Wikipedia

Paraphyly

Paraphylyis ataxonomicterm describing a grouping that consists of the grouping'slast common ancestorand some but not all of its descendant lineages. The grouping is said to be paraphyleticwith respect tothe excluded subgroups. In contrast, amonophyleticgrouping (aclade) includes a common ancestor andallof its descendants.

In thisphylogenetic tree,the green group isparaphyletic;it is composed of a common ancestor (the lowest green vertical stem) and some of its descendants, but it excludes the blue group (a monophyletic group) whichdivergedfrom the green group.

The terms are commonly used inphylogenetics(a subfield ofbiology) and in thetree modelofhistorical linguistics.Paraphyletic groups are identified by a combination ofsynapomorphiesandsymplesiomorphies.If many subgroups are missing from the named group, it is said to be polyparaphyletic.

The term received currency during the debates of the 1960s and 1970s accompanying the rise ofcladistics,having been coined by zoologistWilli Hennigto apply to well-known taxa like Reptilia (reptiles), which is paraphyletic with respect tobirds.Reptilia contains the last common ancestor of reptiles and all descendants of that ancestor except for birds. Other commonly recognized paraphyletic groups includefish,monkeys,andlizards.[1][page needed]

Etymology

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The termparaphyly,orparaphyletic,derives from the twoAncient Greekwordsπαρά(pará), meaning "beside, near", andφῦλον(phûlon), meaning "genus, species",[2][3]and refers to the situation in which one or several monophyletic subgroups of organisms (e.g., genera, species) areleft apartfrom all other descendants of a unique common ancestor.

Conversely, the termmonophyly,ormonophyletic,builds on the Ancient Greek prefixμόνος(mónos), meaning "alone, only, unique",[2][3]and refers to the fact that a monophyletic group includes organisms consisting ofallthe descendants of auniquecommon ancestor.

By comparison, the termpolyphyly,orpolyphyletic,uses the Ancient Greek prefixπολύς(polús), meaning "many, a lot of",[2][3]and refers to the fact that a polyphyletic group includes organisms arising frommultipleancestral sources.

Phylogenetics

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Cladogramof theprimates,showing amonophyly(the simians, in yellow), aparaphyly(the prosimians, in blue, including the red patch), and apolyphyly(the night-active primates, thelorisesand thetarsiers,in red).

In cladistics

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Further information:Cladistics

Groups that include all the descendants of a common ancestor are said to bemonophyletic.A paraphyletic group is a monophyletic group from which one or more subsidiaryclades(monophyletic groups) are excluded to form a separate group. Philosopher of science Marc Ereshefsky has argued that paraphyletic taxa are the result ofanagenesisin the excluded group or groups.[4]A cladistic approach normally does not grant paraphyletic assemblages 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.[5]

A group whose identifying features evolvedconvergentlyin two or more lineages ispolyphyletic(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 is rather arbitrary, since the character states of common ancestors are inferences, not observations.[citation needed]

These terms were developed during the debates of the 1960s and 1970s accompanying the rise ofcladistics.

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.[6][7]Related terms arestem group,chronospecies,budding cladogenesis, anagenesis, or'grade'groupings. Paraphyletic groups are often relics from outdated hypotheses of phylogenic relationships from before the rise of cladistics.[8]

Examples

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Waspsare paraphyletic, consisting of thecladeApocritawithoutantsandbees,which are not usually considered to be wasps; thesawflies( "Symphyta") too are paraphyletic, as the Apocrita are nested inside the Symphytan clades.

