From Wikipedia, the free encyclopedia
In
biology,
taxonomy (from
Ancient Greek τάξις (taxis), meaning 'arrangement', and
-νομία (-nomia), meaning '
method') is the
science of naming, defining (
circumscribing) and classifying groups of biological
organisms on the basis of shared characteristics. Organisms are grouped together into
taxa (singular: taxon) and these groups are given a
taxonomic rank;
groups of a given rank can be aggregated to form a super-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 system known as
Linnaean taxonomy for categorizing organisms and
binomial nomenclature for naming organisms.
With the advent of such fields of study as
phylogenetics,
cladistics, and
systematics, the Linnaean system has progressed to a system of modern biological classification based on the
evolutionary relationships between 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. As points of reference, recent definitions of taxonomy are presented below:
- Theory and practice of grouping individuals into species,
arranging species into larger groups, and giving those groups names,
thus producing a classification.
- A field of science (and major component of systematics) that encompasses description, identification, nomenclature, and classification
- The science of classification, in biology the arrangement of organisms into a classification
- "The science of classification as applied to living organisms, including study of means of formation of species, etc."
- "The analysis of an organism's characteristics for the purpose of classification"
- "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)
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. For example, definition 6 is paired with the following definition of systematics that places nomenclature outside taxonomy:
- 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".
A whole set of terms including taxonomy,
systematic biology, systematics,
biosystematics, scientific classification, biological classification, and
phylogenetics have at times had overlapping meanings – sometimes the same, sometimes slightly different, but always related and intersecting. The broadest meaning of "taxonomy" is used here. The term itself was introduced in 1813 by
de Candolle, in his
Théorie élémentaire de la botanique.
Monograph and taxonomic revision
A
taxonomic revision or
taxonomic review is a novel analysis of the variation patterns in a particular
taxon.
This analysis may be executed on the basis of any combination of the
various available kinds of characters, such as morphological,
anatomical, 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 result in a change in the classification of these subtaxa, the
identification of new subtaxa, or the merger of previous subtaxa.
Alpha and beta taxonomy
The term "
alpha taxonomy" is primarily used today to refer to the discipline of finding, describing, and naming
taxa, particularly species.
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.
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.
… 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.
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 (pp. 365–366).
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. Thus,
Ernst Mayr in 1968 defined "
beta taxonomy" as the classification of ranks higher than species.
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
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. By extension, macrotaxonomy is the study of groups at the higher
taxonomic ranks subgenus and above.
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. 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. Later came systems based on a more complete
consideration of the characteristics of taxa, referred to as "natural
systems", such as those of
de Jussieu (1789), de Candolle (1813) and
Bentham and Hooker (1862–1863). These were pre-
evolutionary in thinking. The publication of
Charles Darwin's
On the Origin of Species (1859) led to new ways of thinking about classification based on evolutionary relationships. This was the concept of
phyletic systems, from 1883 onwards. This approach was typified by those of
Eichler (1883) and
Engler (1886–1892). The advent of
molecular genetics and statistical methodology allowed the creation of the modern era of "phylogenetic systems" based on
cladistics, rather than
morphology alone.
Pre-Linnaean
Early taxonomists
Naming
and classifying our surroundings has probably been taking place as long
as mankind has been able to communicate. It would always have been
important to know the names of poisonous and edible plants and animals
in order to communicate this information to other members of the family
or group. 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.
Ancient times
Organisms were first classified by
Aristotle (
Greece, 384–322 BC) during his stay on the
Island of Lesbos. 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. He divided all living things into two groups: plants and animals. Some of his groups of animals, such as
Anhaima (animals without blood, translated as
invertebrates) and
Enhaima (animals with blood, roughly the
vertebrates), as well as groups like the
sharks and
cetaceans, are still commonly used today. His student
Theophrastus (Greece, 370–285 BC) carried on this tradition, mentioning some 500 plants and their uses in his
Historia Plantarum. Again, several plant groups currently still recognized can be traced back to Theophrastus, such as
Cornus,
Crocus, and
Narcissus.
Medieval
Taxonomy in the
Middle Ages was largely based on the
Aristotelian system, 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,
again deriving ultimately from Aristotle. Aristotelian system did not
classify plants or fungi, due to the lack of microscope at the time, as his ideas were based on arranging the complete world in a single continuum, as per the
scala naturae (the Natural Ladder). This, as well, was taken into consideration in the Great chain of being. Advances were made by scholars such as
Procopius,
Timotheos of Gaza,
Demetrios Pepagomenos, and
Thomas Aquinas.
Medieval thinkers used abstract philosophical and logical
categorizations more suited to abstract philosophy than to pragmatic
taxonomy.
Renaissance and Early Modern
During the
Renaissance, the
Age of Reason, and the Enlightenment, categorizing organisms became more prevalent,
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". His
magnum opus De Plantis came out in 1583, and described more than 1500 plant species. Two large plant families that he first recognized are still in use today: the
Asteraceae and
Brassicaceae. Then in the 17th century
John Ray (
England, 1627–1705) wrote many important taxonomic works. Arguably his greatest accomplishment was
Methodus Plantarum Nova (1682),
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). His work from 1700,
Institutiones Rei Herbariae,
included more than 9000 species in 698 genera, which directly
influenced Linnaeus, as it was the text he used as a young student.
