Phylogenetic nomenclature
Encyclopedia
Phylogenetic nomenclature (PN) or phylogenetic taxonomy
Taxonomy
Taxonomy is the science of identifying and naming species, and arranging them into a classification. The field of taxonomy, sometimes referred to as "biological taxonomy", revolves around the description and use of taxonomic units, known as taxa...

is an alternative to rank-based nomenclature
Biological classification
Biological classification, or scientific classification in biology, is a method to group and categorize organisms by biological type, such as genus or species. Biological classification is part of scientific taxonomy....

, applying definitions from cladistics
Cladistics
Cladistics is a method of classifying species of organisms into groups called clades, which consist of an ancestor organism and all its descendants . For example, birds, dinosaurs, crocodiles, and all descendants of their most recent common ancestor form a clade...

 (or phylogenetic systematics
Systematics
Biological systematics is the study of the diversification of terrestrial life, both past and present, and the relationships among living things through time. Relationships are visualized as evolutionary trees...

). Its two defining features are the use of phylogenetic
Phylogenetics
In biology, phylogenetics is the study of evolutionary relatedness among groups of organisms , which is discovered through molecular sequencing data and morphological data matrices...

 definitions
of biological taxon
Taxon
|thumb|270px|[[African elephants]] form a widely-accepted taxon, the [[genus]] LoxodontaA taxon is a group of organisms, which a taxonomist adjudges to be a unit. Usually a taxon is given a name and a rank, although neither is a requirement...

 names, and the lack of obligatory ranks. It is currently not regulated, but the PhyloCode
PhyloCode
The International Code of Phylogenetic Nomenclature, known as the PhyloCode for short, is a developing draft for a formal set of rules governing phylogenetic nomenclature...

(International Code of Phylogenetic Nomenclature) is intended to regulate it once implemented.

The terms cladism and cladist were first introduced by Ernst W. Mayr in 1965. They sometimes refer to cladistics as a whole.

Definitions

Under the rank-based codes of biological nomenclature
Nomenclature Codes
Nomenclature codes or codes of nomenclature are the various rulebooks that govern biological taxonomic nomenclature, each in their own broad field of organisms...

, names themselves do not have definitions, but are instead usually linked to a type. Some biologists have claimed that this is unsatisfactory and that instability in nomenclature should only reflect instability of our knowledge of phylogeny, not instability in subjective opinions about which ranks should be given to which groups. Phylogenetic nomenclature, on the other hand, uses phylogenetic definitions to tie a name to a clade
Clade
A clade is a group consisting of a species and all its descendants. In the terms of biological systematics, a clade is a single "branch" on the "tree of life". The idea that such a "natural group" of organisms should be grouped together and given a taxonomic name is central to biological...

, a group consisting solely of a species and all its descendants, in such a manner that the meaning of the name is objective under any phylogenetic hypothesis. This prevents splitting and lumping (unless definitions are changed in the process, which will be allowed under the PhyloCode only under carefully restricted circumstances).

Traditionally, groups named in phylogenetic nomenclature are usually monophyletic-that is, they define a natural group made up of all descendants of a single common ancestor. However, it is also possible to create phylogenetic definitions for the names of paraphyletic
Paraphyly
A group of taxa is said to be paraphyletic if the group consists of all the descendants of a hypothetical closest common ancestor minus one or more monophyletic groups of descendants...

 taxa . Assuming Mammalia and Aves are defined, Reptilia could be defined as "the most recent common ancestor of birds and mammals and all its descendants except birds and mammals". This includes taxa that are not currently named and even taxa that cannot be named under the rank-based codes
Nomenclature Codes
Nomenclature codes or codes of nomenclature are the various rulebooks that govern biological taxonomic nomenclature, each in their own broad field of organisms...

 without seriously disrupting existing classifications, such as "all organisms that share a more recent common ancestor with Homo sapiens than with birds and plesiomorphically keep laying eggs". Names of polyphyletic
Polyphyly
A polyphyletic group is one whose members' last common ancestor is not a member of the group.For example, the group consisting of warm-blooded animals is polyphyletic, because it contains both mammals and birds, but the most recent common ancestor of mammals and birds was cold-blooded...

 taxa could be defined by referring to the sum of two or more clades or paraphyletic taxa .

Philosophy

Rank-based and phylogenetic nomenclature differ in philosophical outlook. This manifests itself, particularly, in different approaches to the definitions of taxonomic terms. In providing the definition of "mammal
Mammal
Mammals are members of a class of air-breathing vertebrate animals characterised by the possession of endothermy, hair, three middle ear bones, and mammary glands functional in mothers with young...

