Taxonomy (biology)

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Ancient Greek τάξις (taxis) 'arrangement' and -νομία (-nomia) 'method') is the scientific study of naming, defining (circumscribing) and classifying groups of biological organisms based on shared characteristics. Organisms are grouped into taxa (singular: taxon) and these groups are given a taxonomic rank; groups of a given rank can be aggregated to form a more inclusive group of higher rank, thus creating a taxonomic hierarchy. The principal ranks in modern use are domain, kingdom, phylum (division is sometimes used in botany in place of phylum), class, order, family, genus, and species. The Swedish botanist Carl Linnaeus is regarded as the founder of the current system of taxonomy, as he developed a ranked system known as Linnaean taxonomy for categorizing organisms and binomial nomenclature
for naming organisms.

With advances in the theory, data and analytical technology of biological systematics, the Linnaean system has transformed into a system of modern biological classification intended to reflect the evolutionary relationships among organisms, both living and extinct.

Definition

The exact definition of taxonomy varies from source to source, but the core of the discipline remains: the conception, naming, and classification of groups of organisms.[1] As points of reference, recent definitions of taxonomy are presented below:

  1. Theory and practice of grouping individuals into species, arranging species into larger groups, and giving those groups names, thus producing a classification.[2]
  2. A field of science (and a major component of systematics) that encompasses description, identification, nomenclature, and classification[3]
  3. The science of classification, in biology the arrangement of organisms into a classification[4]
  4. "The science of classification as applied to living organisms, including the study of means of formation of species, etc."[5]
  5. "The analysis of an organism's characteristics for the purpose of classification"[6]
  6. "Systematics studies
    phylogeny to provide a pattern that can be translated into the classification and names of the more inclusive field of taxonomy" (listed as a desirable but unusual definition)[7]

The varied definitions either place taxonomy as a sub-area of systematics (definition 2), invert that relationship (definition 6), or appear to consider the two terms synonymous. There is some disagreement as to whether

biological nomenclature is considered a part of taxonomy (definitions 1 and 2), or a part of systematics outside taxonomy.[8][9] For example, definition 6 is paired with the following definition of systematics that places nomenclature outside taxonomy:[6]

  • Systematics: "The study of the identification, taxonomy, and nomenclature of organisms, including the classification of living things with regard to their natural relationships and the study of variation and the evolution of taxa".

In 1970, Michener et al. defined "systematic biology" and "taxonomy" (terms that are often confused and used interchangeably) in relation to one another as follows:[10]

Systematic biology (hereafter called simply systematics) is the field that (a) provides scientific names for organisms, (b) describes them, (c) preserves collections of them, (d) provides classifications for the organisms, keys for their identification, and data on their distributions, (e) investigates their evolutionary histories, and (f) considers their environmental adaptations. This is a field with a long history that in recent years has experienced a notable renaissance, principally with respect to theoretical content. Part of the theoretical material has to do with evolutionary areas (topics e and f above), the rest relates especially to the problem of classification. Taxonomy is that part of Systematics concerned with topics (a) to (d) above.

A whole set of terms including taxonomy, systematic biology,

alpha taxonomy, is more specifically the identification, description, and naming (i.e., nomenclature) of organisms,[15]
while "classification" focuses on placing organisms within hierarchical groups that show their relationships to other organisms.

Monograph and taxonomic revision

A taxonomic revision or taxonomic review is a novel analysis of the variation patterns in a particular

palynological, biochemical and genetic. A monograph or complete revision is a revision that is comprehensive for a taxon for the information given at a particular time, and for the entire world. Other (partial) revisions may be restricted in the sense that they may only use some of the available character sets or have a limited spatial scope. A revision results in a conformation of or new insights in the relationships between the subtaxa within the taxon under study, which may lead to a change in the classification of these subtaxa, the identification of new subtaxa, or the merger of previous subtaxa.[16]

Taxonomic characters

Taxonomic characters are the taxonomic attributes that can be used to provide the evidence from which relationships (the

phylogeny) between taxa are inferred.[17][18] Kinds of taxonomic characters include:[19]

Alpha and beta taxonomy

Not to be confused with Alpha diversity.

