Species

Species
A species (pl.: species) is a population of organisms in which any two

individuals of the appropriate sexes or mating types can produce fertile
offspring, typically by sexual reproduction.[1] It is the basic unit of
classification and a taxonomic rank of an organism, as well as a unit of
biodiversity. Other ways of defining species include their karyotype, DNA
sequence, morphology, behaviour, or ecological niche. In addition,
palaeontologists use the concept of the chronospecies since fossil
reproduction cannot be examined. The most recent rigorous estimate for the
total number of species of eukaryotes is between 8 and 8.7 million.[2][3][4]
About 14% of these had been described by 2011.[4] All species (except
viruses) are given a two-part name, called 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 the species of the genus Boa, with constrictor being the species' epithet.
While the definitions given above may seem adequate at first glance, when
looked at more closely they represent problematic species concepts. For
example, the boundaries between closely related species become unclear with The hierarchy of
hybridisation, in a species complex of hundreds of similar microspecies, and biological classification's
in a ring species. Also, among organisms that reproduce only asexually, the eight major taxonomic
concept of a reproductive species breaks down, and each clone is potentially ranks. A genus contains
a microspecies. Although none of these are entirely satisfactory definitions, one or more species.
and while the concept of species may not be a perfect model of life, it is still Minor intermediate
ranks are not shown.
a useful tool to scientists and conservationists for studying life on Earth,
regardless of the theoretical difficulties. If species were fixed and distinct
from one another, there would be no problem, but evolutionary processes cause species to change. This
obliges taxonomists to decide, for example, when enough change has occurred to declare that a lineage
should be divided into multiple chronospecies, or when populations have diverged to have enough
distinct character states to be described as cladistic species.
Species and higher taxa were seen from the time of Aristotle until the 18th century as categories 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 On the Origin of Species
explained how species could arise by natural selection. That understanding was greatly extended in the
20th century through genetics and population ecology. Genetic variability arises from mutations and
recombination, while organisms themselves are mobile, leading to geographical isolation and genetic drift
with varying selection pressures. Genes can sometimes be exchanged between species by horizontal gene
transfer; new species can arise rapidly through hybridisation and polyploidy; and species may become
extinct for a variety of reasons. Viruses are a special case, driven by a balance of mutation and selection,
and can be treated as quasispecies.
Definition
Biologists and taxonomists have made many attempts to define species, beginning from morphology and
moving towards genetics. Early taxonomists such as Linnaeus had no option but to describe what they
saw: this was later formalised as the typological or morphological species concept. Ernst Mayr
emphasised reproductive isolation, but this, like other species concepts, is hard or even impossible to
test.[5][6] Later biologists have tried to refine Mayr's definition with the recognition and cohesion
concepts, among others.[7] Many of the concepts are quite similar or overlap, so they are not easy to
count: the biologist R. L. Mayden recorded about 24 concepts,[8] and the philosopher of science John
Wilkins counted 26.[5] Wilkins further grouped the species concepts into seven basic kinds of concepts:
(1) agamospecies for asexual organisms (2) biospecies for reproductively isolated sexual organisms (3)
ecospecies based on ecological niches (4) evolutionary species based on lineage (5) genetic species based
on gene pool (6) morphospecies based on form or phenotype and (7) taxonomic species, a species as
determined by a taxonomist.[9]
Typological or morphological species

A typological species is a group of organisms in which individuals
conform to certain fixed properties (a type, which may be defined
by a chosen 'nominal species'), so that even pre-literate people
often recognise the same taxon as do modern taxonomists.[11][12]
The clusters of variations or phenotypes within specimens (such as
longer or shorter tails) would differentiate the species. This
method was used as a "classical" method of determining species,
such as with Linnaeus, early in evolutionary theory. However,
different phenotypes are not necessarily different species (e.g. a All adult Eurasian blue tits share the
four-winged Drosophila born to a two-winged mother is not a same coloration, unmistakably
different species). Species named in this manner are called identifying the morphospecies.[10]
morphospecies.[13][14]
In the 1970s, Robert R. Sokal, Theodore J. Crovello and Peter Sneath proposed a variation on the
morphological species concept, a phenetic species, defined as a set of organisms with a similar phenotype
to each other, but a different phenotype from other sets of organisms.[15] It differs from the morphological
species concept in including a numerical measure of distance or similarity to cluster entities based on
multivariate comparisons of a reasonably large number of phenotypic traits.[16]
Recognition and cohesion species