," for example, users of rank-based nomenclature will start with the extension of the term that they want to define, perhaps the collection consisting of all animals with hair and mammary glands, and then formulate a definition satisfied by exactly these animals; users of phylogenetic nomenclature, rather, will formulate a definition, perhaps "the least inclusive clade containing brown bear
Brown Bear
The brown bear is a large bear distributed across much of northern Eurasia and North America. It can weigh from and its largest subspecies, the Kodiak Bear, rivals the polar bear as the largest member of the bear family and as the largest land-based predator.There are several recognized...

s and short-beaked echidna
Short-beaked Echidna
The short-beaked echidna , also known as the spiny anteater because of its diet of ants and termites, is one of four living species of echidna and the only member of the genus Tachyglossus...

s," and stipulate that it defines the word "mammal."

Put another way, users of a rank-based system name species and classify them into higher taxa, only some of which are clades, while proponents of phylogenetic nomenclature focus on clades and name them as entities that are of interest in their own right.

The analogy with chemistry

Proponents of phylogenetic nomenclature claim that, as rank-based nomenclature does not delimit taxa precisely, its definitions will need to change as the science of biology advances. They see their approach as preferable in that their definitions are not susceptible to this kind of instability. It is a disputed matter whether such changes are characteristic of sciences outside of biology and whether, if so, biology should follow the example of such sciences. Chemistry has been used as an example.

Michel Laurin
Michel Laurin
Michel Laurin is a Canadian vertebrate paleontologist specialising in the origin and phylogeny of tetrapods, comparative biology and paleobiology. As an undergraduate he worked in the laboratory of Robert L. Carroll, and took his doctor thesis on the osteology of seymouriamorphs under Robert R....

, one of the foremost advocates of phylogenetic nomenclature, considers that the concept of a chemical element has been stable ever since Dmitri Mendeleev
Dmitri Mendeleev
Dmitri Ivanovich Mendeleev , was a Russian chemist and inventor. He is credited as being the creator of the first version of the periodic table of elements...

 put forth the periodic table
Periodic table
The periodic table of the chemical elements is a tabular display of the 118 known chemical elements organized by selected properties of their atomic structures. Elements are presented by increasing atomic number, the number of protons in an atom's atomic nucleus...

 in 1869. Biology should, on Laurin’s view, follow the example of chemistry and define its terms as precisely as possible.

The historian and philosopher Thomas Kuhn
Thomas Kuhn
Thomas Samuel Kuhn was an American historian and philosopher of science whose controversial 1962 book The Structure of Scientific Revolutions was deeply influential in both academic and popular circles, introducing the term "paradigm shift," which has since become an English-language staple.Kuhn...

 argued that changing the meaning of established concepts is central to significant advances in science. Prior to John Dalton
John Dalton
John Dalton FRS was an English chemist, meteorologist and physicist. He is best known for his pioneering work in the development of modern atomic theory, and his research into colour blindness .-Early life:John Dalton was born into a Quaker family at Eaglesfield, near Cockermouth, Cumberland,...

’s work, he pointed out, the criteria for something’s being a chemical compound were such as to include salt water; by new criteria adopted afterwards, this fluid was excluded.

Michael Benton
Michael Benton
Michael J. Benton is a British paleontologist, a Fellow of the Royal Society of Edinburgh, and professor of vertebrate palaeontology in the Department of Earth Sciences at the University of Bristol....

, a prominent defender of rank-based nomenclature, regards biology as an endeavor very different from chemistry. Chemical classification, as he sees it, circumscribes entities in terms of properties that enter into knowable laws; biology, lacking such laws, must look to the usefulness of classifications. From this perspective, he argues, it is less important that the definition of a term classifying organisms remain constant than it is that the term continue to apply to most of the same organisms. As he views phylogenetic nomenclature as seeking the former kind of stability and rank-based nomenclature as seeking the latter, he considers rank-based nomenclature to be preferable.

Lack of ranks

The current codes of biological nomenclature stipulate that taxa cannot be given a valid name without being given a rank. However, the number of generally recognized ranks is limited. Gauthier et al. (1988) claimed that a classification which uses the common array of ranks, while including Aves within Reptilia and keeping Reptilia at its traditional rank of class, is forced to demote Aves substantially, perhaps to the rank of genus. This despite the ~ 12,000 known species of extant and extinct birds that would have to be incorporated into such a genus. To reduce this problem, 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.

