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49-410: Lobosa is a taxonomic group of amoebae in the phylum Amoebozoa . Most lobosans possess broad, bluntly rounded pseudopods , although one genus in the group, the recently discovered Sapocribrum, has slender and threadlike (filose) pseudopodia. In current classification schemes, Lobosa is a subphylum , composed mainly of amoebae that have lobose pseudopods but lack cilia or flagella . The group

98-413: A genus , for example, so there would have to be a genus for every common ancestor of the mammals and the birds. For such a genus to be monophyletic, it would have to include both the class Mammalia and the class Aves. For rank-based nomenclature, however, classes must include genera, not the other way around. The conflict between phylogenetic and traditional nomenclature represents differing opinions of

147-402: A rank cannot be named formally, because the application of a name to a taxon is based on both a type and a rank. Thus for example the "family" Hominidae uses the genus Homo as its type; its rank (family) is indicated by the suffix -idae (see discussion below). The requirement for a rank is a major difference between traditional and phylogenetic nomenclature. It has several consequences: it limits

196-463: A schism among the systematics community, " Gauthier suggested to two members of the ICZN to apply formal taxonomic names ruled by the zoological code only to clades (at least for supraspecific taxa) and to abandon Linnean ranks, but these two members promptly rejected these ideas". The premise of names in traditional nomenclature is based, ultimately, on type specimens, and the circumscription of groups

245-489: A clade are not the same issue. Monophyletic groups are worthy of attention and naming because they share properties of interest -- synapomorphies -- that are the evidence that allows inference of common ancestry. Phylogenetic nomenclature is a semantic extension of the general acceptance of the idea of branching during the course of evolution, represented in the diagrams of Jean-Baptiste Lamarck and later writers like Charles Darwin and Ernst Haeckel . In 1866, Haeckel for

294-428: A controversy about the relative merits of phylogenetic nomenclature versus Linnaean taxonomy, or the related method of evolutionary taxonomy , which has continued to the present. Some of the controversies with which the cladists were engaged had been happening since the 19th century. While Hennig insisted that different classification schemes were useful for different purposes, he gave primacy to his own, claiming that

343-825: A different method. They retained the traditional ranks of family and order, considering them to be of value for teaching and studying relationships between taxa, but also introduced named clades without formal ranks. For phylogenetic nomenclature, ranks have no bearing on the spelling of taxon names (see e.g. Gauthier (1994) and the PhyloCode ). Ranks are, however, not altogether forbidden for phylogenetic nomenclature. They are merely decoupled from nomenclature: they do not influence which names can be used, which taxa are associated with which names, and which names can refer to nested taxa. The principles of traditional rank-based nomenclature are incompatible logically with all taxa being strictly monophyletic. Every organism must belong to

392-407: A group has a different rank in different classifications, its name must have a different suffix. Ereshefsky (1997:512) gave an example. He noted that Simpson in 1963 and Wiley in 1981 agreed that the same group of genera, which included the genus Homo , should be placed together in a taxon. Simpson treated this taxon as a family, and so gave it the name "Hominidae": "Homin-" from "Homo" and "-idae" as

441-549: A lower rank than any named taxon in which it is nested, so the number of names that can be assigned in a nested set of taxa can be no greater than the number of generally recognized ranks. Gauthier et al. (1988) suggested that, if Reptilia is assigned its traditional rank of "class", then a phylogenetic classification has to assign the rank of genus to Aves. In such a classification, all ~12,000 known species of extant and extinct birds would then have to be incorporated into this genus. Various solutions have been proposed while keeping

490-613: A paraphyletic group that includes some Permian tetrapods but not their extant descendants, it cannot be admitted as a valid taxon name. Again, while not disagreeing with the notion that only monophyletic groups should be named, empiricist systematists counter this ancestry essentialism by pointing out that pelycosaurs are recognized as paraphyletic precisely because they exhibit a combination of synapomorphies and symplesiomorphies indicating that some of them are more closely related to mammals than they are to other pelycosaurs. The material existence of an assemblage of fossils and its status as

539-449: A priori premises. If there are no characteristics that allow scientists to recognize a fossil as belonging to a taxonomic group, then it is just an unclassifiable piece of rock. If ancestry is sufficient for the continuity of a taxon, then all descendants of a taxon member will also be included in the taxon, so all bona fide taxa are monophyletic; the names of paraphyletic groups do not merit formal recognition. As " Pelycosauria " refers to

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588-727: Is paraphyletic , consisting of a grade of three clades: Discosea , Tubulinea and Cutosea . The first two are part of a paraphyletic superclass Glycopoda , while the latter constitutes the monophyletic superclass Cutosa . The clade uniting Tubulinea + Cutosea + Conosa is named Tevosa , while the clade uniting Cutosa + Conosa is named Evosea . Flabellinia Centramoebia Corycidia Echinamoebia Elardia Cutosea Archamoebea Variosea Dictyostelea Ceratiomyxea Myxogastrea Opisthokonta Taxon Initial attempts at classifying and ordering organisms (plants and animals) were presumably set forth in prehistoric times by hunter-gatherers, as suggested by