The term "alpha taxonomy" is primarily used to refer to the discipline of finding, describing, and naming taxa, particularly species.[20] In earlier literature, the term had a different meaning, referring to morphological taxonomy, and the products of research through the end of the 19th century.[21]

William Bertram Turrill introduced the term "alpha taxonomy" in a series of papers published in 1935 and 1937 in which he discussed the philosophy and possible future directions of the discipline of taxonomy.[22]

... there is an increasing desire amongst taxonomists to consider their problems from wider viewpoints, to investigate the possibilities of closer co-operation with their cytological, ecological and genetics colleagues and to acknowledge that some revision or expansion, perhaps of a drastic nature, of their aims and methods, may be desirable ... Turrill (1935) has suggested that while accepting the older invaluable taxonomy, based on structure, and conveniently designated "alpha", it is possible to glimpse a far-distant taxonomy built upon as wide a basis of morphological and physiological facts as possible, and one in which "place is found for all observational and experimental data relating, even if indirectly, to the constitution, subdivision, origin, and behaviour of species and other taxonomic groups". Ideals can, it may be said, never be completely realized. They have, however, a great value of acting as permanent stimulants, and if we have some, even vague, ideal of an "omega" taxonomy we may progress a little way down the Greek alphabet. Some of us please ourselves by thinking we are now groping in a "beta" taxonomy.[22]

Turrill thus explicitly excludes from alpha taxonomy various areas of study that he includes within taxonomy as a whole, such as ecology, physiology, genetics, and cytology. He further excludes phylogenetic reconstruction from alpha taxonomy.[23]

Later authors have used the term in a different sense, to mean the delimitation of species (not subspecies or taxa of other ranks), using whatever investigative techniques are available, and including sophisticated computational or laboratory techniques.[24][20] Thus, Ernst Mayr in 1968 defined "beta taxonomy" as the classification of ranks higher than species.[25]

An understanding of the biological meaning of variation and of the evolutionary origin of groups of related species is even more important for the second stage of taxonomic activity, the sorting of species into groups of relatives ("taxa") and their arrangement in a hierarchy of higher categories. This activity is what the term classification denotes; it is also referred to as "beta taxonomy".

Microtaxonomy and macrotaxonomy

Main article:
Species problem

How species should be defined in a particular group of organisms gives rise to practical and theoretical problems that are referred to as the

species problem. The scientific work of deciding how to define species has been called microtaxonomy.[26][27][20] By extension, macrotaxonomy is the study of groups at the higher taxonomic ranks subgenus and above,[20] or simply in clades that include more than one taxon considered a species, expressed in terms of phylogenetic nomenclature.[28]

History

While some descriptions of taxonomic history attempt to date taxonomy to ancient civilizations, a truly scientific attempt to classify organisms did not occur until the 18th century, with the possible exception of Aristotle, whose works hint at a taxonomy.[29][30] Earlier works were primarily descriptive and focused on plants that were useful in agriculture or medicine.

There are a number of stages in this scientific thinking. Early taxonomy was based on arbitrary criteria, the so-called "artificial systems", including Linnaeus's system of sexual classification for plants (Linnaeus's 1735 classification of animals was entitled "Systema Naturae" ("the System of Nature"), implying that he, at least, believed that it was more than an "artificial system").

Later came systems based on a more complete consideration of the characteristics of taxa, referred to as "natural systems", such as those of

Bentham and Hooker (1862–1863). These classifications described empirical patterns and were pre-evolutionary
in thinking.

The publication of

Eichler (1883) and Engler
(1886–1892).