The current codes also each have rules saying that names must have certain endings if they are applied to taxa that have certain ranks. When a taxon changes rank from one classification to another, its name must change its suffix. Ereshefsky (1997:512) stated:
The Linnaean rule of assigning rank-specific suffices [sic] gives rise to even more confusing cases. Simpson (1963, 29–30) and Wiley (1981, 238) agree that the genus Homo belongs to a particular taxon. They disagree, however, on that taxon's rank. Acting in accord with the Linnaean system, they attach different suffixes to the root Homini [actually Homin-] and give the taxon in question different names: Wiley calls it 'Hominini' [tribe rank] and Simpson calls it 'Hominidae' [family rank]. Their disagreement does not stop there. Wiley believes that the taxon just cited is a part of a more inclusive taxon which is a family. Using the root Homini, and following the rules of the Linnaean system [more precisely, the zoological code], he names the more inclusive taxon 'Hominidae.' So for Wiley and Simpson, the name 'Hominidae' refers to two different taxa. In brief, the Linnaean system causes Wiley and Simpson to assign different names to what they agree is the same taxon, and it causes them to give the same name to what they agree are different taxa.


In phylogenetic nomenclature, ranks have no bearing on the spelling of taxon names (see e.g.; see also the PhyloCode
PhyloCode
The International Code of Phylogenetic Nomenclature, known as the PhyloCode for short, is a developing draft for a formal set of rules governing phylogenetic nomenclature...

). Ranks are, however, not altogether forbidden in 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 (e.g.).

History

Ultimately, phylogenetic nomenclature is a result of Darwin
Charles Darwin
Charles Robert Darwin FRS was an English naturalist. He established that all species of life have descended over time from common ancestry, and proposed the scientific theory that this branching pattern of evolution resulted from a process that he called natural selection.He published his theory...

's discovery that the diversity and history of life is best represented in tree-shaped diagrams. This discovery immediately led to changes in the existing classifications. For example, John Hogg
John Hogg (biologist)
John Hogg was a British naturalist who wrote about amphibians, birds, plants, and protist. In 1839 he became a member of the Royal Society....

 proposed the term Protoctista
Protist
Protists are a diverse group of eukaryotic microorganisms. Historically, protists were treated as the kingdom Protista, which includes mostly unicellular organisms that do not fit into the other kingdoms, but this group is contested in modern taxonomy...

 in 1860 for organisms that did not seem closely related to either animals or plants. In 1866, the controversial biologist Ernst Haeckel
Ernst Haeckel
The "European War" became known as "The Great War", and it was not until 1920, in the book "The First World War 1914-1918" by Charles à Court Repington, that the term "First World War" was used as the official name for the conflict.-Research:...

 for the first time reconstructed a single tree of all life (see figure) and immediately proceeded to translate it into a classification. This classification was rank-based, in accordance with the only code of biological nomenclature that existed at the time, but did not contain taxa that Haeckel considered polyphyletic
Polyphyly
A polyphyletic group is one whose members' last common ancestor is not a member of the group.For example, the group consisting of warm-blooded animals is polyphyletic, because it contains both mammals and birds, but the most recent common ancestor of mammals and birds was cold-blooded...

; in it, Haeckel introduced the rank of phylum
Phylum
In biology, a phylum The term was coined by Georges Cuvier from Greek φῦλον phylon, "race, stock," related to φυλή phyle, "tribe, clan." is a taxonomic rank below kingdom and above class. "Phylum" is equivalent to the botanical term division....

 which carries a connotation of monophyly
Clade
A clade is a group consisting of a species and all its descendants. In the terms of biological systematics, a clade is a single "branch" on the "tree of life". The idea that such a "natural group" of organisms should be grouped together and given a taxonomic name is central to biological...

 in its name.

Ever since it has been debated in which ways and to what extent the phylogeny of life should be used as a basis for its classification, with views ranging from "numerical taxonomy" (phenetics
Phenetics
In biology, phenetics, also known as taximetrics, is an attempt to classify organisms based on overall similarity, usually in morphology or other observable traits, regardless of their phylogeny or evolutionary relation. It is closely related to numerical taxonomy which is concerned with the use of...

) over "evolutionary taxonomy
Evolutionary taxonomy
Evolutionary taxonomy, evolutionary systematics or Darwinian classification is a branch of biological classification that seeks to classify organisms using a combination of phylogenetic relationship and overall similarity. This type of taxonomy considers taxa rather than single species, so that...

" (gradistics) to "phylogenetic systematics" (cladistics
Cladistics
Cladistics is a method of classifying species of organisms into groups called clades, which consist of an ancestor organism and all its descendants . For example, birds, dinosaurs, crocodiles, and all descendants of their most recent common ancestor form a clade...