A mate-recognition species is a group of sexually reproducing organisms that recognise one another as
potential mates.[17][18] Expanding on this to allow for post-mating isolation, a cohesion species is the
most inclusive population of individuals having the potential for phenotypic cohesion through intrinsic
cohesion mechanisms; no matter whether populations can hybridise successfully, they are still distinct
cohesion species if the amount of hybridisation is insufficient to completely mix their respective gene
pools.[19] A further development of the recognition concept is provided by the biosemiotic concept of
species.[20]
Genetic similarity and barcode species

In microbiology, genes can move freely even between distantly
related bacteria, possibly extending to the whole bacterial domain.
As a rule of thumb, microbiologists have assumed that members
of Bacteria or Archaea with 16S ribosomal RNA gene sequences
more similar than 97% to each other need to be checked by DNA–
DNA hybridisation to decide if they belong to the same
species.[21] This concept was narrowed in 2006 to a similarity of
98.7%.[22]
A region of the gene for the
cytochrome c oxidase enzyme is
The average nucleotide identity (ANI) method quantifies genetic
used to distinguish species in the
distance between entire genomes, using regions of about 10,000 Barcode of Life Data Systems
base pairs. With enough data from genomes of one genus, database.
algorithms can be used to categorize species, as for Pseudomonas
avellanae in 2013,[23] and for all sequenced bacteria and archaea
since 2020.[24] Observed ANI values among sequences appear to have an "ANI gap" at 85–95%,
suggesting that a genetic boundary suitable for defining a species concept is present.[25]
DNA barcoding has been proposed as a way to distinguish species suitable even for non-specialists to
use.[26] One of the barcodes is a region of mitochondrial DNA within the gene for cytochrome c oxidase.
A database, Barcode of Life Data System, contains DNA barcode sequences from over 190,000
species.[27][28] However, scientists such as Rob DeSalle have expressed concern that classical taxonomy
and DNA barcoding, which they consider a misnomer, need to be reconciled, as they delimit species
differently.[29] Genetic introgression mediated by endosymbionts and other vectors can further make
barcodes ineffective in the identification of species.[30]
Phylogenetic or cladistic species