The advent of

synapomorphies. Since then, the evidentiary basis has been expanded with data from molecular genetics that for the most part complements traditional morphology.[31][page needed][32][page needed][33][page needed
]

Pre-Linnaean

Early taxonomists

Naming and classifying human surroundings likely began with the onset of language. Distinguishing poisonous plants from edible plants is integral to the survival of human communities. Medicinal plant illustrations show up in Egyptian wall paintings from c. 1500 BC, indicating that the uses of different species were understood and that a basic taxonomy was in place.[34]

Ancient times

Further information: Aristotle's biology § Classification
Description of rare animals (写生珍禽图), by Song dynasty painter Huang Quan (903–965)

Organisms were first classified by Aristotle (Greece, 384–322 BC) during his stay on the Island of Lesbos.[35][36][37] He classified beings by their parts, or in modern terms attributes, such as having live birth, having four legs, laying eggs, having blood, or being warm-bodied.[38] He divided all living things into two groups: plants and animals.[36]

Some of his groups of animals, such as Anhaima (animals without blood, translated as

cetaceans, are commonly used.[39][40][41]

His student

Cornus, Crocus, and Narcissus.[36]

Medieval

Taxonomy in the

Aristotelian system,[38] with additions concerning the philosophical and existential order of creatures. This included concepts such as the great chain of being in the Western scholastic tradition,[38]
again deriving ultimately from Aristotle.

The Aristotelian system did not classify plants or

fungi, due to the lack of microscopes at the time,[37] as his ideas were based on arranging the complete world in a single continuum, as per the scala naturae (the Natural Ladder).[36] This, as well, was taken into consideration in the great chain of being.[36]

Advances were made by scholars such as Procopius, Timotheus of Gaza, Demetrios Pepagomenos, and Thomas Aquinas. Medieval thinkers used abstract philosophical and logical categorizations more suited to abstract philosophy than to pragmatic taxonomy.[36]

Renaissance and early modern

During the Renaissance and the Age of Enlightenment, categorizing organisms became more prevalent,[36] and taxonomic works became ambitious enough to replace the ancient texts. This is sometimes credited to the development of sophisticated optical lenses, which allowed the morphology of organisms to be studied in much greater detail.

One of the earliest authors to take advantage of this leap in technology was the Italian physician Andrea Cesalpino (1519–1603), who has been called "the first taxonomist".[42] His magnum opus De Plantis came out in 1583, and described more than 1,500 plant species.[43][44] Two large plant families that he first recognized are in use: the Asteraceae and Brassicaceae.[45]

In the 17th century, John Ray (England, 1627–1705) wrote many important taxonomic works.[37] Arguably his greatest accomplishment was Methodus Plantarum Nova (1682),[46] in which he published details of over 18,000 plant species. At the time, his classifications were perhaps the most complex yet produced by any taxonomist, as he based his taxa on many combined characters.

The next major taxonomic works were produced by Joseph Pitton de Tournefort (France, 1656–1708).[47] His work from 1700, Institutiones Rei Herbariae, included more than 9,000 species in 698 genera, which directly influenced Linnaeus, as it was the text he used as a young student.[34]

Linnaean era

Main article: Linnaean taxonomy
Title page of Systema Naturae, Leiden, 1735

The Swedish botanist

Linnaean system).[51]

Plant and animal taxonomists regard Linnaeus' work as the "starting point" for valid names (at 1753 and 1758 respectively).[52] Names published before these dates are referred to as "pre-Linnaean", and not considered valid (with the exception of spiders published in Svenska Spindlar[53]). Even taxonomic names published by Linnaeus himself before these dates are considered pre-Linnaean.[34]

The digital era of taxonomy

Modern taxonomy is heavily influenced by technology such as DNA sequencing, bioinformatics, databases, and imaging.