 – today, the term "cladistics" is only used for the method of phylogeny reconstruction, but its inventor, Willi Hennig
Willi Hennig
Emil Hans Willi Hennig was a German biologist who is considered the founder of phylogenetic systematics, also known as cladistics. With his works on evolution and systematics he revolutionised the view of the natural order of beings...

, regarded this method as a mere tool for the purpose of classification). From the 1960s onwards, rankless classifications were occasionally proposed, but in general the principles of rank-based nomenclature were used by all three schools of thought.

Most of 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
Sauropsida
Sauropsida is a group of amniotes that includes all existing reptiles and birds and their fossil ancestors, including the dinosaurs, the immediate ancestors of birds...

, erected 40 years earlier by Huxley, to include the birds (Aves
Bird
Birds are feathered, winged, bipedal, endothermic , egg-laying, vertebrate animals. Around 10,000 living species and 188 families makes them the most speciose class of tetrapod vertebrates. They inhabit ecosystems across the globe, from the Arctic to the Antarctic. Extant birds range in size from...

) as well as part of Reptilia
Reptile
Reptiles are members of a class of air-breathing, ectothermic vertebrates which are characterized by laying shelled eggs , and having skin covered in scales and/or scutes. They are tetrapods, either having four limbs or being descended from four-limbed ancestors...

, and coined the new name Theropsida
Synapsid
Synapsids are a group of animals that includes mammals and everything more closely related to mammals than to other living amniotes. They are easily separated from other amniotes by having an opening low in the skull roof behind each eye, leaving a bony arch beneath each, accounting for their name...

 to include the mammals as well as another part of Reptilia, but did not give them ranks, and treated them exactly as if they had phylogenetic definitions, using neither contents nor diagnostic characters to decide whether a given animal should belong to Theropsida, Sauropsida, or something else once its phylogenetic position was agreed upon. Goodrich also opined that the name Reptilia should be abandoned once the phylogeny of the reptiles would be better known. The lack of compatibility of his scheme with the existing rank-based classifications (despite agreement on the phylogeny in all but details), and the lack of a method of phylogenetics
Phylogenetics
In biology, phylogenetics is the study of evolutionary relatedness among groups of organisms , which is discovered through molecular sequencing data and morphological data matrices...

 at this time, are the most likely reasons why Goodrich's suggestions were largely ignored.

The principle that only clade
Clade
A clade is a group consisting of a species and all its descendants. In the terms of biological systematics, a clade is a single "branch" on the "tree of life". The idea that such a "natural group" of organisms should be grouped together and given a taxonomic name is central to biological...

s (monophyletic taxa – an ancestor plus all its descendants) should be formally named became popular in the second half of the 20th century. It spread together with the methods for discovering clades (cladistics
Cladistics
Cladistics is a method of classifying species of organisms into groups called clades, which consist of an ancestor organism and all its descendants . For example, birds, dinosaurs, crocodiles, and all descendants of their most recent common ancestor form a clade...

) and is an integral part of phylogenetic systematics (see above). At the same time, it became apparent that the obligatory ranks that are part of the traditional systems of nomenclature produced problems. Some authors suggested abandoning them altogether, starting with Willi Hennig
Willi Hennig
Emil Hans Willi Hennig was a German biologist who is considered the founder of phylogenetic systematics, also known as cladistics. With his works on evolution and systematics he revolutionised the view of the natural order of beings...

's abandonment of his earlier proposal to define ranks as geological age classes.

The origin of phylogenetic nomenclature can be dated to 1986, when Jacques Gauthier
Jacques Gauthier
Jacques Armand Gauthier is a vertebrate paleontologist, comparative morphologist, and systematist, and one of the founders of the use of cladistics in biology....

 used phylogenetic definitions for the first time in a published work. 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 a schism
Schism (religion)
A schism , from Greek σχίσμα, skhísma , is a division between people, usually belonging to an organization or movement religious denomination. The word is most frequently applied to a break of communion between two sections of Christianity that were previously a single body, or to a division within...

 in the biologist community, "Gauthier suggested to two members of the ICZN
International Commission on Zoological Nomenclature
The International Commission on Zoological Nomenclature is an organization dedicated to "achieving stability and sense in the scientific naming of animals". Founded in 1895, it currently comprises 28 members from 20 countries, mainly practicing zoological taxonomists...

 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" (Laurin, 2008: 224). This led him, Kevin de Queiroz, and the botanist Philip Cantino to start drafting their own code of nomenclature, the PhyloCode
PhyloCode
The International Code of Phylogenetic Nomenclature, known as the PhyloCode for short, is a developing draft for a formal set of rules governing phylogenetic nomenclature...