A phylogenetic or cladistic species is "the smallest aggregation of populations (sexual) or lineages
(asexual) diagnosable by a unique combination of character states in comparable individuals
(semaphoronts)".[31] The empirical basis – observed character states – provides the evidence to support
hypotheses about evolutionarily divergent lineages that have maintained their hereditary integrity through
time and space.[32][33][34][35] Molecular markers may be used to determine diagnostic genetic differences
in the nuclear or mitochondrial DNA of various species.[36][31][37] For example, in a study done on fungi,
studying the nucleotide characters using cladistic species produced the most accurate results in
recognising the numerous fungi species of all the concepts studied.[37][38] Versions of the phylogenetic
species concept that emphasise monophyly or diagnosability[39] may lead to splitting of existing species,
for example in Bovidae, by recognising old subspecies as species, despite
the fact that there are no reproductive barriers, and populations may
intergrade morphologically.[40] Others have called this approach
taxonomic inflation, diluting the species concept and making taxonomy
unstable.[41] Yet others defend this approach, considering "taxonomic
inflation" pejorative and labelling the opposing view as "taxonomic
conservatism"; claiming it is politically expedient to split species and
recognise smaller populations at the species level, because this means they
can more easily be included as endangered in the IUCN red list and can
attract conservation legislation and funding.[42]
Unlike the biological species concept, a cladistic species does not rely on
reproductive isolation – its criteria are independent of processes that are The cladistic or
integral in other concepts.[31] Therefore, it applies to asexual phylogenetic species
lineages.[36][37] However, it does not always provide clear cut and concept is that a species is
the smallest lineage which
intuitively satisfying boundaries between taxa, and may require multiple
is distinguished by a unique
sources of evidence, such as more than one polymorphic locus, to give set of either genetic or
plausible results.[37] morphological traits. No
claim is made about
reproductive isolation,
Evolutionary species making the concept useful
An evolutionary species, suggested by George Gaylord Simpson in 1951, also in palaeontology where
is "an entity composed of organisms which maintains its identity from only fossil evidence is
available.
other such entities through time and over space, and which has its own
independent evolutionary fate and historical tendencies".[8][43] This differs
from the biological species concept in embodying persistence over time. Wiley and Mayden stated that
they see the evolutionary species concept as "identical" to Willi Hennig's species-as-lineages concept, and
asserted that the biological species concept, "the several versions" of the phylogenetic species concept,
and the idea that species are of the same kind as higher taxa are not suitable for biodiversity studies (with
the intention of estimating the number of species accurately). They further suggested that the concept
works for both asexual and sexually-reproducing species.[44] A version of the concept is Kevin de
Queiroz's "General Lineage Concept of Species".[45]
Ecological species
An ecological species is a set of organisms adapted to a particular set of resources, called a niche, in the
environment. According to this concept, populations form the discrete phenetic clusters that we recognise
as species because the ecological and evolutionary processes controlling how resources are divided up
tend to produce those clusters.[46]
Genetic species
A genetic species as defined by Robert Baker and Robert Bradley is a set of genetically isolated
interbreeding populations. This is similar to Mayr's Biological Species Concept, but stresses genetic
rather than reproductive isolation.[47] In the 21st century, a genetic species could be established by
comparing DNA sequences. Earlier, other methods were available, such as comparing karyotypes (sets of
chromosomes) and allozymes (enzyme variants).[48]
Evolutionarily significant unit

An evolutionarily significant unit (ESU) or "wildlife species"[49] is a population of organisms considered
distinct for purposes of conservation.[50]
Chronospecies
In palaeontology, with only comparative anatomy (morphology) and
histology[51] from fossils as evidence, the concept of a chronospecies can
be applied. During anagenesis (evolution, not necessarily involving
branching), some palaeontologists seek to identify a sequence of species,
each one derived from the phyletically extinct one before through
continuous, slow and more or less uniform change. In such a time
sequence, some palaeontologists assess how much change is required for a
morphologically distinct form to be considered a different species from its
ancestors.[52][53][54][55]
Viral quasispecies
Viruses have enormous populations, are doubtfully living since they
A chronospecies is defined
consist of little more than a string of DNA or RNA in a protein coat, and
in a single lineage (solid
mutate rapidly. All of these factors make conventional species concepts line) whose morphology
largely inapplicable.[56] A viral quasispecies is a group of genotypes changes with time. At some
related by similar mutations, competing within a highly mutagenic point, palaeontologists
environment, and hence governed by a mutation–selection balance. It is judge that enough change
predicted that a viral quasispecies at a low but evolutionarily neutral and has occurred that two
species (A and B),
highly connected (that is, flat) region in the fitness landscape will
separated in time and
outcompete a quasispecies located at a higher but narrower fitness peak in
anatomy, once existed.
which the surrounding mutants are unfit, "the quasispecies effect" or the
"survival of the flattest". There is no suggestion that a viral quasispecies
resembles a traditional biological species.[57][58][59] The International Committee on Taxonomy of
Viruses has since 1962 developed a universal taxonomic scheme for viruses; this has stabilised viral
taxonomy.[60][61][62]
Mayr's biological species concept

Most modern textbooks make use of Ernst Mayr's 1942 definition,[63][64] known as the biological species
concept, as a basis for further discussion on the definition of species. It is also called a reproductive or
isolation concept. This defines a species as[65]
groups of actually or potentially interbreeding natural
populations, which are reproductively isolated from
other such groups.[65]
It has been argued that this definition is a natural consequence of

the effect of sexual reproduction on the dynamics of natural
selection.[66][67][68][69] Mayr's use of the adjective "potentially"
has been a point of debate; some interpretations exclude unusual Ernst Mayr proposed the widely
or artificial matings that occur only in captivity, or that involve used Biological Species Concept of
animals capable of mating but that do not normally do so in the reproductive isolation in 1942.
wild.[65]
The species problem