Modern system of classification

Main articles: Evolutionary taxonomy and Phylogenetic nomenclature
vertebrates at class level, width of spindles indicating number of families. Spindle diagrams are typical for evolutionary taxonomy
.
The same relationship, expressed as a cladogram typical for cladistics

A pattern of groups nested within groups was specified by Linnaeus' classifications of plants and animals, and these patterns began to be represented as

dendrograms of the animal and plant kingdoms toward the end of the 18th century, well before Charles Darwin's On the Origin of Species was published.[37] The pattern of the "Natural System" did not entail a generating process, such as evolution, but may have implied it, inspiring early transmutationist thinkers. Among early works exploring the idea of a transmutation of species were Zoonomia in 1796 by Erasmus Darwin (Charles Darwin's grandfather), and Jean-Baptiste Lamarck's Philosophie zoologique of 1809.[20] The idea was popularized in the Anglophone world by the speculative but widely read Vestiges of the Natural History of Creation, published anonymously by Robert Chambers in 1844.[54]

With Darwin's theory, a general acceptance quickly appeared that a classification should reflect the Darwinian principle of

Tree of life representations became popular in scientific works, with known fossil groups incorporated. One of the first modern groups tied to fossil ancestors was birds.[56] Using the then newly discovered fossils of Archaeopteryx and Hesperornis, Thomas Henry Huxley pronounced that they had evolved from dinosaurs, a group formally named by Richard Owen in 1842.[57][58] The resulting description, that of dinosaurs "giving rise to" or being "the ancestors of" birds, is the essential hallmark of evolutionary taxonomic thinking. As more and more fossil groups were found and recognized in the late 19th and early 20th centuries, palaeontologists worked to understand the history of animals through the ages by linking together known groups.[59] With the modern evolutionary synthesis of the early 1940s, an essentially modern understanding of the evolution of the major groups was in place. As evolutionary taxonomy is based on Linnaean taxonomic ranks, the two terms are largely interchangeable in modern use.[60]

The

synapomorphies, shared derived character states.[63]

Cladistic classifications are compatible with traditional Linnean taxonomy and the Codes of

Zoological and Botanical nomenclature, to a certain extent.[64] An alternative system of nomenclature, the International Code of Phylogenetic Nomenclature or PhyloCode has been proposed, which regulates the formal naming of clades.[65][28][9] Linnaean ranks are optional and have no formal standing under the PhyloCode, which is intended to coexist with the current, rank-based codes.[28] While popularity of phylogenetic nomenclature has grown steadily in the last few decades,[9] it remains to be seen whether a majority of systematists will eventually adopt the PhyloCode or continue using the current systems of nomenclature that have been employed (and modified, but arguably not as much as some systematists wish)[66][67]
for over 250 years.

Kingdoms and domains

The basic scheme of modern classification. Many other levels can be used; domain, the highest level within life, is both new and disputed.
Main articles: Kingdom (biology) and Domain (biology)

Well before Linnaeus, plants and animals were considered separate Kingdoms.[68][unreliable source?] Linnaeus used this as the top rank, dividing the physical world into the vegetable, animal and mineral kingdoms. As advances in microscopy made the classification of microorganisms possible, the number of kingdoms increased, five- and six-kingdom systems being the most common.

nucleus.[70] A small number of scientists include a sixth kingdom, Archaea, but do not accept the domain method.[68]

archaeobacteria as part of a subkingdom of the kingdom Bacteria, i.e., he rejected the three-domain system entirely.[72] Stefan Luketa in 2012 proposed a five "dominion" system, adding Prionobiota (acellular and without nucleic acid) and Virusobiota (acellular but with nucleic acid) to the traditional three domains.[73]

Linnaeus
1735[74] 		Haeckel
1866[75] 		Chatton
1925[76] 		Copeland
1938[77] 		Whittaker
1969[78] 		Woese et al.
1990[79] 		Cavalier-Smith
1998,[72] 2015[80]
2 kingdoms 3 kingdoms 2 empires 4 kingdoms 5 kingdoms 3 domains 2 empires,
6/7 kingdoms
(not treated) Protista
Prokaryota
 Monera Monera Bacteria Bacteria
Archaea Archaea (2015)
Eukaryota
Protoctista
 Protista Eucarya "Protozoa"
"Chromista"
Vegetabilia
 Plantae Plantae Plantae Plantae
Fungi Fungi
Animalia Animalia Animalia Animalia Animalia

Recent comprehensive classifications

Partial classifications exist for many individual groups of organisms and are revised and replaced as new information becomes available; however, comprehensive, published treatments of most or all life are rarer; recent examples are that of Adl et al., 2012 and 2019,

NCBI taxonomy database, the Interim Register of Marine and Nonmarine Genera, the Open Tree of Life, and the Catalogue of Life. The Paleobiology Database
is a resource for fossils.