Theprokaryotes(single-celled life forms without cell nuclei) are a paraphyletic grouping, because they exclude theeukaryotes,a descendant group.BacteriaandArchaeaare prokaryotes, but archaea and eukaryotes share a common ancestor that is not ancestral to the bacteria. The prokaryote/eukaryote distinction was proposed byEdouard Chattonin 1937[9]and was generally accepted after being adopted by Roger Stanier and C.B. van Niel in 1962. The botanical code (the ICBN, now theICN) abandoned consideration of bacterial nomenclature in 1975; currently, prokaryotic nomenclature is regulated under theICNBwith a starting date of 1 January 1980 (in contrast to a 1753 start date under the ICBN/ICN).[10]

Among plants,dicotyledons(in the traditional sense) are paraphyletic because the group excludesmonocotyledons."Dicotyledon" has not been used as a botanic classification for decades, but is allowed as a synonym of Magnoliopsida.[note 1]Phylogenetic analysis indicates that themonocotsare a development from adicotancestor. Excluding monocots from the dicots makes the latter a paraphyletic group.[11]

Among animals, several familiar groups are not, in fact, clades. The orderArtiodactyla(even-toed ungulates) as traditionally defined is paraphyletic because it excludesCetaceans(whales, dolphins, etc.). Under the ranks of theICZN 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.[12] The classReptiliais paraphyletic because it excludes birds (classAves). Under a traditional classification, these two taxa are separate classes. However birds are sister taxon to a group of dinosaurs (part ofDiapsida), both of which are "reptiles".[13]

Osteichthyes,bony fish, are paraphyletic when circumscribed to include onlyActinopterygii(ray-finned fish) andSarcopterygii(lungfish, etc.), and to excludetetrapods;more recently, Osteichthyes is treated as a clade, including the tetrapods.[14][15]

The "wasps"are paraphyletic, consisting of the narrow-waistedApocritawithout theantsandbees.[16]The sawflies (Symphyta) are similarly paraphyletic, forming all of theHymenopteraexcept for the Apocrita, a clade deep within the sawfly tree.[14] Crustaceansare not a clade because theHexapoda(insects) are excluded. The modern clade that spans all of them is theTetraconata.[17][18]

One of the goals of modern taxonomy over the past fifty years has been to eliminate paraphyletic "groups", such as the examples given here, from formal classifications.[19][20]

Paraphyly in species

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Main article:Paraspecies

Specieshave a special status in systematics as being an observable feature of nature itself and as the basic unit of classification.[21]Some articulations of thephylogenetic species conceptrequire 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 a "single common ancestor" organism. Paraphyly is common inspeciation,whereby a mother species (aparaspecies) gives rise to a daughter species without itself becoming extinct.[22]Research indicates as many as 20 percent of all animal species and between 20 and 50 percent of plant species are paraphyletic.[23][24]Accounting for these facts, some taxonomists argue that paraphyly is a trait of nature that should be acknowledged at higher taxonomic levels.[25][26]

Cladists advocate a phylogenetic species concept[27]that does not consider species to exhibit the properties of monophyly or paraphyly, concepts under that perspective which apply only to groups of species.[28]They consider Zander's extension of the "paraphyletic species" argument to higher taxa to represent acategory error[29]

Uses for paraphyletic groups

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When the appearance of significant traits has led a subclade on an evolutionary path very divergent from that of a more inclusive clade, it often makes sense to study the paraphyletic group that remains without considering the larger clade. For example, theNeogeneevolution of theArtiodactyla(even-toed ungulates, like deer, cows, pigs and hippopotamuses -Cervidae,Bovidae,SuidaeandHippopotamidae,the families that contain these various artiodactyls, are all monophyletic groups) has taken place in environments so different from that of theCetacea(whales, dolphins, and porpoises) that the Artiodactyla are often studied in isolation even though the cetaceans are a descendant group. Theprokaryotegroup is another example; it is paraphyletic because it is composed of two Domains (Eubacteria and Archaea) and excludes (theeukaryotes). It is very useful because it has a clearly defined and significant distinction (absence of a cell nucleus, aplesiomorphy) from its excluded descendants.[citation needed]

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.[30]Other systematists consider reification of paraphyletic groups to obscure inferred patterns of evolutionary history.[31]

The term "evolutionary grade"is sometimes used for paraphyletic groups.[32] Moreover, the concepts ofmonophyly,paraphyly, andpolyphylyhave been used in deducing key genes forbarcodingof diverse group of species.[33]

