Está en la página 1de 13

Species

1 History and development of the


concept

For other uses, see Species (disambiguation).


The hierarchy of biological classication's eight major
taxonomic ranks. A genus contains one or more species.
Intermediate minor rankings are not shown.
In biology, a species (abbreviated sp., with the plural
form species abbreviated spp.) is one of the basic units of
biological classication and a taxonomic rank. A species
is often dened as the largest group of organisms capable of interbreeding and producing fertile ospring.
While in many cases this denition is adequate, the diculty of dening species is known as the species problem.
Diering measures are often used, such as similarity of
DNA, morphology, or ecological niche. Presence of specic locally adapted traits may further subdivide species
into "infraspecic taxa" such as subspecies (and in botany
other taxa are used, such as varieties, subvarieties, and
formae).
Species hypothesized to have the same ancestors are
placed in one genus, based on similarities. The similarity
of species is judged based on comparison of physical attributes, and where available, their DNA sequences. All
species are given a two-part name, a binomial name, or
just binomial. The rst part of a binomial is the generic
name, the genus to which the species belongs. The second part is either called the specic name (a term used
only in zoology) or the specic epithet (the term used in
botany, which can also be used in zoology). For example,
Boa constrictor is one of four species of the Boa genus.
While the genus gets capitalized, the species name does
not. The binomial is written in italics when printed and
underlined when handwritten.

John Ray

In the earliest works of science, a species was simply


an individual organism that represented a group of similar or nearly identical organisms. No other relationships beyond that group were implied. Aristotle used
the words genus and species to mean generic and specic categories. Aristotle and other pre-Darwinian scientists took the species to be distinct and unchanging,
with an "essence", like the chemical elements. When
early observers began to develop systems of organization
for living things, they began to place formerly isolated
species into a context. Many of these early delineation
schemes would now be considered whimsical and these
included consanguinity based on color (all plants with yellow owers) or behavior (snakes, scorpions and certain
biting ants).

A usable denition of the word species and reliable


methods of identifying particular species are essential for
stating and testing biological theories and for measuring
biodiversity, though other taxonomic levels such as families may be considered in broad-scale studies.[1] Extinct
species known only from fossils are generally dicult to
assign precise taxonomic rankings, which is why higher
taxonomic levels such as families are often used for fossilbased studies.[1][2]

The total number of non-bacterial and non-archaeal


John Ray (1686), an English naturalist, was the rst to
species in the world has been estimated at 8.7
give a biological denition of the term species.[6]
[3][4]
with previous estimates ranging from two
million,
In the 18th century Swedish scientist Carl Linnaeus clasmillion to 100 million.[5]
sied organisms according to shared physical characteristics, and not simply based upon dierences.[7] He is also
established the idea of a taxonomic hierarchy of clas1

1 HISTORY AND DEVELOPMENT OF THE CONCEPT


Charles Darwin and Alfred Wallace provided what scientists now consider as the most powerful and compelling
theory of evolution. Darwin argued that it was populations that evolved, not individuals. His argument relied
on a radical shift in perspective from that of Linnaeus:
rather than dening species in ideal terms (and searching
for an ideal representative and rejecting deviations), Darwin considered variation among individuals to be natural.
He further argued that variation, far from being problematic, actually provides the explanation for the existence of
distinct species.

Carl Linnaeus believed in the xity of species.

sication based upon observable characteristics and intended to reect natural relationships.[8][9] 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 inuenced by European scholarly and religious education at the time, which
held that the categories of life are dictated by God, in a
hierarchical scheme. Although there are always dierences (although sometimes minute) between individual
organisms, Linnaeus strove to identify individual organisms that were exemplary of the species, and considered
other non-exemplary organisms to be deviant and imperfect.
By the 19th century most 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, oered one of the rst logical arguments against
creationism. The new emphasis was on determining how
a species could change over time. Lamarck suggested
that an organism could pass on an acquired trait to its
ospring (i.e. he attributed the girae's long neck to
generations of giraes stretching to reach the leaves of
higher treetops). With the acceptance of the natural selection idea of Charles Darwin in the 1860s, however,
Lamarcks view of goal-oriented evolution, also known
as a teleological process, was eclipsed. Recent interest
in inheritance of acquired characteristics centers around
epigenetic processes (e.g. methylation) that do not aect
DNA sequences, but instead alter expression in an inheritable manner. Thus, Neo-Lamarckism, as it is sometimes
termed, is not a challenge to the theory of evolution by
natural selection.

Darwins work drew on Thomas Malthus' insight that the


rate of growth of a biological population will always outpace the rate of growth of the resources in the environment, such as the food supply. As a result, Darwin argued, not all the members of a population will be able to
survive and reproduce. Those that did will, on average,
be the ones possessing variationshowever slightthat
make them slightly better adapted to the environment. If
these variable traits are heritable, then the ospring of the
survivors will also possess them. Thus, over many generations, adaptive variations will accumulate in the population, while counter-adaptive traits will tend to be eliminated.
Whether a variation is adaptive or non-adaptive depends
on the environment: dierent environments favor dierent traits. Since the environment eectively selects which
organisms live to reproduce, it is the environment (the
ght for existence) that selects the traits to be passed
on. This is the theory of evolution by natural selection.
In this model, the length of a giraes neck would be explained by positing that proto-giraes with longer necks
would have had a signicant reproductive advantage to
those with shorter necks. Over many generations, the entire population would be a species of long-necked animals.
In 1859, when Darwin published his theory of natural selection, the mechanism behind the inheritance of individual traits was unknown. Although Darwin made some
speculations on how traits are inherited (pangenesis), his
theory relies only on the fact that inheritable traits exist,
and are variable (which makes his accomplishment even
more remarkable.) Although Gregor Mendel's paper on
genetics was published in 1866, its signicance was not
recognized. It was not until 1900 that his work was rediscovered by Hugo de Vries, Carl Correns and Erich von
Tschermak, who realised that the inheritable traits in
Darwins theory are genes.
The theory of the evolution of species through natural selection has two important implications for discussions of
speciesconsequences that fundamentally challenge the
assumptions behind Linnaeus taxonomy. First, it suggests that species are not just similar, they may actually be related. Some students of Darwin argue that all
species are descended from a common ancestor. Second, it supposes that species are not homogeneous,