Application

Biological taxonomy is a sub-discipline of biology, and is generally practiced by biologists known as "taxonomists", though enthusiastic naturalists are also frequently involved in the publication of new taxa.[85] Because taxonomy aims to describe and organize life, the work conducted by taxonomists is essential for the study of biodiversity and the resulting field of conservation biology.[86][87]

Classifying organisms

Main article: Taxonomic rank

Biological classification is a critical component of the taxonomic process. As a result, it informs the user as to what the relatives of the taxon are hypothesized to be. Biological classification uses taxonomic ranks, including among others (in order from most inclusive to least inclusive): domain, kingdom, phylum, class, order, family, genus, species, and strain.[88][note 1]

Taxonomic descriptions

See also: Species description
Type specimen for Nepenthes smilesii, a tropical pitcher plant

The "definition" of a taxon is encapsulated by its description or its diagnosis or by both combined. There are no set rules governing the definition of taxa, but the naming and publication of new taxa is governed by sets of rules.

superfamily to subspecies), is regulated by the International Code of Zoological Nomenclature (ICZN Code).[89] In the fields of phycology, mycology, and botany, the naming of taxa is governed by the International Code of Nomenclature for algae, fungi, and plants (ICN).[90]

The initial description of a taxon involves five main requirements:[91]

  1. The taxon must be given a name based on the 26 letters of the Latin alphabet (a binomial for new species, or uninomial for other ranks).
  2. The name must be unique (i.e. not a homonym).
  3. The description must be based on at least one name-bearing type specimen.
  4. It should include statements about appropriate attributes either to describe (define) the taxon or to differentiate it from other taxa (the diagnosis, ICZN Code, Article 13.1.1, ICN, Article 38, which may or may not be based on morphology[92]). Both codes deliberately separate defining the content of a taxon (its circumscription) from defining its name.
  5. These first four requirements must be published in a work that is obtainable in numerous identical copies, as a permanent scientific record.

However, often much more information is included, like the geographic range of the taxon, ecological notes, chemistry, behavior, etc. How researchers arrive at their taxa varies: depending on the available data, and resources, methods vary from simple

DNA sequence data.[93]

Author citation

Main articles: Author citation (botany) and Author citation (zoology)

An "authority" may be placed after a scientific name.

list of botanists by author abbreviation).[96] The system for assigning authorities differs slightly between botany and zoology.[8] However, it is standard that if the genus of a species has been changed since the original description, the original authority's name is placed in parentheses.[97]

Phenetics

A comparison of phylogenetic and phenetic (character-based) concepts
Main article: Phenetics

In phenetics, also known as taximetrics, or numerical taxonomy, organisms are classified based on overall similarity, regardless of their phylogeny or evolutionary relationships.

apomorphic) traits.[98] However, certain phenetic methods, such as neighbor joining, have persisted, as rapid estimators of relationships when more advanced methods (such as Bayesian inference) are too computationally expensive.[99]

Databases

Main article: Taxonomic database

Modern taxonomy uses database technologies to search and catalogue classifications and their documentation.[100] While there is no commonly used database, there are comprehensive databases such as the Catalogue of Life, which attempts to list every documented species.[101] The catalogue listed 1.64 million species for all kingdoms as of April 2016, claiming coverage of more than three-quarters of the estimated species known to modern science.[102]

See also

Notes

  1. ^ This ranking system, except for "Strain", can be remembered by the mnemonic "Do Kings Play Chess On Fine Glass Sets?"

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Bibliography
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