In biology, a species (abbreviated sp., with the plural form species abbreviated spp.

) is the basic
unit of biological classification and a taxonomic rank. A species is often defined as the largest group
of organisms in which two individuals can produce fertile offspring, typically by sexual reproduction.
While this definition is often adequate, looked at more closely it is problematic. For example,
with hybridisation, in a species complex of hundreds of similar microspecies, or in a ring species, the
boundaries between closely related species become unclear. Other ways of defining species include
similarity of DNA, morphology, or ecological niche.
All species are given a two-part name, a "binomial". The first part of a binomial is the genus to which
the species belongs. The second part is called the specific name or the specific epithet (in botanical
nomenclature, also sometimes in zoological nomenclature). For example, Boa constrictor is one of
four species of the Boa genus.
Species were seen from the time of Aristotle until the 18th century as fixed kinds that could be
arranged in a hierarchy, the great chain of being. In the 19th century, biologists grasped that species
could evolve given sufficient time. Charles Darwin's 1859 book The Origin of Species explained
how species could arise by natural selection. Genes can sometimes be exchanged between species
by horizontal gene transfer; and species may become extinct for a variety of reasons.

Contents
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1History
o 1.1Classical forms
o 1.2Fixed species
o 1.3Species that could change
2Naming
o 2.1Common and scientific names
o 2.2Species description
o 2.3Abbreviations
o 2.4Identification codes
o 2.5Lumping and splitting
3Mayr's biological species concept
4The species problem
o 4.1When Mayr's concept breaks down
o 4.2Aggregates of microspecies
o 4.3Hybridisation
o 4.4Ring species
5Attempts at definition
o 5.1Typological or morphological species
o 5.2Recognition and cohesion species
o 5.3Vavilovian species
o 5.4Genetic similarity
o 5.5Evolutionary species
o 5.6Phylogenetic or cladistic species
o 5.7Ecological species
o 5.8Genetic species
o 5.9Evolutionarily significant unit
o 5.10Phenetic species
6Change
o 6.1Speciation
o 6.2Exchange of genes between species
 o 6.3Extinction
7See also
8Notes
9References
10External links

History[edit]
Classical forms[edit]
Main article: Aristotle's biology
In his biology, Aristotle used the term (gnos) to mean a kind, such as a bird or fish, and
(eidos) to mean a specific form within a kind, such as (within the birds) the crane, eagle, crow,
or sparrow. These terms were translated into Latin as "genus" and "species", though they do not
correspond to the Linnean terms thus named; today the birds are a class, the cranes are a family,
and the crows a genus. A kind was distinguished by its attributes; for instance, a bird has feathers, a
beak, wings, a hard-shelled egg, and warm blood. A form was distinguished by being shared by all
its members, the young inheriting any variations they might have from their parents. Aristotle
believed all kinds and forms to be distinct and unchanging. His approach remained influential until
the Renaissance.[1]

Fixed species[edit]

John Ray believed that species breed true, and do not change.

Main article: Great chain of being

When observers in the Early Modern period began to develop systems of organization for living
things, they placed each kind of animal or plant into a context. Many of these early delineation
schemes would now be considered whimsical: schemes included consanguinity based on colour (all
plants with yellow flowers) or behaviour (snakes, scorpions and certain biting ants). John
Ray (1686), an English naturalist, was the first to give a biological definition of the term "species", as
follows:
No surer criterion for determining species has occurred to me than the distinguishing features that
perpetuate themselves in propagation from seed. Thus, no matter what variations occur in the
individuals or the species, if they spring from the seed of one and the same plant, they are
accidental variations and not such as to distinguish a species ... Animals likewise that differ
specifically preserve their distinct species permanently; one species never springs from the seed of
another nor vice versa.[2]
Carl Linnaeus created the binomial system for naming species.

In the 18th century, the Swedish scientist Carl Linnaeus classified organisms according to shared
physical characteristics, and not simply based upon differences.[3] He established the idea of
a taxonomic hierarchy of classification based upon observable characteristics and intended to reflect
natural relationships.[4][5] At the time, however, it was still widely believed that there was no organic
connection between species, no matter how similar they appeared. This view was influenced by
European scholarly and religious education, which held that the categories of life are dictated by
God, forming an Aristotelian hierarchy, the scala naturae or great chain of being. However, whether
or not it was supposed to be fixed, the scala (a ladder) inherently implied the possibility of climbing.[6]

Species that could change[edit]

By the 19th century, naturalists understood that species could change form over time, and that the
history of the planet provided enough time for major changes. Jean-Baptiste Lamarck, in his
1809 Zoological Philosophy, described the transmutation of species, proposing that a species could
change over time, in a radical departure from Aristotelian thinking.[7]
In 1859, Charles Darwin and Alfred Russel Wallace provided a compelling account of evolution and
the formation of new species. Darwin argued that it was populations that evolved, not individuals,
by natural selection from naturally occurring variation among individuals.[8] This required a new
definition of species. Darwin concluded that species are what they appear to be: ideas, provisionally
useful for naming groups of interacting individuals. "I look at the term species", he wrote, "as one
arbitrarily given for the sake of convenience to a set of individuals closely resembling each other ... It
does not essentially differ from the word variety, which is given to less distinct and more fluctuating
forms. The term variety, again, in comparison with mere individual differences, is also applied
arbitrarily, and for convenience sake."[9]

A cougar, mountain lion, panther, or puma, among other common names: its scientific name is Puma concolor.