3
xed, permanent things; members of a species are all
dierent, and over time species change. This suggests
that species do not have any clear boundaries but are
rather momentary statistical eects of constantly changing gene-frequencies. One may still use Linnaeus taxonomy to identify individual plants and animals, but one can
no longer think of species as independent and immutable.
The rise of a new species from a parental line is called
speciation. There is no clear line demarcating the ancestral species from the descendant species.
Although the current scientic understanding of species
suggests that there is no rigorous and comprehensive way
to distinguish between dierent species in all cases, biologists continue to seek concrete ways to operationalize the idea. One of the most popular biological denitions of species is in terms of reproductive isolation;
if two creatures cannot reproduce to produce fertile ospring of both sexes, then they are in dierent species.
This denition captures a number of intuitive species
boundaries, but it remains imperfect. It has nothing to
say about species that reproduce asexually, for example,
and it is very dicult to apply to extinct species. Moreover, boundaries between species are often fuzzy: there
are examples where members of one population can produce fertile ospring of both sexes with a second population, and members of the second population can produce
fertile ospring of both sexes with members of a third
population, but members of the rst and third population cannot produce fertile ospring, or can only produce
fertile ospring of the homozygous sex. Consequently,
some people reject this denition of a species.
Richard Dawkins denes two organisms as conspecic if
and only if they have the same number of chromosomes
and, for each chromosome, both organisms have the same
number of nucleotides (The Blind Watchmaker, p. 118).
However, most taxonomists would disagree. For example, in many amphibians, most notably in New Zealands
Leiopelma frogs, the genome consists of core chromosomes that are mostly invariable and accessory chromosomes, of which exist a number of possible combinations. Even though the chromosome numbers are highly
variable between populations, these can interbreed successfully and form a single evolutionary unit. In plants,
polyploidy is extremely commonplace with few restrictions on interbreeding; as individuals with an odd number
of chromosome sets are usually sterile, depending on the
actual number of chromosome sets present, this results
in the odd situation where some individuals of the same
evolutionary unit can interbreed with certain others and
some cannot, with all populations being eventually linked
as to form a common gene pool.
The classication of species has been profoundly affected by technological advances that have allowed researchers to determine relatedness based on molecular
markers, starting with the comparatively crude blood
plasma precipitation assays in the mid-20th century to

Charles Sibley's DNA-DNA hybridization studies in the


1970s leading to DNA sequencing techniques. The results of these techniques caused revolutionary changes
in the higher taxonomic categories (such as phyla and
classes), resulting in the reordering of many branches of
the phylogenetic tree (see also: molecular phylogeny).
For taxonomic categories below genera, the results have
been mixed so far; the pace of evolutionary change on the
molecular level is rather slow, yielding clear dierences
only after considerable periods of reproductive separation. DNA-DNA hybridization results have led to misleading conclusions, the Pomarine Skua Great Skua
phenomenon being a famous example.[10][11] Turtles have
been determined to evolve with just one-eighth of the
speed of other reptiles on the molecular level, and the
rate of molecular evolution in albatrosses is half of what
is found in the rather closely related storm-petrels. The
hybridization technique is now obsolete and is replaced
by more reliable computational approaches for sequence
comparison. Molecular taxonomy is not directly based
on the evolutionary processes, but rather on the overall
change brought upon by these processes. The processes
that lead to the generation and maintenance of variation
such as mutation, crossover and selection are not uniform (see also molecular clock). DNA is only extremely
rarely a direct target of natural selection rather than
changes in the DNA sequence enduring over generations
being a result of the latter; for example, silent transitiontransversion combinations would alter the melting point
of the DNA sequence, but not the sequence of the encoded proteins and thus are a possible example where, for
example in microorganisms, a mutation confers a change
in tness all by itself.

2 Biologists working denition


A usable denition of the word species and reliable
methods of identifying particular species is essential for
stating and testing biological theories and for measuring
biodiversity. Traditionally, multiple examples of a proposed species must be studied for unifying characters before it can be regarded as a species. It is generally dicult to give precise taxonomic rankings to extinct species
known only from fossils.
Some biologists may view species as statistical phenomena, as opposed to the traditional idea, with a species seen
as a class of organisms. In that case, a species is dened as a separately evolving lineage that forms a single
gene pool. Although properties such as DNA-sequences
and morphology are used to help separate closely related lineages,[12] this denition has fuzzy boundaries.[13]
However, the exact denition of the term species is still
controversial, particularly in prokaryotes,[14] and this is
called the species problem.[15] Biologists have proposed a
range of more precise denitions, but the denition used
is a pragmatic choice that depends on the particularities

of the species of concern.[15]

2.1

Common names and species

The commonly used names for plant and animal taxa


sometimes correspond to species:[16] for example, "lion",
"walrus", and "Camphor tree" each refers to a species.
In other cases common names do not: for example,
"deer" refers to a family of 34 species, including Elds
Deer, Red Deer and Elk (as the use in American English meaning Wapiti, not the use in British English meaning moose). The last two species were once considered
a single species, illustrating how species boundaries may
change with increased scientic knowledge.

2.2

Placement within genera

BIOLOGISTS WORKING DEFINITION

This binomial naming convention, later formalized in


the biological codes of nomenclature, was rst used
by Leonhart Fuchs and introduced as the standard by
Carolus Linnaeus in his 1753 Species Plantarum (followed by his 1758 Systema Naturae, 10th edition).

2.3 Abbreviated names


Books and articles sometimes intentionally do not identify species fully and use the abbreviation sp. in the singular or spp. (Species pluralis, Latin abbreviation for
multiple species) in the plural in place of the specic epithet (e.g. Canis sp.) This commonly occurs in the following situations:
The authors are condent that some individuals belong to a particular genus but are not sure to which
exact species they belong. This is particularly common in paleontology.