It is difficult to define a species in a way that applies to all organisms.[70] The debate about species
concepts is called the species problem.[65][71][72][73] The problem was recognised even in 1859, when
Darwin wrote in On the Origin of Species:
I was much struck how entirely vague and arbitrary is the distinction between species and
varieties.[74]
He went on to write:
No one definition has satisfied all naturalists; yet every naturalist knows vaguely what he
means when he speaks of a species. Generally the term includes the unknown element of a
distinct act of creation.[75]
When Mayr's concept breaks down

Many authors have argued that a simple textbook definition, following Mayr's concept, works well for
most multi-celled organisms, but breaks down in several situations:
When organisms reproduce asexually, as in single-celled organisms such as bacteria and

other prokaryotes,[76] and parthenogenetic or apomictic multi-celled organisms. DNA
barcoding and phylogenetics are commonly used in these cases.[77][78][79] The term
quasispecies is sometimes used for rapidly mutating entities like viruses.[80][81]
When scientists do not know whether two morphologically similar groups of organisms are
capable of interbreeding; this is the case with all extinct life-forms in palaeontology, as
breeding experiments are not possible.[82]
When hybridisation permits substantial gene flow between species.[83]
In ring species, when members of adjacent populations in a widely continuous distribution
range interbreed successfully but members of more distant populations do not.[84]
Species identification is made difficult by discordance between
molecular and morphological investigations; these can be
categorised as two types: (i) one morphology, multiple lineages
(e.g. morphological convergence, cryptic species) and (ii) one
lineage, multiple morphologies (e.g. phenotypic plasticity,
multiple life-cycle stages).[85] In addition, horizontal gene transfer
(HGT) makes it difficult to define a species.[86] All species
definitions assume that an organism acquires its genes from one or
two parents very like the "daughter" organism, but that is not what
happens in HGT.[87] There is strong evidence of HGT between
very dissimilar groups of prokaryotes, and at least occasionally
between dissimilar groups of eukaryotes,[86] including some
Palaeontologists are limited to
crustaceans and echinoderms.[88]
morphological evidence when
deciding whether fossil life-forms
The evolutionary biologist James Mallet concludes that
like these Inoceramus bivalves
formed a separate species.
there is
no easy
way to
tell
whether
related
geographic
or
temporal
forms The willow warbler and chiffchaff are almost identical in appearance but do not interbreed.
belong
to the
same
or
different
species.
Species
gaps
can be
verified
only
locally
and at
a point
of
time.
One is
forced
to
admit
that
Darwin's insi
geographic ra
The botanist Brent Mishler[89] argued that the species concept is not valid, notably because gene flux
decreases gradually rather than in discrete steps, which hampers objective delimitation of species.[90]
Indeed, complex and unstable patterns of gene flux have been observed in cichlid teleosts of the East
African Great Lakes.[91] Wilkins argued that "if we were being true to evolution and the consequent
phylogenetic approach to taxa, we should replace it with a 'smallest clade' idea" (a phylogenetic species
concept).[92] Mishler and Wilkins [93] and others [94] concur with this approach, even though this would
raise difficulties in biological nomenclature. Wilkins cited the ichthyologist Charles Tate Regan's early
20th century remark that "a species is whatever a suitably qualified biologist chooses to call a
species".[92] Wilkins noted that the philosopher Philip Kitcher called this the "cynical species
concept",[95] and arguing that far from being cynical, it usefully leads to an empirical taxonomy for any
given group, based on taxonomists' experience.[92] Other biologists have gone further and argued that we
should abandon species entirely, and refer to the "Least Inclusive Taxonomic Units" (LITUs),[96] a view
that would be coherent with current evolutionary theory.[94]
Aggregates of microspecies
The species concept is further weakened by the existence of microspecies, groups of organisms, including
many plants, with very little genetic variability, usually forming species aggregates.[97] For example, the
dandelion Taraxacum officinale and the blackberry Rubus fruticosus are aggregates with many
microspecies—perhaps 400 in the case of the blackberry and over 200 in the dandelion,[98] complicated
by hybridisation, apomixis and polyploidy, making gene flow between populations difficult to determine,
and their taxonomy debatable.[99][100][101] Species complexes occur in insects such as Heliconius
butterflies,[102] vertebrates such as Hypsiboas treefrogs,[103] and fungi such as the fly agaric.[104]
Blackberries belong to The butterfly genus The Hypsiboas