The Linnaean era
The Swedish botanist
Carl Linnaeus (1707–1778) ushered in a new era of taxonomy. With his major works
Systema Naturae 1st Edition in 1735,
Species Plantarum in 1753, and
Systema Naturae 10th Edition, he revolutionized modern taxonomy. His works implemented a standardized binomial naming system for animal and plant species,
which proved to be an elegant solution to a chaotic and disorganized
taxonomic literature. He not only introduced the standard of class,
order, genus, and species, but also made it possible to identify plants
and animals from his book, by using the smaller parts of the flower. Thus the
Linnaean system was born, and is still used in essentially the same way today as it was in the 18th century.
Currently, plant and animal taxonomists regard Linnaeus' work as the
"starting point" for valid names (at 1753 and 1758 respectively).
Names published before these dates are referred to as "pre-Linnaean",
and not considered valid (with the exception of spiders published in
Svenska Spindlar). Even taxonomic names published by Linnaeus himself before these dates are considered pre-Linnaean.
Modern system of classification
With Darwin's theory, a general acceptance quickly appeared that a classification should reflect the Darwinian principle of
common descent.
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. 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. 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. 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.
The
cladistic method has emerged since the 1960s. In 1958,
Julian Huxley used the term clade. Later, in 1960, Cain and Harrison introduced the term cladistic. The salient feature is arranging taxa in a hierarchical
evolutionary tree, ignoring ranks. A taxon is called monophyletic, if it includes all the descendants of an ancestral form. Groups that have descendant groups removed from them are termed
paraphyletic, while groups representing more than one branch from the tree of life are called
polyphyletic. The
International Code of Phylogenetic Nomenclature or
PhyloCode is intended to regulate the formal naming of
clades. Linnaean ranks will be optional under the
PhyloCode, which is intended to coexist with the current, rank-based codes.
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.
Well before Linnaeus, plants and animals were considered separate Kingdoms.
Linnaeus used this as the top rank, dividing the physical world into
the plant, animal and mineral kingdoms. As advances in microscopy made
classification of microorganisms possible, the number of kingdoms
increased, five- and six-kingdom systems being the most common.
Domains are a relatively new grouping. First proposed in 1977,
Carl Woese's
three-domain system was not generally accepted until later. One main characteristic of the three-domain method is the separation of
Archaea and
Bacteria, previously grouped into the single kingdom Bacteria (a kingdom also sometimes called
Monera), with the
Eukaryota for all organisms whose cells contain
a nucleus. A small number of scientists include a sixth kingdom, Archaea, but do not accept the domain method.
Thomas Cavalier-Smith, who has published extensively on the classification of
protists, has recently proposed that the
Neomura, the clade that groups together the Archaea and
Eucarya, would have evolved from Bacteria, more precisely from
Actinobacteria. His 2004 classification treated the
archaeobacteria as part of a subkingdom of the kingdom Bacteria, i.e., he rejected the three-domain system entirely. 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.
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, which covers eukaryotes only with an emphasis on protists, and Ruggiero et al., 2015, covering both eukaryotes and
prokaryotes to the rank of Order, although both exclude fossil representatives. A separate compilation (Ruggiero, 2014) covers extant taxa to the rank of family. Other, database-driven treatments include the
Encyclopedia of Life, the
Global Biodiversity Information Facility, the
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. 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.
Classifying organisms
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.
Taxonomic descriptions
The initial description of a taxon involves five main requirements:
- 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).
- The name must be unique (i.e. not a homonym).
- The description must be based on at least one name-bearing type specimen.
- 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). Both codes deliberately separate defining the content of a taxon (its circumscription) from defining its name.
- 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
quantitative or
qualitative comparisons of striking features, to elaborate computer analyses of large amounts of
DNA sequence data.
Author citation
An "authority" may be placed after a scientific name. The authority is the name of the scientist or scientists who first validly published the name. For example, in 1758 Linnaeus gave the
Asian elephant the scientific name
Elephas maximus, so the name is sometimes written as "
Elephas maximus Linnaeus, 1758". The names of authors are frequently abbreviated: the abbreviation
L., for Linnaeus, is commonly used. In botany, there is, in fact, a regulated list of standard abbreviations. The system for assigning authorities differs slightly between
botany and
zoology.
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.
Phenetics
In phenetics, also known as taximetrics, or numerical taxonomy,
organisms are classified based on overall similarity, regardless of
their phylogeny or evolutionary relationships.
It results in a measure of evolutionary "distance" between taxa.
Phenetic methods have become relatively rare in modern times, largely
superseded by cladistic analyses, as phenetic methods do not distinguish
common ancestral (or
plesiomorphic) traits from new common (or apomorphic) traits. However, certain phenetic methods, such as
neighbor joining, have found their way into cladistics, as a reasonable approximation of phylogeny when more advanced methods (such as
Bayesian inference) are too computationally expensive.
Databases
Modern taxonomy uses
database technologies to search and catalogue classifications and their documentation. While there is no commonly used database, there are comprehensive databases such as the
Catalogue of Life, which attempts to list every documented species.
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.