Ideally, a species is given a formal, scientic name, although in practice there are very many unnamed species
(which have only been described, not named). When a
species is named, it is placed within a genus. From a sci The authors use spp. as a short way of saying that
entic point of view this can be regarded as a hypothesis
something applies to many species within a genus,
that the species is more closely related to other species
but do not wish to say that it applies to all species
within its genus (if any) than to species of other genwithin that genus. If scientists mean that something
era. Species and genus are usually dened as part of a
applies to all species within a genus, they use the
larger taxonomic hierarchy. The best-known taxonomic
genus name without the specic epithet.
ranks are, in order: life, domain, kingdom, phylum, class,
order, family, genus, and species. This assignment to a
Sometimes, the aforementioned plural is rendered as
genus is not immutable; later a dierent (or the same)
sps., which may lead to confusion with ssp., this one
taxonomist may assign it to a dierent genus, in which
standing for subspecies instead. In books and articles,
case the name will also change.
genus and species names are usually printed in italics. AbIn biological nomenclature, the name for a species is a breviations such as sp., spp., sps., ssp., subsp.,
two-part name (a binomial name), treated as Latin, al- etc. should not be italicized.[17]
though roots from any language can be used as well as
names of locales or individuals. The generic name is
listed rst (with its leading letter capitalized), followed by 2.4 Identication codes
a second term. The terminology used for the second term
diers between zoological and botanical nomenclature.
Various codes have been devised for identifying particu In zoological nomenclature, the second part of the
name can be called the specic name or the specic epithet. For example, gray wolves belong to the
species Canis lupus, coyotes to Canis latrans, golden
jackals to Canis aureus, etc., and all of those belong
to the genus Canis (which also contains many other
species). For the gray wolf, the genus name is Canis,
the specic name or specic epithet is lupus, and the
binomen, the name of the species, is Canis lupus.
In botanical nomenclature, the second part of the
name can only be called the specic epithet. The
'specic name' in botany is always the combination
of genus name and specic epithet. For example,
the species commonly known as the longleaf pine is
Pinus palustris; the genus name is Pinus, the specic
epithet is palustris, the specic name is Pinus palustris.

lar species. For example:


National Center for Biotechnology Information
(NCBI) employs a numeric 'taxid' or Taxonomy
identier, a stable unique identier, e.g. the taxid
of H. sapiens is 9606.[18]
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.[19]
UniProt employs an organism mnemonic of not
more than ve alphanumeric characters, e.g. HUMAN for H. sapiens.[20]
Integrated Taxonomic Information System (ITIS)
provides a unique number for each species.

Diculty dening or identifying


species

Main article: Species problem


It is surprisingly dicult to dene the word species in a
way that applies to all naturally occurring organisms,[21]
and the debate among biologists about how to dene
species and how to identify actual species is called the
species problem. Over two dozen distinct denitions of
species are in use amongst biologists.[22]
This problem dates as early as to the writings of Charles
Darwin. While Darwin wrote the following in On the Origin of Species, Chapter II:
No one denition has satised 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.[23]
He readdressed the question in The Descent of Man,
specically discussing the question whether mankind
consists of one or several species, where he revised his
opinion, writing:
it is a hopeless endeavour to decide this
point on sound grounds, until some denition
of the term species is generally accepted; and
the denition must not include an element that
cannot possibly be ascertained, such as an act
of creation.[24]

By denition it applies only to organisms that


reproduce sexually. So it does not work for asexually
reproducing single-celled organisms and for the
relatively few parthenogenetic or apomictic multicelled organisms.[29] The term "phylotype" is often
applied to such organisms.
Biologists frequently do not know whether two morphologically similar groups of organisms are potentially capable of interbreeding.
There is considerable variation in the degree to
which hybridization may succeed under natural conditions, or even in the degree to which some organisms use sexual reproduction between individuals to
breed.
In ring species, members of adjacent populations
interbreed successfully but members of some nonadjacent populations do not.
Among microorganisms, in particular, the problem of
species identication is made dicult by discordance
between molecular and morphological investigations;
these can be categorized 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).[30] In addition, in these and other organisms,
horizontal gene transfer (HGT) makes it dicult to dene the term species. All species denitions assume that
an organism acquires its genes from one or two parents
very like the daughter organism, but HGT makes that
assumption false. There is strong evidence of HGT between very dissimilar groups of prokaryotes, and at least
occasionally between dissimilar groups of eukaryotes.
Williamson argues that there is also evidence for HGT
in some crustaceans and echinoderms.[31]

Most modern textbooks follow Ernst Mayrs denition,


known as the Biological Species Concept (BSC) of a
species as groups of actually or potentially interbreeding natural populations, which are reproductively isolated
from other such groups.[15] It has been argued that this 4 Denitions of species
denition of species is not only a useful formulation, but
is also a natural consequence of the eect of sexual repro- Prior to Darwin, naturalists viewed species as ideal or
duction on the dynamics of natural selection.[25][26][27][28] general types, which could be exemplied by an ideal
(Also see Speciation.)
specimen bearing all the traits general to the species. DarVarious parts of this denition serve to exclude some un- wins theories shifted attention from uniformity to variation and from the general to the particular. According to
usual or articial matings:
intellectual historian Louis Menand,
Those that as a result of deliberate human action,
or occur only in captivity (when the animals normal
mating partners may not be available)
Those that involve animals that may be physically
and physiologically capable of mating but, for various reasons, do not normally do so in the wild
The typical textbook denition above works well for most
multi-celled organisms, but there are several types of situations in which it breaks down:

Once our attention is redirected to the individual, we need another way of making generalizations. We are no longer interested in the
conformity of an individual to an ideal type;
we are now interested in the relation of an individual to the other individuals with which it
interacts. To generalize about groups of interacting individuals, we need to drop the language of types and essences, which is prescriptive (telling us what nches should be), and

4 DEFINITIONS OF SPECIES
adopt the language of statistics and probability,
which is predictive (telling us what the average
nch, under specied conditions, is likely to
do). Relations will be more important than categories; functions, which are variable, will be
more important than purposes; transitions will
be more important than boundaries; sequences
will be more important than hierarchies.[32]

ulations to either side, but individuals of the populations


on either side cannot interbreed. Thus, one could argue
that these populations constitute a single species, or two
distinct species. This is not a paradox; it is evidence that
species are dened by gene frequencies, and thus have
fuzzy boundaries.