any of hundreds of Heliconius contains many calcaratus–fasciatus
microspecies of the similar species. species complex contains
Rubus fruticosus species at least six species of
aggregate. treefrog.
Hybridisation
Natural hybridisation presents a challenge to the concept of a reproductively isolated species, as fertile
hybrids permit gene flow between two populations. For example, the carrion crow Corvus corone and the
hooded crow Corvus cornix appear and are classified as separate species, yet they can hybridise where
their geographical ranges overlap.[105]
Hybridisation of carrion and hooded crows permits gene flow between 'species'
Carrion crow Hooded crow
Hybrid with dark belly
Ring species
A ring species is a connected series of neighbouring populations, each of which can sexually interbreed
with adjacent related populations, but for which there exist at least two "end" populations in the series,
which are too distantly related to interbreed, though there is a potential gene flow between each "linked"
population.[106] Such non-breeding, though genetically connected, "end" populations may co-exist in the
same region thus closing the ring. Ring species thus present a difficulty for any species concept that relies
on reproductive isolation.[107] However, ring species are at best rare. Proposed examples include the
herring gull–lesser black-backed gull complex around the North pole, the Ensatina eschscholtzii group of
19 populations of salamanders in America,[108] and the greenish warbler in Asia,[109] but many so-called
ring species have turned out to be the result of misclassification leading to questions on whether there
really are any ring species.[110][111][112][113]
A greenish warbler,
Opposite ends of the ring: Phylloscopus trochiloides
a herring gull (Larus
Seven "species" of Larus argentatus) (front) and a
gulls interbreed in a ring lesser black-backed gull
around the Arctic. (Larus fuscus) in Norway
Presumed evolution of
five "species" of greenish
warblers around the
Himalayas
Taxonomy and naming
Common and scientific names

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 A cougar, mountain lion, panther, or
mean the jaguar (Panthera onca) of Latin America or the leopard puma, among other common
names: its scientific name is Puma
(Panthera pardus) of Africa and Asia. In contrast, the scientific
concolor.
names of species are chosen to be unique and universal (except for
some inter-code homonyms); they are in two parts used together:
the genus as in Puma, and the specific epithet as in concolor.[114][115]
Species description
A species is given a taxonomic name when a type specimen is described formally, in a publication that
assigns it a unique scientific name. The description typically provides means for identifying the new
species, which may not be based solely on morphology[116] (see cryptic species), differentiating it from
other previously described and related or confusable species and provides
a validly published name (in botany) or an available name (in zoology)
when the paper is accepted for publication. The type material is usually
held in a permanent repository, often the research collection of a major
museum or university, that allows independent verification and the means
to compare specimens.[117][118][119] Describers of new species 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".[120]
Abbreviations
Books and articles sometimes intentionally do not identify species fully,
using the abbreviation "sp." in the singular or "spp." (standing for species
pluralis, 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 The type specimen
confident that some individuals belong to a particular genus but are not (holotype) of Lacerta plica,
sure to which exact species they belong, as is common in described by Linnaeus in
1758
paleontology.[121]
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. However, abbreviations such as "sp." should not be
italicised.[121]
When a species' identity is not clear, a specialist may use "cf." before the epithet to indicate that
confirmation is required. The abbreviations "nr." (near) or "aff." (affine) may be used when the identity is
unclear but when the species appears to be similar to the species mentioned after.[121]
Identification codes
With the rise of online databases, codes have been devised to provide identifiers for species that are
already defined, including:
National Center for Biotechnology Information (NCBI) employs a numeric 'taxid' or