637-701: Is a method of nomenclature for taxa in biology that uses phylogenetic definitions for taxon names as explained below. This contrasts with the traditional method , by which taxon names are defined by a type , which can be a specimen or a taxon of lower rank , and a description in words. Phylogenetic nomenclature is regulated currently by the International Code of Phylogenetic Nomenclature ( PhyloCode ). Phylogenetic nomenclature associates names with clades , groups consisting of an ancestor and all its descendants. Such groups are said to be monophyletic . There are slightly different methods of specifying

686-548: Is an example of a paraphyletic group, a clade minus one or more subordinate clades. Names of polyphyletic groups, characterized by a trait that evolved convergently in two or more subgroups, can be defined similarly as the sum of multiple clades. Using the traditional nomenclature codes , such as the International Code of Zoological Nomenclature and the International Code of Nomenclature for algae, fungi, and plants , taxa that are not associated explicitly with

735-528: Is considered a taxonomic choice made by the systematists working on particular groups, rather than a nomenclatural decision made based on a priori rules of the Codes on Nomenclature . The desire to subsume taxonomic circumscriptions within nomenclatural definitions caused Kevin de Queiroz and the botanist Philip Cantino to start drafting their own code of nomenclature, the PhyloCode , to regulate phylogenetic nomenclature. Willi Hennig 's pioneering work provoked

784-554: Is included in the Reptilia (birds are traditionally placed in the class Aves , and mammals in the class Mammalia ). The term taxon was first used in 1926 by Adolf Meyer-Abich for animal groups, as a back-formation from the word taxonomy ; the word taxonomy had been coined a century before from the Greek components τάξις ( táxis ), meaning "arrangement", and νόμος ( nómos ), meaning " method ". For plants, it

833-461: Is no attempt to be exhaustive. Phylogenetic nomenclature allows the use, not only of ancestral relations , but also of the property of being extant . One of the many methods of specifying the Neornithes (modern birds), for example, is: Neornithes is a crown clade , a clade for which the last common ancestor of its extant members is also the last common ancestor of all its members. For

882-440: Is relative, and restricted to a particular systematic schema. For example, liverworts have been grouped, in various systems of classification, as a family, order, class, or division (phylum). The use of a narrow set of ranks is challenged by users of cladistics ; for example, the mere 10 ranks traditionally used between animal families (governed by the International Code of Zoological Nomenclature (ICZN)) and animal phyla (usually

931-491: Is to designate the ancestor. There are a number of methods of doing this. Commonly, the ancestor is indicated by its relation to two or more specifiers (species, specimens, or traits) that are mentioned explicitly. The diagram shows three common ways of doing this. For previously defined clades A, B, and C, the clade X can be defined as: Several other alternatives are provided in the PhyloCode , (see below ) though there

980-489: The Foraminifera . The name Lobosa was chosen for these amoebae "as expressing the lobe-like character of their pseudopodial extensions". As currently defined, the subphylum Lobosa includes both shelled ( testate ) and naked amoebae (gymnamoebae), but excludes some organisms traditionally regarded as "lobosean", such as Pelomyxa and Entamoeba (Amoebozoa) and some Heterolobosea ( Excavata ). The subphylum Lobosa

1029-529: The PhyloCode , only a clade can receive a "phylogenetic definition", and this restriction is observed in the present article. However, it is also possible to create definitions for the names of other groups that are phylogenetic in the sense that they use only ancestral relations based on species or specimens. For example, assuming Mammalia and Aves (birds) are defined in this manner, Amniotes could be defined as "the most recent common ancestor of Mammalia and Aves and all its descendants except Mammalia and Aves". This

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1078-463: The clade , is equivalent to the taxon, assuming that taxa should reflect evolutionary relationships. Similarly, among those contemporary taxonomists working with the traditional Linnean (binomial) nomenclature, few propose taxa they know to be paraphyletic . An example of a long-established taxon that is not also a clade is the class Reptilia , the reptiles; birds and mammals are the descendants of animals traditionally classed as reptiles, but neither

1127-464: The metaphysics and epistemology of taxa. For the advocates of phylogenetic nomenclature, a taxon is an individual entity, an entity that may gain and lose attributes as time passes. Just as a person does not become somebody else when his or her properties change through maturation, senility, or more radical changes like amnesia, the loss of a limb, or a change of sex, so a taxon remains the same entity whatever characteristics are gained or lost. Given