Practically, biologists dene species as populations of organisms that have a high level of genetic similarity. This
may reect an adaptation to the same niche, and the transfer of genetic material from one individual to others,
through a variety of possible means. The exact level of
similarity used in such a denition is arbitrary, but this is
the most common denition used for organisms that reproduce asexually (asexual reproduction), such as some
plants and microorganisms.

BSC subspecies are considered species under the PSC;


the dierence between the BSC and the PSC can be
summed up insofar as that the BSC denes a species as a
consequence of manifest evolutionary history, while the
PSC denes a species as a consequence of manifest evolutionary potential. Thus, a PSC species is made as soon
as an evolutionary lineage has started to separate, while a
BSC species starts to exist only when the lineage separation is complete. Accordingly, there can be considerable
conict between alternative classications based upon the
PSC versus BSC, as they dier completely in their treatment of taxa that would be considered subspecies under
the latter model (e.g. the numerous subspecies of honey
bees).

Consequently, any single, universal denition of species


is necessarily arbitrary. Instead, biologists have proposed
a range of denitions; which denition a biologists uses
This shift results in a new approach to species"; Darwin
is a pragmatic choice, depending on the particularities of
concluded that species are what they appear to be: ideas,
that biologists research.
which are provisionally useful for naming groups of interacting individuals. I look at the term species, he wrote, In practice, these denitions often coincide, and the difas one arbitrarily given for the sake of convenience to a ferences between them are more a matter of emphasis
set of individuals closely resembling each other ... It does than of outright contradiction. Nevertheless, no species
not essentially dier from the word variety, which is given concept yet proposed is entirely objective, or can be apto less distinct and more uctuating forms. The term plied in all cases without resorting to judgment.
variety, again, in comparison with mere individual dif- For most vertebrates, this is the biological species conferences, is also applied arbitrarily, and for convenience cept (BSC), and to a lesser extent (or for dierent pursake.[32]
poses) the phylogenetic species concept (PSC). Many

This lack of any clear species concept in microbiology


has led to some authors arguing that the term species
is not useful when studying bacterial evolution. Instead
they see genes as moving freely between even distantly
related bacteria, with the entire bacterial domain being a
single gene pool. Nevertheless, a kind of rule of thumb
has been established, saying that species of Bacteria or
Archaea with 16S rRNA gene sequences more similar
than 97% to each other need to be checked by DNADNA Hybridization if they belong to the same species
or not.[33] This concept has been updated recently, saying
that the border of 97% was too low and can be raised to
98.7%.[34]
In the study of sexually reproducing organisms, where genetic material is shared through the process of reproduction, the ability of two organisms to interbreed and produce fertile ospring of both sexes is generally accepted
as a simple indicator that the organisms share enough
genes to be considered members of the same species.
Thus a species is a group of interbreeding organisms.
This denition can be extended to say that a species is
a group of organisms that could potentially interbreed
sh could still be classed as the same species even if they
live in dierent lakes, as long as they could still interbreed were they ever to come into contact with each other.
On the other hand, there are many examples of series of
three or more distinct populations, where individuals of
the population in the middle can interbreed with the pop-

4.1 Typological species


A group of organisms in which individuals are members of the species if they suciently conform to certain xed properties. The clusters of variations or phenotypes within specimens (i.e. longer or shorter tails)
would dierentiate the species. This method was used as
a classical method of determining species, such as with
Linnaeus early in evolutionary theory. However, we now
know that dierent phenotypes do not always constitute
dierent species (e.g. a four-winged Drosophila born to
a 2-winged mother is not a dierent species). Species
named in this manner are called morphospecies.[35][36]

4.2 Evolutionary species


A single evolutionary lineage of organisms within which
genes can be shared, and that maintains its integrity with
respect to other lineages through both time and space. At
some point in the evolution of such a group, some members may diverge from the main population and evolve

7
into a subspecies, a process that may eventually lead to
gauge whether or not the results of such experiments
the formation of a new species if isolation (geographical
are meaningful in reference to natural populations.
or ecological) is maintained. The process through which
species are formed by evolution is called speciation. A Genetic species Based on similarity of DNA of individuals or populations. Techniques to compare similarspecies that gives rise to another species is a paraphyletic
ity of DNA include DNA-DNA hybridization, and
species, or paraspecies.[37]
genetic ngerprinting (or DNA barcoding).

4.3

Phylogenetic (cladistic) species

Cohesion species Most inclusive population of individuals having the potential for phenotypic cohesion
through intrinsic cohesion mechanisms. This is an
expansion of the mate-recognition species concept
to allow for post-mating isolation mechanisms; no
matter whether populations can hybridize successfully, they are still distinct cohesion species if the
amount of hybridization is insucient to completely
mix their respective gene pools.

