12426-589: 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

12540-526: Was Hylonomus , a small and superficially lizard-like animal which lived in Nova Scotia during the Bashkirian age of the Late Carboniferous , around 318  million years ago . Genetic and fossil data argues that the two largest lineages of reptiles, Archosauromorpha (crocodilians, birds, and kin) and Lepidosauromorpha (lizards, and kin), diverged during the Permian period. In addition to

12654-588: 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

12768-547: Was driven primarily by amphibians in the Palaeozoic, dominated by reptiles in the Mesozoic and expanded by the explosive growth of birds and mammals in the Cenozoic. As biodiversity has grown, so has the number of species and the number of niches that tetrapods have occupied. The first tetrapods were aquatic and fed primarily on fish. Today, the Earth supports a great diversity of tetrapods that live in many habitats and subsist on

12882-417: Was more subdued during the Permian, but they did begin to fracture into several lineages ancestral to modern reptiles. Amniotes were not the only tetrapods to experiment with prolonged life on land. Some temnospondyls, seymouriamorphs , and diadectomorphs also successfully filled terrestrial niches in the earlier part of the Permian. Non-amniote tetrapods declined in the later part of the Permian. The end of

12996-626: 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 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

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