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55-468: Cavalier-Smith proposed the term 'Archezoa' for a paraphyletic (see Paraphyly ) territory of eukaryotes that primitively lacked mitochondria. Like Margulis and others before (see Pelomyxa ), Cavalier-Smith argued that the initial ancestor of eukaryotes emerged prior to the endosymbiotic acquisition (see endosymbiosis ) of mitochondria. The same paraphyletic territory was referred to as 'Hypochondria' by others. The argument for Archezoa sensu Cavalier-Smith

110-562: 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")

165-447: 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

220-479: A "single common ancestor" organism. Paraphyly is common in speciation , whereby a mother species (a paraspecies ) gives rise to a daughter species without itself becoming extinct. Research indicates as many as 20 percent of all animal species and between 20 and 50 percent of plant species are paraphyletic. Accounting for these facts, some taxonomists argue that paraphyly is a trait of nature that should be acknowledged at higher taxonomic levels. Cladists advocate

275-592: A cell nucleus, a plesiomorphy ) from its excluded descendants. Also, some systematists recognize paraphyletic groups as being involved in evolutionary transitions, the development of the first tetrapods from their ancestors for example. Any name given to these hypothetical ancestors to distinguish them from tetrapods—"fish", for example—necessarily picks out a paraphyletic group, because the descendant tetrapods are not included. Other systematists consider reification of paraphyletic groups to obscure inferred patterns of evolutionary history. The term " evolutionary grade "

330-421: A class of systematic errors in phylogenetic analysis called " long branch attraction ". This eukaryote -related article is a stub . You can help Misplaced Pages by expanding it . Paraphyly Paraphyly is a taxonomic term describing a grouping that consists of the grouping's last common ancestor and some but not all of its descendant lineages. The grouping is said to be paraphyletic with respect to

385-407: A common ancestor are said to be monophyletic . A paraphyletic group is a monophyletic group from which one or more subsidiary clades (monophyletic groups) are excluded to form a separate group. Philosopher of science Marc Ereshefsky has argued that paraphyletic taxa are the result of anagenesis in the excluded group or groups. A cladistic approach normally does not grant paraphyletic assemblages

440-419: A group of dinosaurs (part of Diapsida ), both of which are "reptiles". Osteichthyes , bony fish, are paraphyletic when circumscribed to include only Actinopterygii (ray-finned fish) and Sarcopterygii (lungfish, etc.), and to exclude tetrapods ; more recently, Osteichthyes is treated as a clade, including the tetrapods. The " wasps " are paraphyletic, consisting of the narrow-waisted Apocrita without

495-439: A kind of lizard). Put another way, viviparity is a synapomorphy for Theria within mammals, and an autapomorphy for Eulamprus tympanum (or perhaps a synapomorphy, if other Eulamprus species are also viviparous). Groupings based on independently-developed traits such as these examples of viviparity represent examples of polyphyly , not paraphyly. The following list recapitulates a number of paraphyletic groups proposed in

550-560: A more inclusive clade, it often makes sense to study the paraphyletic group that remains without considering the larger clade. For example, the Neogene evolution of the Artiodactyla (even-toed ungulates, like deer, cows, pigs and hippopotamuses - Cervidae , Bovidae , Suidae and Hippopotamidae , the families that contain these various artiodactyls, are all monophyletic groups) has taken place in environments so different from that of

605-424: A phylogenetic species concept that does not consider species to exhibit the properties of monophyly or paraphyly, concepts under that perspective which apply only to groups of species. They consider Zander's extension of the "paraphyletic species" argument to higher taxa to represent a category error When the appearance of significant traits has led a subclade on an evolutionary path very divergent from that of

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660-495: 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

715-569: 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

770-433: Is allowed as a synonym of Magnoliopsida. Phylogenetic analysis indicates that the monocots are a development from a dicot ancestor. Excluding monocots from the dicots makes the latter a paraphyletic group. Among animals, several familiar groups are not, in fact, clades. The order Artiodactyla ( even-toed ungulates ) as traditionally defined is paraphyletic because it excludes Cetaceans (whales, dolphins, etc.). Under