1176-526: The Theropsidan or the Sauropsidan line of evolution." Goodrich concluded his paper: "The possession of these characters shows that all living Reptilia belong to the Sauropsidan group, while the structure of the foot enables us to determine the affinities of many incompletely known fossil genera, and to conclude that only certain extinct orders can belong to the Theropsidan branch." Goodrich opined that

1225-496: The ancestor, which are discussed below. Once the ancestor is specified, the meaning of the name is fixed: the ancestor and all organisms which are its descendants are included in the taxon named. Listing all these organisms (i.e. providing a full circumscription ) requires the complete phylogenetic tree to be known. In practice, there are almost always one or more hypotheses as to the correct relationship. Different hypotheses result in different organisms being thought to be included in

1274-607: The basic tenets of phylogenetic nomenclature (lack of obligatory ranks, and something close to phylogenetic definitions) can, however, be traced to 1916, when Edwin Goodrich interpreted the name Sauropsida , defined 40 years earlier by Thomas Henry Huxley , to include the birds ( Aves ) as well as part of Reptilia , and invented the new name Theropsida to include the mammals as well as another part of Reptilia. As these taxa were separate from traditional zoological nomenclature, Goodrich did not emphasize ranks, but he clearly discussed

1323-532: The body traces a continuous path through the world and it is this continuity, rather than any characteristics of the individual, that associates the baby with the octogenarian. This is similar to the well-known philosophical problem of the Ship of Theseus . For a taxon, IF characteristics are not relevant, THEN it can only be ancestral relations that associate the Devonian Rhyniognatha hirsti with

1372-541: The categories of his system had "individuality and reality" in contrast to the "timeless abstractions" of classifications based on overall similarity. Formal classifications based on cladistic reasoning are said to emphasize ancestry at the expense of descriptive characteristics. Nonetheless, most taxonomists presently avoid paraphyletic groups whenever they think it is possible within Linnaean taxonomy; polyphyletic taxa have long been unfashionable. Many cladists claim that

1421-410: The diagnostic features necessary to recognize and classify fossils belonging to the various groups. For example, in regard to the fifth metatarsal of the hind leg, he said "the facts support our view, for these early reptiles have normal metatarsals like their Amphibian ancestors. It is clear, then, that we have here a valuable corroborative character to help us to decide whether a given species belongs to

1470-483: The fairly sophisticated folk taxonomies. Much later, Aristotle, and later still, European scientists, like Magnol , Tournefort and Carl Linnaeus 's system in Systema Naturae , 10th edition (1758), , as well as an unpublished work by Bernard and Antoine Laurent de Jussieu , contributed to this field. The idea of a unit-based system of biological classification was first made widely available in 1805 in

1519-432: The first time constructed a single relational diagram of all life based on the existing classification of life accepted at the time. This classification was rank-based, but did not contain taxa that Haeckel considered polyphyletic . In it, Haeckel introduced the rank of phylum which carries a connotation of monophyly in its name (literally meaning "stem"). Ever since, it has been debated in which ways and to what extent

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1568-470: The highest relevant rank in taxonomic work) often cannot adequately represent the evolutionary history as more about a lineage's phylogeny becomes known. In addition, the class rank is quite often not an evolutionary but a phenetic or paraphyletic group and as opposed to those ranks governed by the ICZN (family-level, genus-level and species -level taxa), can usually not be made monophyletic by exchanging

1617-687: The introduction of Jean-Baptiste Lamarck 's Flore françoise , and Augustin Pyramus de Candolle 's Principes élémentaires de botanique . Lamarck set out a system for the "natural classification" of plants. Since then, systematists continue to construct accurate classifications encompassing the diversity of life; today, a "good" or "useful" taxon is commonly taken to be one that reflects evolutionary relationships . Many modern systematists, such as advocates of phylogenetic nomenclature , use cladistic methods that require taxa to be monophyletic (all descendants of some ancestor). Therefore, their basic unit,

1666-411: The known biodiversity. In fact, calls to reform biological nomenclature were made even before phylogenetic nomenclature was developed. The ICPN , or PhyloCode , is a code of rules and recommendations for phylogenetic nomenclature. The number of supporters for widespread adoption of the PhyloCode is still small, and it is uncertain how widely it will be followed. A few publications not cited in

1715-441: The metaphysical claims regarding unobservable entities made by advocates of phylogenetic nomenclature, critics have referred to their method as origin essentialism. For any individual, there has to be something that associates its temporal stages with each other by virtue of which it remains the same entity. For a person, the spatiotemporal continuity of the body provides the relevant conceptual continuity; from infancy to old age,

1764-399: The modern monarch butterfly as representatives, separated by 400 million years, of the taxon Insecta. The opposing opinion questions the premise of that syllogism, and argues, from an epistemological perspective, that members of taxa are only recognizable empirically on the basis of their observable characteristics, and hypotheses of common ancestry are results of theoretical systematics, not