Taxonomy identifier, a "stable unique identifier", e.g., the taxid of Homo sapiens is 9606.[122]
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.[123]
UniProt employs an "organism mnemonic" of not more than five alphanumeric characters,
e.g., HUMAN for H. sapiens.[124]
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.[125]
Lumping and splitting

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 corroborated 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 species names are
discovered to apply to the same species, the older species name is given priority and usually retained, and
the newer name considered as a junior synonym, a process called synonymy. 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 recognising differences or commonalities
between organisms.[126][127][121] The circumscription of taxa, considered a taxonomic decision at the
discretion of cognizant specialists, is not governed by the Codes of Zoological or Botanical
Nomenclature, in contrast to the PhyloCode, and contrary to what is done in several other fields, in which
the definitions of technical terms, like geochronological units and geopolitical entities, are explicitly
delimited.[128][94]
Broad and narrow senses

The nomenclatural codes that guide the naming of species, including the ICZN for animals and the ICN
for plants, do not make rules for defining the boundaries of the species. Research can change the
boundaries, also known as circumscription, based on new evidence. Species may then need to be
distinguished by the boundary definitions used, and in such cases the names may be qualified with sensu
stricto ("in the narrow sense") to denote usage in the exact meaning given by an author such as the person
who named the species, while the antonym sensu lato ("in the broad sense") denotes a wider usage, for
instance including other subspecies. Other abbreviations such as "auct." ("author"), and qualifiers such as
"non" ("not") may be used to further clarify the sense in which the specified authors delineated or
described the species.[121][129][130]
Change
Species are subject to change, whether by evolving into new species,[131] exchanging genes with other
species,[132] merging with other species or by becoming extinct.[133]
Speciation
The evolutionary process by which biological populations of sexually-reproducing organisms evolve to
become distinct or reproductively isolated as species is called speciation.[134][135] Charles Darwin was
the first to describe the role of natural selection in speciation in his 1859 book The Origin of Species.[136]
Speciation depends on a measure of reproductive isolation, a reduced gene flow. This occurs most easily
in allopatric speciation, where populations are separated geographically and can diverge gradually as
mutations accumulate. Reproductive isolation is threatened by hybridisation, but this can be selected
against once a pair of populations have incompatible alleles of the same gene, as described in the
Bateson–Dobzhansky–Muller model.[131] A different mechanism, phyletic speciation, involves one
lineage gradually changing over time into a new and distinct form (a chronospecies), without increasing
the number of resultant species.[137]
Exchange of genes between species

Horizontal gene transfer between organisms of different species,
either through hybridisation, antigenic shift, or reassortment, is
sometimes an important source of genetic variation. Viruses can
transfer genes between species. Bacteria can exchange plasmids
with bacteria of other species, including some apparently distantly
related ones in different phylogenetic domains, making analysis of
their relationships difficult, and weakening the concept of a
bacterial species.[138][86][139][132]
Louis-Marie Bobay and Howard Ochman suggest, based on