Naming[edit]
Common and scientific names[edit]
The commonly used names for kinds of organisms are often ambiguous: "cat" could mean the
domestic cat, Felis catus, or the cat family, Felidae. Another problem with common names is that
they often vary from place to place, so that puma, cougar, catamount, panther, painter and mountain
lion all mean Puma concolor in various parts of America, while "panther" may also mean
the jaguar (Panthera onca) of Latin America or the leopard (Panthera pardus) of Africa and Asia. In
contrast, the scientific names of species are chosen to be unique and universal; they are in two parts
used together: the genus as in Puma, and the specific epithet as in concolor.[10][11]

Species description[edit]

The type specimen(holotype) of Lacerta plica, described by Linnaeus in 1758

Main article: Species description

A species is given a name when a type specimen is described formally by a scientist, in a paper that
assigns it a scientific name. The name becomes a validly published name(in botany) or an available
name (in zoology) when the paper is accepted for publication. The type material is provided for other
scientists to examine, often in the research collection of a major museum.[12][13][14] Scientists are asked
to choose names that, in the words of the International Code of Zoological Nomenclature, are
"appropriate, compact, euphonious, memorable, and do not cause offence."[15]

Abbreviations[edit]
Books and articles sometimes intentionally do not identify species fully and use the abbreviation
"sp." in the singular or "spp." (standing for species pluralis, the Latin for multiple species) in the plural
in place of the specific name or epithet (e.g. Canis sp.) This commonly occurs when authors are
confident that some individuals belong to a particular genus but are not sure to which exact species
they belong, as is common in paleontology. Authors may also use "spp." as a short way of saying
that something applies to many species within a genus, but not to all. If scientists mean that
something applies to all species within a genus, they use the genus name without the specific name
or epithet. The names of genera and species are usually printed in italics. Abbreviations such as
"sp." should not be italicized.[16]

Identification codes[edit]
Various codes have been devised to provide identifiers for species, including:

National Center for Biotechnology Information (NCBI) employs a numeric 'taxid' or Taxonomy
identifier, a "stable unique identifier", e.g., the taxid of H. sapiens is 9606.[17]
Kyoto Encyclopedia of Genes and Genomes (KEGG) employs a three- or four-letter code for a
limited number of organisms; in this code, for example, H. sapiens is simply hsa.[18]
UniProt employs an "organism mnemonic" of not more than five alphanumeric characters,
e.g., HUMAN for H. sapiens.[19]
Integrated Taxonomic Information System (ITIS) provides a unique number for each species.
The LSID for Homo sapiens is urn:lsid:catalogueoflife.org:taxon:4da6736d-d35f-11e6-9d3f-
bc764e092680:col20170225.[20]
Lumping and splitting[edit]
Main article: Lumpers and splitters
The naming of a particular species, including which genus (and higher taxa) it is placed in, is
a hypothesis about the evolutionary relationships and distinguishability of that group of organisms.
As further information comes to hand, the hypothesis may be confirmed or refuted. Sometimes,
especially in the past when communication was more difficult, taxonomists working in isolation have
given two distinct names to individual organisms later identified as the same species. When two
named species are discovered to be of the same species, the older species name is usually
retained, and the newer species name dropped, a process called synonymization, or colloquially,
as lumping. Dividing a taxon into multiple, often new, taxa is called splitting. Taxonomists are often
referred to as "lumpers" or "splitters" by their colleagues, depending on their personal approach to
recognizing differences or commonalities between organisms.[21][22]

Mayr's biological species concept[edit]

Ernst Mayr proposed the widely used Biological Species Concept.

Main article: Biological Species Concept

Most modern textbooks use Ernst Mayr's definition, known as the Biological Species Concept. It is
also called a reproductive or isolation concept. This defines a species as[23]
groups of actually or potentially interbreeding natural populations, which are reproductively isolated
from other such groups".[23]
It can be argued that this definition is a natural consequence of the effect of sexual reproduction on
the dynamics of natural selection.[24][25][26][27] Mayr's definition excludes unusual or artificial matings that
result from deliberate human action, or occur only in captivity, or that involve animals capable of
mating but that do not normally do so in the wild.[23]

The species problem[edit]

Main article: Species Problem