825-479: 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

880-629: Is rather arbitrary, since the character states of common ancestors are inferences, not observations. These terms were developed during the debates of the 1960s and 1970s accompanying the rise of cladistics . Paraphyletic groupings are considered problematic by many taxonomists, as it is not possible to talk precisely about their phylogenetic relationships, their characteristic traits and literal extinction. Related terms are stem group , chronospecies , budding cladogenesis, anagenesis, or 'grade' groupings. Paraphyletic groups are often relics from outdated hypotheses of phylogenic relationships from before

935-548: Is said to be polyparaphyletic. The term received currency during the debates of the 1960s and 1970s accompanying the rise of cladistics , having been coined by zoologist Willi Hennig to apply to well-known taxa like Reptilia ( reptiles ), which is paraphyletic with respect to birds . Reptilia contains the last common ancestor of reptiles and all descendants of that ancestor except for birds. Other commonly recognized paraphyletic groups include fish , monkeys , and lizards . The term paraphyly , or paraphyletic , derives from

990-482: Is sometimes used for paraphyletic groups. Moreover, the concepts of monophyly , paraphyly, and polyphyly have been used in deducing key genes for barcoding of diverse group of species. Current phylogenetic hypotheses of tetrapod relationships imply that viviparity , the production of offspring without the external laying of a fertilized egg, developed independently in the lineages that led to humans ( Homo sapiens ) and southern water skinks ( Eulampus tympanum ,

1045-514: The Cetacea (whales, dolphins, and porpoises) that the Artiodactyla are often studied in isolation even though the cetaceans are a descendant group. The prokaryote group is another example; it is paraphyletic because it is composed of two Domains (Eubacteria and Archaea) and excludes (the eukaryotes ). It is very useful because it has a clearly defined and significant distinction (absence of

1100-581: The ICN ) abandoned consideration of bacterial nomenclature in 1975; currently, prokaryotic nomenclature is regulated under the ICNB with a starting date of 1 January 1980 (in contrast to a 1753 start date under the ICBN/ICN). Among plants, dicotyledons (in the traditional sense) are paraphyletic because the group excludes monocotyledons . "Dicotyledon" has not been used as a botanic classification for decades, but

1155-861: 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

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1210-418: 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

1265-660: The ants and bees . The sawflies ( Symphyta ) are similarly paraphyletic, forming all of the Hymenoptera except for the Apocrita, a clade deep within the sawfly tree. Crustaceans are not a clade because the Hexapoda (insects) are excluded. The modern clade that spans all of them is the Tetraconata . One of the goals of modern taxonomy over the past fifty years has been to eliminate paraphyletic "groups", such as

1320-1741: 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

1375-453: 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

1430-484: 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

1485-657: 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

1540-421: 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

1595-963: The diplomonads, Entamoeba , Microsporidia , oxymonads, parabasalids ( Parabasalids ), pelobionts (see Pelomyxa ), retortamonads, trichomonads, and Trimastix (see Cavalier-Smith's system of classification ). With the rejection of 'Archeozoa', the meaning of the term 'Metakaryota' became the same as 'Eukaryota' (see Eukaryote ), and Metakaryota became superfluous. Eukaryotic protists lacking mitochondria were discovered to have experienced secondary mitochondrial loss, meaning that their ancestors once possessed mitochondria but that these mitochondria had, over time, been transformed, reduced, or lost. In some of these organisms, mitochondria had degraded into simpler double-membrane bound organelles known as mitosomes and hydrogenosomes . Some of both types of organelles are known to have fully lost their genome. Initial discoveries found that amitochondriate organisms appeared to express mitochondrial Hsp60 and Hsp70 proteins from

1650-554: 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

1705-450: The examples given here, from formal classifications. Species have a special status in systematics as being an observable feature of nature itself and as the basic unit of classification. Some articulations of the phylogenetic species concept require species to be monophyletic, but paraphyletic species are common in nature, to the extent that they do not have a single common ancestor. Indeed, for sexually reproducing taxa, no species has