analysis of the genomes of many types of bacteria, that they can
often be grouped "into communities that regularly swap genes", in
much the same way that plants and animals can be grouped into
reproductively isolated breeding populations. Bacteria may thus
form species, analogous to Mayr's biological species concept, Horizontal gene transfers between
consisting of asexually reproducing populations that exchange widely separated species
genes by homologous recombination.[140][141] complicate the phylogeny of
bacteria.
Extinction
A species is extinct when the last individual of that species dies, but it may be functionally extinct well
before that moment. It is estimated that over 99 percent of all species that ever lived on Earth, some five
billion species, are now extinct. Some of these were in mass extinctions such as those at the ends of the
Ordovician, Devonian, Permian, Triassic and Cretaceous periods. Mass extinctions had a variety of
causes including volcanic activity, climate change, and changes in oceanic and atmospheric chemistry,
and they in turn had major effects on Earth's ecology, atmosphere, land surface and waters.[142][143]
Another form of extinction is through the assimilation of one species by another through hybridization.
The resulting single species has been termed as a "compilospecies".[144]
Practical implications
Biologists and conservationists need to categorise and identify organisms in the course of their work.
Difficulty assigning organisms reliably to a species constitutes a threat to the validity of research results,
for example making measurements of how abundant a species is in an ecosystem moot. Surveys using a
phylogenetic species concept reported 48% more species and accordingly smaller populations and ranges
than those using nonphylogenetic concepts; this was termed "taxonomic inflation",[145] which could
cause a false appearance of change to the number of endangered species and consequent political and
practical difficulties.[146][147] Some observers claim that there is an inherent conflict between the desire
to understand the processes of speciation and the need to identify and to categorise.[147]
Conservation laws in many countries make special provisions to prevent species from going extinct.
Hybridization zones between two species, one that is protected and one that is not, have sometimes led to
conflicts between lawmakers, land owners and conservationists. One of the classic cases in North
America is that of the protected northern spotted owl which hybridises with the unprotected California
spotted owl and the barred owl; this has led to legal debates.[148]
It has been argued, that since species are not comparable, simply counting them is not a valid measure of
biodiversity; alternative measures of phylogenetic biodiversity have been proposed.[149][90][150]
History
Classical forms
In his biology, Aristotle used the term γένος (génos) to mean a kind, such as a bird or fish, and εἶδος
(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. More importantly, in Aristotle's works, the terms γένος (génos) and εἶδος (eidos)
are relative; a taxon that is considered an eidos in a given context can be considered a génos in another,
and be further subdivided into eide (plural of eidos).[151][152] His approach remained influential until the
Renaissance,[153] and still, to a lower extent, today.[154]
Fixed species
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, an English naturalist, was the first to
attempt a biological definition of species in 1686, 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 John Ray believed that
the individuals or the species, if they spring from the seed of one species breed true and do
and the same plant, they are accidental variations and not such not change, even though
variations exist.
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.[155]
In the 18th century, the Swedish scientist Carl Linnaeus classified
organisms according to shared physical characteristics, and not simply
based upon differences.[156] Like many contemporary
systematists, [157][158][159] he established the idea of a taxonomic hierarchy
of classification based upon observable characteristics and intended to
reflect natural relationships.[160][161] At the time, however, it was still
widely believed that there was no organic connection between species
(except, possibly, between those of a given genus),[94] no matter how
similar they appeared. This view was influenced by European scholarly
and religious education, which held that the taxa had been created by God,
forming an Aristotelian hierarchy, the scala naturae or great chain of Carl Linnaeus created the
being. However, whether or not it was supposed to be fixed, the scala (a binomial system for naming
ladder) inherently implied the possibility of climbing.[162] species.
Mutability
In viewing evidence of hybridisation, Linnaeus recognised that species were not fixed and could change;
he did not consider that new species could emerge and maintained a view of divinely fixed species that
may alter through processes of hybridisation or acclimatisation.[163] 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.[164]
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.[165] 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, writing:
I look at the term species 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.[166]
See also
Cline Lists of plant species
Encyclopedia of Life Outline of zoology
Endangered species Systematics
Global biodiversity Speciation
Lists of animal species Pseudospeciation
Gallery
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ne-documents-archive.info/entries/species/) at the Stanford Encyclopedia of Philosophy
VisualTaxa (http://visualtaxa.redgolpe.com/)
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Species

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Species

A species (pl.: species) is a population of organisms in which any two

Typological or morphological species

Recognition and cohesion species

Genetic similarity and barcode species

Phylogenetic or cladistic species

Evolutionarily significant unit

Mayr's biological species concept

It has been argued that this definition is a natural consequence of

The species problem

When Mayr's concept breaks down

When organisms reproduce asexually, as in single-celled organisms such as bacteria and

Blackberries belong to The butterfly genus The Hypsiboas

Carrion crow Hooded crow

Hybrid with dark belly

Taxonomy and naming

Common and scientific names

National Center for Biotechnology Information (NCBI) employs a numeric 'taxid' or

Lumping and splitting

Broad and narrow senses

Exchange of genes between species

Louis-Marie Bobay and Howard Ochman suggest, based on

No surer criterion for determining species has occurred to me

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