A phylogenetic or cladistic species is a group of organisms that shares an ancestora lineage that maintains its
hereditary integrity with respect to other lineages through
both time and space. At some point in the evolution of
such a group, members may diverge from one another:
when such a divergence becomes suciently clear, the
two populations are regarded as separate species. This
category of species denition diers from evolutionary
species in that the parent of the phylogenetic species goes Evolutionarily signicant unit (ESU) An
evolutionarily signicant unit is a population
extinct taxonomically when a new species evolves; the
of organisms that is considered distinct for purposes
mother and daughter populations now forming two new
of conservation. Often referred to as a species or
species. Subspecies as such are not recognized under this
a wildlife species, an ESU also has several possible
denition; either a population is a phylogenetic species
denitions, which coincide with denitions of
or it is not taxonomically distinguishable.
species.
It has been argued, that operation of the phylogenetic
species concept (PSC) will lead to taxonomic ination, Phenetic species Based on phenetics.
since smaller and smaller units of its population can be
distinguishedeven down to individuals. Species of Microspecies A species with very little genetic variabilbovine (i.e., cattle) for example, could be split up into
ity, usually one that reproduces by apomixis.
any number of species based on this concept.[38]
Recognition species Based on shared reproductive systems, including mating behavior. The Recognition
4.4 Other species concepts
concept of species has been introduced by Hugh E.
H. Paterson, after earlier work by Wilhelm Petersen.
Ecological species A set of organisms adapted to a particular set of resources, called a niche, in the en- Mate-recognition species A group of organisms that
vironment. According to this concept, populations
are known to recognize one another as potential
form the discrete phenetic clusters that we recognize
mates. Like the isolation species concept above,
as species because the ecological and evolutionary
it applies only to organisms that reproduce sexuprocesses controlling how resources are divided up
ally. Unlike the isolation species concept, it focuses
tend to produce those clusters.[39]
specically on pre-mating reproductive isolation.

Reproductive species Two organisms that are able to


reproduce naturally to produce fertile ospring of 5 Numbers of species
both sexes. Organisms that can reproduce but almost always make infertile hybrids of at least one Main article: Global biodiversity
sex, such as a mule, hinny or F1 male cattalo are not Bearing in mind the aforementioned problems with cateconsidered to be the same species.
gorizing species, the following numbers are only a guide.
Based on various discussions from the rst decade of the
Isolation species A set of actually or potentially inter- new millennium, counts can roughly be broken down as
breeding populations. This is generally a useful for- follows:[40]
mulation for scientists working with living examples
of the higher taxa like mammals, sh, and birds, but
more problematic for organisms that do not repro- 5.1 Number of prokaryotic species, doduce sexually. The results of breeding experiments
main Bacteria
done in articial conditions may or may not reect
what would happen if the same organisms encoun- This number is very dicult to assess, but the
tered each other in the wild, making it dicult to discussed range varies from tens of thousands to

NUMBERS OF SPECIES

5.3 Number of eukaryotic species


This number has historically varied from a few million to about 100 millions. However these higher numbers, which were based on the potential deep marine and
arthropod diversity, are now considered unlikely. The
total number of eukaryotic species is likely to be 5
3 million of which about 1.5 million have been already
named.[54] Some older estimates for various eukaryote
phyla are:
As many as 1.5 million fungi[55]
3,067 brown algae
An estimate of the number of some undiscovered and discovered
eukaryotic species.

17,000 lichens
321,212 plants, including:

billions;[41][42][43][44] most recent approaches and studies


appear to favor the larger magnitude number.[45][46][47]
Smaller numbers arise from assumptions based on a
plateauing of identication of new species (which has
technical explanations other than that fewer species remain to be identied).[41] Larger numbers address the fact
that success in culturing bacteria has only been achieved
in half of identied Bacterial phyla (where lack of success
in attempts to culture a bacterial isolate limits abilities to
study and delineate new species),[48] and address the diculty of applying traditional botanic and zoologic denitions of species to asexually reproducing bacteria (where
more modern sequencing and molecular approaches support higher species tallies).[43][49]

5.2

Number of prokaryotic species, domain Archaea

As a further microbial domain, the issues and diculties of domain Bacteria also pertain to any counting
of species of Archaea, all the more given their various extreme habitats. The classication of archaea into
species is also controversial, as they also reproduce asexually (likewise eliminating applicability of species definitions based on interbreeding),[50] and face the same
diculties associated with organism isolation and culturing (see citations for Bacteria, above).[48][49][51] Archaebacteria have been shown to exhibit high rates of
horizontal gene transfer (resulting from a bacterial cognate of sex), including between organisms quite separate based on genomic analysis.[52] As the Archaea article
notes, "[c]urrent knowledge on genetic diversity is fragmentary and the total number of archaean species cannot
be estimated with any accuracy ... though like domain
Bacteria, the number of cultured and studied phyla relative to the total is low (as of 2005, less than 50% of known
phyla cultured).[53] Taken together, very high numbers of
unique archaebacterial types are likely, as in the case of
domain Bacteria.

10,134 red and green algae


16,236 mosses
12,000 ferns and horsetails
1,021 gymnosperms
281,821 angiosperms
1,367,555 non-insect animals, including:
1,305,250 invertebrates
2,175 corals
85,000 mollusks
As many as 1.1 million arachnids, including ~1 million mites and ~100,000 other
arachnids[56]
47,000 crustaceans
68,827 other invertebrates
63,649 vertebrates

31,300 sh
7,093 amphibians[57]
9,768 reptiles[58]
9,998 birds
5,490 mammals

As many as 1030 million insects[59]


At present, organisations such as the Global Taxonomy
Initiative, the European Distributed Institute of Taxonomy and the Census of Marine Life (the last of these only
for marine organisms) are trying to improve taxonomy
and add previously undiscovered species to the taxonomy
system.[60] Current knowledge covers only a portion of
the organisms in the biosphere and thus does not enable a
complete understanding of the workings of the environment. Humankind is also currently wiping out undiscovered species at an unprecedented rate,[61] which means
that even before a new species has had the chance of being studied and classied, it may already be extinct.