1813-414: The name Reptilia should be abandoned once the phylogeny of the reptiles was better known. The principle that only clades should be named formally became popular among some researchers during the second half of the 20th century. It spread together with the methods for discovering clades ( cladistics ) and is an integral part of phylogenetic systematics (see above). At the same time, it became apparent that

1862-401: The named taxon, but application to the name in the context of various phylogenies generally remains unambiguous. Possible exceptions occur for apomorphy-based definitions, when optimization of the defining apomorphy is ambiguous. Phylogenetic nomenclature assigns names to clades , groups consisting solely of an ancestor and all its descendants. All that is needed to specify a clade, therefore,

1911-419: The number of nested levels at which names can be applied; it causes the endings of names to change if a group has its rank changed, even if it has precisely the same members (i.e. the same circumscription ); and it is logically inconsistent with all taxa being monophyletic. The current codes have rules stating that names must have certain endings depending on the rank of the taxa to which they are applied. When

1960-594: The obligatory ranks that are part of the traditional systems of nomenclature produced problems. Some authors suggested abandoning them altogether, starting with Willi Hennig 's abandonment of his earlier proposal to define ranks as geological age classes. The first use of phylogenetic nomenclature in a publication can be dated to 1986. Theoretical papers outlining the principles of phylogenetic nomenclature, as well as further publications containing applications of phylogenetic nomenclature (mostly to vertebrates), soon followed (see Literature section). In an attempt to avoid

2009-539: The rank-based nomenclature codes. Patterson and Rosen (1977) suggested nine new ranks between family and superfamily in order to be able to classify a clade of herrings, and McKenna and Bell (1997) introduced a large array of new ranks in order to cope with the diversity of Mammalia; these have not been adopted widely. For botany, the Angiosperm Phylogeny Group , responsible for the currently most widely used classification of flowering plants , chose

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2058-412: The same name had different circumscriptions (Simpson's Hominidae and Wiley's Hominidae). Especially in recent decades (due to advances in phylogenetics ), taxonomists have named many "nested" taxa (i.e. taxa which are contained inside other taxa). No system of nomenclature attempts to name every clade; this would be particularly difficult with traditional nomenclature since every named taxon must be given

2107-425: The suffix for family using the zoological code. Wiley considered it to be at the rank of "tribe", and so gave it the name "Hominini", "-ini" being the suffix for tribe. Wiley's tribe Hominini formed only part of a family which he termed "Hominidae". Thus, using the zoological code, two groups with precisely the same circumscription were given different names (Simpson's Hominidae and Wiley's Hominini), and two groups with

2156-489: The taxa contained therein. This has given rise to phylogenetic taxonomy and the ongoing development of the PhyloCode , which has been proposed as a new alternative to replace Linnean classification and govern the application of names to clades . Many cladists do not see any need to depart from traditional nomenclature as governed by the ICZN, International Code of Nomenclature for algae, fungi, and plants , etc. Phylogenetic nomenclature Phylogenetic nomenclature

2205-402: The traditional Codes of Zoological and Botanical Nomenclature are fully compatible with cladistic methods, and that there is no need to reinvent a system of names that has functioned well for 250 years, but others argue that this system is not as effective as it should be and that it is time to adopt nomenclatural principles that represent divergent evolution as a mechanism that explains much of

2254-438: The understanding of the phylogeny of life should be used as a basis for its classification, with opinions including "numerical taxonomy" ( phenetics ), " evolutionary taxonomy " (gradistics), and "phylogenetic systematics". From the 1960s onwards, rankless classifications were occasionally proposed, but in general the principles and common language of traditional nomenclature have been used by all three schools of thought. Most of

2303-420: Was always used for animals, whereas "division" was traditionally often used for plants , fungi , etc. A prefix is used to indicate a ranking of lesser importance. The prefix super- indicates a rank above, the prefix sub- indicates a rank below. In zoology , the prefix infra- indicates a rank below sub- . For instance, among the additional ranks of class are superclass, subclass and infraclass. Rank

2352-444: Was originally proposed in 1861 by William B. Carpenter , who created it as a taxonomic order containing the single family Amoebina. Carpenter's Lobosa consisted of amoeboid organisms whose endoplasm (endosarc) flows into lobe-like "pseudopodian prolongations." This type of pseudopod, which was understood to be typical of the genus Amoeba "and its allies," differed from the filose (thread-like) or reticulose (netlike) pseudopods of

2401-498: Was proposed by Herman Johannes Lam in 1948, and it was adopted at the VII International Botanical Congress , held in 1950. The glossary of the International Code of Zoological Nomenclature (1999) defines a A taxon can be assigned a taxonomic rank , usually (but not necessarily) when it is given a formal name. " Phylum " applies formally to any biological domain , but traditionally it

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