Independently evolved traits

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Current phylogenetic hypotheses of tetrapod relationships imply thatviviparity,the production of offspring without the external laying of a fertilized egg, developedindependentlyin the lineages that led to humans (Homo sapiens) andsouthern water skinks(Eulampus tympanum,a kind of lizard). Put another way, viviparity is a synapomorphy forTheriawithin mammals, and an autapomorphy forEulamprus tympanum(or perhaps a synapomorphy, if otherEulamprusspecies are also viviparous).[citation needed]

Groupings based on independently-developed traits such as these examples of viviparity represent examples ofpolyphyly,not paraphyly.[citation needed]

Not paraphyly

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Further information:Convergent evolution
  • Amphibious fishare polyphyletic, not paraphyletic. Although they appear similar, several different groups of amphibious fishes such asmudskippersandlungfishesevolved independently in a process ofconvergent evolutionin distant relatives faced with similar ecological circumstances.[34]
  • Flightless birdsare polyphyletic because they independently (in parallel) lost the ability to fly.[35]
  • Animals with adorsal finare not paraphyletic, even though their last common ancestor may have had such a fin, because theMesozoicancestors of porpoises did not have such a fin, whereas pre-Mesozoic fish did have one.
  • Quadrupedalarchosaursare not a paraphyletic group.Bipedaldinosaurs likeEoraptor,ancestral to quadrupedal ones, were descendants of the last common ancestor of quadrupedal dinosaurs and other quadrupedal archosaurs like the crocodilians.

Non-exhaustive list of paraphyletic groups

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The following list recapitulates a number of paraphyletic groups proposed in the literature, and provides the corresponding monophyletic taxa.

Paraphyletic taxon Excluded clades Corresponding monophyletic taxon References and notes
Prokaryotes Eukaryota Cellular organisms [36]
Protista Animalia,Plantae,andFungi Eukaryota [37]
Invertebrates Vertebrata Animalia [38]
Worm Multiple groups Nephrozoa [39][40]
Radiata Bilateria Eumetazoa [41]
Platyzoa Lophotrochozoa,Mesozoa Spiralia [42]
Fish Tetrapoda Vertebrate [43]
Reptiles Birds Sauropsida [44]
Nonavian Dinosaur Birds Dinosauria
Lizard Snakes,Amphisbaenia Squamates [45]
Plagiaulacidans Cimolodonta,Arginbaataridae Multituberculata [46]
Pelycosaurs Therapsida Synapsida [47]
Even-toed ungulates Cetacea Cetartiodactyla [12][48]
Archaeoceti Neoceti Cetacea [49]
Prosimians Simiiformes Primates [50]
Crustaceans Hexapoda Tetraconata [17][18]
Wasps Ants,Bees Apocrita [16]
Sawfly Apocrita Hymenoptera [14]
Vespoidea Apoidea,Ants Euaculeata [16]
Parasitica Aculeata Apocrita [51]
Nautiloidea Ammonoidea,Coleoidea Cephalopoda [52]
Charophyte Embryophyte(Land plants) Streptophyta [53]
Dicotyledon Monocotyledon Angiosperm [11]
Moth Butterfly Lepidoptera [54]
Coral Medusozoa,Myxozoa Cnidaria [55][56]
Jellyfish Hydroidolina Medusozoa [57][58][59]
Cycloneuralia Panarthropoda Ecdysozoa [60][61]
Rotifera Acanthocephala Syndermata [62][63]
Mecoptera Siphonaptera Mecopteroidea
Anthoathecata Leptothecata,Siphonophorae Hydroidolina
Monkey Hominoidea Simiiformes [64][65]
Antelope Bovini,Caprinae Bovidae [66]

Linguistics

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Main article:Tree model

The concept of paraphyly has also been applied tohistorical linguistics,where the methods ofcladisticshave found some utility in comparing languages. For instance, theFormosan languagesform a paraphyletic group of theAustronesian languagesbecause they consist of the nine branches of the Austronesian family that are notMalayo-Polynesianand are restricted to the island ofTaiwan.[67]

See also

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Explanatory notes

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  1. ^The history of flowering plant classification can be found underHistory of the classification of flowering plants.

References

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