Lumping and splitting of taxa

Main article: Lumpers and splitters

Koinophilia
Ring species
Speciation

The naming of a particular species may be regarded as a


hypothesis about the evolutionary relationships and distinguishability of that group of organisms. As further
information comes to hand, the hypothesis may be conrmed or refuted. Sometimes, especially in the past when
communication was more dicult, taxonomists working
in isolation have given two distinct names to individual
organisms later identied 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, taxons is called splitting. Taxonomists are often referred to as lumpers or splitters by their colleagues, depending on their personal approach to recognizing dierences or commonalities between organisms.[62][63]
Traditionally, researchers relied on observations of
anatomical dierences, and on observations of whether
dierent populations were able to interbreed successfully, to distinguish species; both anatomy and breeding
behavior are still important to assigning species status. As
a result of the revolutionary (and still ongoing) advance
in microbiological research techniques, including DNA
analysis, in the last few decades, a great deal of additional
knowledge about the dierences and similarities between
species has become available. Many populations formerly
regarded as separate species are now considered a single
taxon, and many formerly grouped populations have been
split. Any taxonomic level (species, genus, family, etc.)
can be synonymized or split, and at higher taxonomic levels, these revisions have been still more profound.
From a taxonomical point of view, groups within a
species can be dened as being of a taxon hierarchically
lower than a species. In zoology only the subspecies is
used, while in botany the variety, subvariety, and form
are used as well. In conservation biology, the concept of
evolutionary signicant units (ESU) is used, which may
dene either species or smaller distinct population segments. Identifying and naming species is the providence
of alpha taxonomy.

See also
Cline
Cryptic species complex
Encyclopedia of Life
Endangered species
Global biodiversity

Species description
Species naming
Species problem
Systematics

8 References
[1] Sahney, S.; Benton, M.J.; Ferry, P.A. (2010). Links Between Global Taxonomic Diversity, Ecological Diversity
and the Expansion of Vertebrates on Land. Biology Letters 6 (4): 544547. doi:10.1098/rsbl.2009.1024. PMC
2936204. PMID 20106856.
[2] Sahney, S. and Benton, M.J. (2008). Recovery from the
most profound mass extinction of all time (PDF). Proceedings of the Royal Society: Biological 275 (1636): 759
65. doi:10.1098/rspb.2007.1370. PMC 2596898. PMID
18198148.
[3] Mora, C. et al.; Tittensor, Derek P.; Adl, Sina; Simpson, Alastair G. B.; Worm, Boris (August 23, 2011).
Mace, Georgina M, ed. How Many Species Are There
on Earth and in the Ocean?". PLoS Biology 9 (8):
e1001127. doi:10.1371/journal.pbio.1001127. PMC
3160336. PMID 21886479.
[4] Goldenberg, Suzanne (2011-08-23). Planet Earth is
home to 8.7 million species, scientists estimate. The
Guardian (London). Archived from the original on 201309-30. Retrieved 2014-02-24.
[5] Just How Many Species Are There, Anyway?". Science
Daily. 2003-05-26. Archived from the original on 200305-28. Retrieved 2014-02-24.
[6] Historia plantarum generalis, in the volume published in
1686, Tome I, Libr. I, Chap. XX, page 40 (Quoted in
Mayr, Ernst. 1982. The growth of biological thought:
diversity, evolution, and inheritance. Cambridge, Mass.:
Belknap Press: 256)
[7] Davis, P. H.; Heywood, V. H. (1973). Principles of Angiosperm Taxonomy. Huntington, New York: Robert E.
Krieger Publishing Company. p. 17.
[8] Reveal, James L.; Pringle, James S. (1993). 7. Taxonomic Botany and Floristics. Flora of North America.
New York and Oxford: Oxford University Press. pp.
160161. ISBN 0-19-505713-9.
[9] Simpson, George Gaylord (1961). Principles of Animal
Taxonomy. New York and London: Columbia University
Press. pp. 5657.
[10] Newton, Ian (2003). Speciation and Biogeography of
Birds. Academic Press. p. 69. ISBN 9780080924991.

10

[11] Andersson, Malte (1999). Hybridization and skua phylogeny. Proceedings of the Royal Society B 266 (1428):
15791585. doi:10.1098/rspb.1999.0818.
[12] Koch, H. (2010). Combining morphology and DNA
barcoding resolves the taxonomy of Western Malagasy
Liotrigona Moure, 1961. African Invertebrates 51 (2):
413421. doi:10.5733/an.051.0210.
[13] De Queiroz K (2007). Species concepts and species
delimitation.
Syst.
Biol.
56 (6): 87986.
doi:10.1080/10635150701701083. PMID 18027281.
[14] Fraser C, Alm EJ, Polz MF, Spratt BG, Hanage WP
(February 2009). The bacterial species challenge: making sense of genetic and ecological diversity. Science 323
(5915): 7416. doi:10.1126/science.1159388. PMID
19197054.
[15] de Queiroz K (2005). Ernst Mayr and the modern
concept of species. Proc. Natl. Acad. Sci. U.S.A.
102 (Suppl 1): 66007. doi:10.1073/pnas.0502030102.
PMC 1131873. PMID 15851674.
[16] Bailey, L.H. (1933). How plants get their names. New
York: Macmillan. ISBN 9780486207964.
[17] Hardy, Jay (2011). Naming Conventions. Nomenclature
of Microorganisms, Hardydiagnostics.com.
[18] USA (2012-10-19). Home Taxonomy NCBI.
Ncbi.nlm.nih.gov. Retrieved 2012-11-25.
[19] KEGG Organisms: Complete Genomes. Genome.jp.
Retrieved 2012-11-25.
[20] Taxonomy. Uniprot.org. Retrieved 2012-11-25.
[21] Hanage, William P. (April 2013). Fuzzy species revisited. BMC Biology 11 (41). Retrieved 2013-04-22. A
coherent species concept that can be applied throughout
the kingdoms of life is still elusive.
[22] Wilkins, John (2010-10-20). How many species concepts are there?". London: The Guardian. Retrieved
2010-10-19.
[23] Darwin 1859 p. 59. Darwin-online.org.uk. Retrieved
2012-11-25.
[24] Darwin 1871 p. 24. Darwin-online.org.uk. Retrieved
2012-11-25.
[25] Hopf FA, Hopf FW. (1985). The role of the Allee eect
on species packing. Theor. Pop. Biol. 27, 27-50.
[26] Bernstein H, Byerly HC, Hopf FA, Michod RE (December 1985). Sex and the emergence of species. J.
Theor. Biol. 117 (4): 66590. doi:10.1016/S00225193(85)80246-0. PMID 4094459.