, for regulating phylogenetic nomenclature.

Further reading

A few seminal publications not cited in the references are cited here. An exhaustive list of publications about phylogenetic nomenclature can be found on the website of the International Society for Phylogenetic Nomenclature
International Society for Phylogenetic Nomenclature
The International Society for Phylogenetic Nomenclature was established to encourage and facilitate the development and use of, and communication about, phylogenetic nomenclature. It organizes periodic scientific meetings and is overseeing the completion and implementation of the PhyloCode....

.
  • Bryant, Harold N. (1994). Comments on the phylogenetic definition of taxon names and conventions regarding the naming of crown clades. Syst. Biol. 43:124–129.
  • Cantino, Philip D., and Richard G. Olmstead (2008). Application of phylogenetically defined names does not require that every specifier be present on a tree. Syst. Biol. 57:157–160.
  • de Queiroz, Kevin (1992). Phylogenetic definitions and taxonomic philosophy. Biol. Philos. 7:295–313.
  • Gauthier, Jacques A., Arnold G. Kluge, and Timothy Rowe (1988). The early evolution of the Amniota. Pages 103–155 in Michael J. Benton (ed.): The Phylogeny and Classification of the Tetrapods, Volume 1: Amphibians, Reptiles, Birds. Syst. Ass. Spec. Vol. 35A. Clarendon Press, Oxford.
  • Gauthier, Jacques, David Cannatella, Kevin de Queiroz, Arnold G. Kluge, and Timothy Rowe (1989). Tetrapod phylogeny. Pages 337–353 in B. Fernholm, K. Bremer, and H. Jörnvall (eds.): The Hierarchy of Life. Elsevier Science B. V. (Biomedical Division), New York.
  • Ghiselin, M. T. (1984). "Definition," "character," and other equivocal terms. Syst. Zool. 33:104–110.
  • Keesey, T. Michael (2007). A mathematical approach to defining clade names, with potential applications to computer storage and processing. Zool. Scr.
    Zoologica Scripta
    Zoologica Scripta is a bi-monthly scientific journal on systematic zoology, published by Blackwell Publishing Limited, on behalf of the Norwegian Academy of Science and Letters and the Royal Swedish Academy of Sciences...

    36:607–621.
  • Laurin, Michel (2005). The advantages of phylogenetic nomenclature over Linnean nomenclature. Pages 67–97 in A. Minelli, G. Ortalli, and G. Sanga (eds): Animal Names. Instituto Veneto di Scienze, Lettere ed Arti; Venice.
  • Lee, Michael S. Y. (2005). Choosing reference taxa in phylogenetic nomenclature. Zool. Scr.
    Zoologica Scripta
    Zoologica Scripta is a bi-monthly scientific journal on systematic zoology, published by Blackwell Publishing Limited, on behalf of the Norwegian Academy of Science and Letters and the Royal Swedish Academy of Sciences...

    34:329–331.
  • Rowe, Timothy (1987). Definition and diagnosis in the phylogenetic system. Syst. Zool. 36:208–211.
  • Rowe, Timothy, and Jacques Gauthier (1992). Ancestry, paleontology and definition of the name Mammalia. Syst. Biol. 41:372–378.
  • Sereno, Paul C. (1998). A rationale for phylogenetic definitions, with application to the higher-level taxonomy of Dinosauria. N. Jb. Geol. Paläont. Abh. 210:41–83.
  • Sereno, Paul C. (1999). Definitions in phylogenetic taxonomy: critique and rationale. Syst. Biol. 48:329–351.
  • Sereno, Paul C. (2005). The Logical Basis of Phylogenetic Taxonomy [sic]. Syst. Biol. 54:595–619.
  • Taylor, Michael P. (2007). Phylogenetic definitions in the pre-PhyloCode era; implications for naming clades under the PhyloCode. PaleoBios 27:1–6.
  • Wilkinson, Mark (2006). Identifying stable reference taxa for phylogenetic nomenclature. Zool. Scr.
    Zoologica Scripta
    Zoologica Scripta is a bi-monthly scientific journal on systematic zoology, published by Blackwell Publishing Limited, on behalf of the Norwegian Academy of Science and Letters and the Royal Swedish Academy of Sciences...

    35:109–112.
  • Wyss, A. R., and J. Meng (1996). Application of phylogenetic taxonomy to poorly resolved crown clades: a stem-modified node-based definition of Rodentia. Syst. Biol. 45:559–568.
The source of this article is wikipedia, the free encyclopedia.  The text of this article is licensed under the GFDL.
 
x
OK