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1760-411: The excluded subgroups. In contrast, a monophyletic grouping (a clade ) includes a common ancestor and all of its descendants. The terms are commonly used in phylogenetics (a subfield of biology ) and in the tree model of historical linguistics . Paraphyletic groups are identified by a combination of synapomorphies and symplesiomorphies . If many subgroups are missing from the named group, it

1815-480: The fact that a monophyletic group includes organisms consisting of all the descendants of a unique common ancestor. By comparison, the term polyphyly , or polyphyletic , uses the Ancient Greek prefix πολύς ( polús ), meaning "many, a lot of", and refers to the fact that a polyphyletic group includes organisms arising from multiple ancestral sources. Groups that include all the descendants of

1870-453: 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

1925-466: 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

1980-504: The island of Taiwan . Reptile This is an accepted version of this page 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

2035-508: 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

2090-508: 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

2145-622: The literature, and provides the corresponding monophyletic taxa. The concept of paraphyly has also been applied to historical linguistics , where the methods of cladistics have found some utility in comparing languages. For instance, the Formosan languages form a paraphyletic group of the Austronesian languages because they consist of the nine branches of the Austronesian family that are not Malayo-Polynesian and are restricted to

2200-796: 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  –

2255-405: The nuclear DNA of the organism. This indicated that the ancestors of these organisms once possessed mitochondria which expressed these proteins, but that these genes had migrated to their nuclear DNA over time as a result of endosymbiotic gene transfer. As a result, the argument that some extant eukaryotes lacking mitochondria had emerged from the eukaryotic lineage before mitochondria were acquired

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2310-422: 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,

2365-416: 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

2420-696: The ranks of the ICZN Code , the two taxa are separate orders. Molecular studies, however, have shown that the Cetacea descend from artiodactyl ancestors, although the precise phylogeny within the order remains uncertain. Without the Cetaceans the Artiodactyls are paraphyletic. The class Reptilia is paraphyletic because it excludes birds (class Aves ). Under a traditional classification, these two taxa are separate classes. However birds are sister taxon to

2475-438: 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

2530-567: The rise of cladistics. The prokaryotes (single-celled life forms without cell nuclei) are a paraphyletic grouping, because they exclude the eukaryotes , a descendant group. Bacteria and Archaea are prokaryotes, but archaea and eukaryotes share a common ancestor that is not ancestral to the bacteria. The prokaryote/eukaryote distinction was proposed by Edouard Chatton in 1937 and was generally accepted after being adopted by Roger Stanier and C.B. van Niel in 1962. The botanical code (the ICBN, now

2585-460: The status of "groups", nor does it reify them with explanations, as in cladistics they are not seen as the actual products of evolutionary events. A group whose identifying features evolved convergently in two or more lineages is polyphyletic (Greek πολύς [ polys ], "many"). More broadly, any taxon that is not paraphyletic or monophyletic can be called polyphyletic. Empirically, the distinction between polyphyletic groups and paraphyletic groups

2640-789: 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

2695-477: The two Ancient Greek words παρά ( pará ), meaning "beside, near", and φῦλον ( phûlon ), meaning "genus, species", and refers to the situation in which one or several monophyletic subgroups of organisms (e.g., genera, species) are left apart from all other descendants of a unique common ancestor. Conversely, the term monophyly , or monophyletic , builds on the Ancient Greek prefix μόνος ( mónos ), meaning "alone, only, unique", and refers to

2750-590: 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

2805-650: 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

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2860-592: 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

2915-459: Was falsified. An argument for the Archezoa group was that amitochondriate protists appeared to branch off early on from the eukaryotic lineage in phylogenetic analyses. This corroborated the supposition that Archezoa were more closely linked to primitive eukaryotes that evolved prior to the endosymbiotic process that generated the mitochondria. However, this early divergence later turned out to be

2970-538: Was never universally accepted because of conflicting information, and was dropped when the contrary argument, that amitochondriates were descendants of eukaryotes with mitochondria, became dominant. Eukaryotes that eventually acquired a bacterial endosymbiont that became the mitochondria were placed in a taxonomic group which Cavalier-Smith called the Metakaryota, whereas the Archezoa represented an earlier paraphyletic group to which Cavalier-Smith variously assigned

3025-627: 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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