REFERENCES

[29] Templeton, A.R. (1989). The meaning of species and


speciation: A genetic perspective. In D. Otte; J.A.
Endler. Speciation and its consequences. Sunderland: Sinauer Associates Inc. pp. 327.
[30] Lahr, D. J., Laughinghouse, H. D., Oliverio, A. M., Gao,
F., & Katz, L. A. (2014). . BioEssays, 36(10), 950-959.
[31] Williamson, David I. (2003). The Origins of Larvae.
Kluwer. ISBN 1-4020-1514-3.
[32] Menand, Louis (2001). The Metaphysical Club: A Story
of Ideas in America. Farrar, Straus and Giroux. pp. 123
124. ISBN 0-374-70638-7.
[33] Stackebrandt E, Goebel BM (1994). Taxonomic note:
a place for DNA-DNA reassociation and 16S rRNA sequence analysis in the present species denition in bacteriology. Int. J. Syst. Bacteriol. 44 (4): 8469.
doi:10.1099/00207713-44-4-846.
[34] Stackebrandt E, Ebers J (2006). Taxonomic parameters
revisited: tarnished gold standards. Microbiol. Today 33:
1525.
[35] Michael Ruse (1969). Denitions of Species in Biology.
The British Journal for the Philosophy of Science 20 (2):
97119. doi:10.1093/bjps/20.2.97. JSTOR 686173.
[36] Lewin, Ralph A. (1981). Three Species Concepts.
Taxon 30 (3): 609613. doi:10.2307/1219942.
[37] Albert, James S.; Reis, Roberto E. (2011). Historical Biogeography of Neotropical Freshwater Fishes. ISBN 9780-520-26868-5.
[38] Heller, R., Frandsen, P., Lorenzen, E. D., & Siegismund,
H. R. (2013). Are there really twice as many bovid species
as we thought?. Systematic biology, 62(3), 490-493.
[39] Ridley, Mark. The Idea of Species. Evolution (2nd ed.).
Cambridge, Massachusetts: Blackwell Science. p. 719.
ISBN 0-86542-495-0.
[40] Osborn, Liz. Current Results Number of Species on
Earth. Currentresults.com. Retrieved 2012-11-25.
[41] Counting in a bacterial world. New Scientist, June 10,
2008.
[42] Torsvik V, vres L & Thingstad TF (2002).
Prokaryotic diversitymagnitude, dynamics, and
controlling factors. Science 296 (5570): 10646.
doi:10.1126/science.1071698. PMID 12004116.
[43] Pedrs-Ali, C (2006). Marine microbial diversity: Can
it be determined?". Trends in microbiology 14 (6): 257
63. doi:10.1016/j.tim.2006.04.007. PMID 16679014.

[27] Bernstein, Carol; Bernstein, Harris; (1991). Aging, sex,


and DNA repair. Boston: Academic Press. ISBN 0-12092860-4.

[44] Sogin ML, Morrison HG, Huber JA, et al. (August 2006).
Microbial diversity in the deep sea and the underexplored
rare biosphere"". Proc. Natl. Acad. Sci. U.S.A. 103
(32): 1211520. doi:10.1073/pnas.0605127103. PMC
1524930. PMID 16880384. Cheung L (31 July 2006).
Thousands of microbes in one gulp. BBC.

[28] Michod, Richard E. (1995). Eros and Evolution: A Natural Philosophy of sex. Reading, Mass: Addison-Wesley
Pub. Co. ISBN 0-201-44232-9.

[45] Dykhuizen, D. (2005). Species Numbers in Bacteria.


Proceedings. California Academy of Sciences 56 (6): 62
71. PMC 3160642. PMID 21874075.

11

[46] Zhong, S, Kodursky A, Dykhuizen DE, Dean AM


(2004). Evolutionary genomics of ecological specialization. Proceedings of the National Academy of Sciences of the United States of America 101 (32): 1171924.
doi:10.1073/pnas.0404397101. PMC 511043. PMID
15289609.
[47] Dykhuizen, DE & Baranton G (2001). Evolutionary
implications of promiscuous recombination in Borrelia.
Trends Microbiol 9 (7): 344350. doi:10.1016/S0966842X(01)02066-2. PMID 11435109.
[48] Rapp MS, Giovannoni SJ (2003). Theunculturedmicrobialmajority. Annual Review of Microbiology 57:
36994. doi:10.1146/annurev.micro.57.030502.090759.
PMID 14527284.
[49] Stackebrandt E, Frederiksen W, Garrity GM, Grimont
AD, Ka P, Maiden MCJ, Nesme X, Swings J, Tru HG,
Ward AC & Whitman WB (2002). Taxonomic Report of the ad hoc committee for the re-evaluation of the
species denition in bacteriology. Int. J. Syst. Evol. Microbiol. 52 (Pt 3): 10431047. doi:10.1099/ijs.0.023600. PMID 12054223.
[50] de Queiroz K (2005). Ernst Mayr and the modern concept of species. Proceedings of the National
Academy of Sciences of the United States of America 102
(Suppl 1): 66007. Bibcode:2005PNAS..102.6600D.
doi:10.1073/pnas.0502030102. PMC 1131873. PMID
15851674.
[51] Gevers D, Cohan FM, Lawrence JG, Spratt BG, Coenye
T, Feil EJ, Stackebrandt E, Van De Peer Y, Vandamme P,
Thompson FL, Swings J (2005). Opinion: Re-evaluating
prokaryotic species. Nature Reviews Microbiology 3 (9):
7339. doi:10.1038/nrmicro1236. PMID 16138101.
[52] Papke RT, Zhaxybayeva O, Feil EJ, Sommerfeld K, Muise
D, Doolittle WF (2007). Searching for species in haloarchaea. Proceedings of the National Academy of Sciences
104 (35): 140927. doi:10.1073/pnas.0706358104.
PMC 1955782. PMID 17715057.
[53] Robertson CE, Harris JK, Spear JR, Pace NR (2005).
Phylogenetic diversity and ecology of environmental Archaea. Current Opinion in Microbiology 8 (6): 63842.
doi:10.1016/j.mib.2005.10.003. PMID 16236543.
[54] Costello, M. J.; May, R. M.; Stork, N. E. (2013). Can We
Name Earths Species Before They Go Extinct?". Science
339 (6118): 413416. doi:10.1126/science.1230318.
PMID 23349283.
[55] Hawksworth, David L. (2001). The magnitude of
fungal diversity: The 1.5 million species estimate
revisited.
Mycological Research 105 (12): 1422.
doi:10.1017/S0953756201004725.
[56] Acari at University of Michigan Museum of Zoology
Web Page. Insects.ummz.lsa.umich.edu. 2003-11-10.
Retrieved 2012-11-25.
[57] http://www.amphibiaweb.org Amphibiaweb, accessed 27
Feb 2013
[58] http://www.reptile-database.org The Reptile Database,
accessed 27 Feb 2013

[59] Encyclopedia Smithsonian: Numbers of Insects. Si.edu.


Retrieved 2012-11-25.
[60] Census of marine life. Coml.org. Retrieved 2010-0423.
[61] Robin McKie and Zoe Corbyn (2005-09-25). Discovery
of new species and extermination at high rate. London:
Guardian. Retrieved 2010-04-23.
[62] Simpson, George G. (1945). The Principles of Classication and a Classication of Mammals. Bulletin of the
AMNH (New York: American Museum of Natural History) 85: 23.
[63] Chase, Bob (Autumn 2005). Upstart Antichrist. History Workshop Journal (60): 202206.

9 Further reading
Other Species Concepts U.C. Berkeley
2003-12-31, ScienceDaily: Working On The
'Porsche Of Its Time': New Model For Species Determination Oered
2003-08-08, ScienceDaily: Cross-species Mating
May Be Evolutionarily Important And Lead To
Rapid Change
2004-01-09 ScienceDaily: Mayo Researchers Observe Genetic Fusion Of Human, Animal Cells; May
Help Explain Origin Of AIDS
2000-09-18, ScienceDaily: Scientists Unravel Ancient Evolutionary History Of Photosynthesis

10 External links
Barcoding of species
Catalogue of Life
European Species Names in Linnaean, Czech, English, German and French
Speciation
Stanford Encyclopedia of Philosophy entry: Species
VisualTaxa
Wikispecies The free species directory that anyone
can edit from the Wikimedia Foundation

12

11

11
11.1

TEXT AND IMAGE SOURCES, CONTRIBUTORS, AND LICENSES

Text and image sources, contributors, and licenses


Text

Species Source: http://en.wikipedia.org/wiki/Species?oldid=639442869 Contributors: AxelBoldt, Tobias Hoevekamp, CYD, Mav, Bryan
Derksen, Zundark, Timo Honkasalo, Stephen Gilbert, Slrubenstein, Ap, Ed Poor, Andre Engels, XJaM, PierreAbbat, SimonP, Anthere,
Schewek, Graft, Heron, Ryguasu, Hephaestos, Ram-Man, Patrick, Smelialichu, Michael Hardy, Lexor, Gdarin, Shyamal, Menchi, Tannin,
Ixfd64, GTBacchus, Karada, Arpingstone, Goatasaur, Card, Ahoerstemeier, Jimfbleak, Bueller 007, Julesd, Glenn, Nikai, Andres, Evercat,
Crusadeonilliteracy, Andrewman327, Steinsky, Tpbradbury, Marshman, Samsara, Jni, Robbot, Fredrik, Peak, Seglea, Naddy, UtherSRG,
GerardM, SoLando, Pengo, Alan Liefting, Giftlite, DocWatson42, Jacoplane, MPF, Fastssion, Tom Radulovich, Alterego, Everyking,
Dratman, Curps, Jorend, Niteowlneils, Nikita Borisov, Semorrison, Bobblewik, Alanl, OldakQuill, Gdr, Zeimusu, Antandrus, Beland,
Onco p53, Lesgles, Kaldari, Vina, Szajd, Icairns, Jmeppley, Neutrality, Golnazfotohabadi, Thorwald, Mike Rosoft, Ta bu shi da yu, Spiy
sperry, Jiy, Rich Farmbrough, Cfailde, Vsmith, D-Notice, Dbachmann, AlanBarrett, Dmeranda, Brian0918, Tompw, El C, Shanes, Art
LaPella, RoyBoy, Adambro, Bobo192, Circeus, Shenme, Cmdrjameson, Wisdom89, Jerryseinfeld, Rajah, Nsaa, Schnolle, Orangemarlin,
Jumbuck, Alansohn, Gary, Plumbago, Wangry, Pippu d'Angelo, Malo, Wtmitchell, BanyanTree, Inge-Lyubov, R6MaY89, Dave.Dunford,
Pauli133, Ceyockey, Jackhynes, Stemonitis, Woohookitty, 2004-12-29T22:45Z, Kzollman, Niqueco, Benbest, WadeSimMiser, Kmg90,
Brendanconway, Dover, Ashmoo, BD2412, Edison, Sj, Rjwilmsi, Mayumashu, Koavf, , Wikibofh, Avia, HappyCamper, Neuron132, Boccobrock, Ucucha, Yamamoto Ichiro, FlaBot, Daderot, Nivix, RexNL, Otets, Choess, Alphachimp, Chobot,
Gdrbot, Roboto de Ajvol, YurikBot, Wavelength, RobotE, Adam1213, Phantomsteve, RussBot, Petiatil, Reo On, Spaully, Lalalalalala,
Bhny, Stephenb, Eleassar, Tavilis, Pelago, Captain B, Dysmorodrepanis, Snek01, Grafen, ErkDemon, ONEder Boy, Joelr31, Robert McClenon, Bobbo, Voidxor, Bota47, Trainra, Wknight94, FF2010, 21655, Citynoise, Lt-wiki-bot, Redgolpe, Noodleman, ArielGold, Allens,
Katieh5584, Kungfuadam, GrinBot, Elliskev, Victor falk,
robot, Snalwibma, SmackBot, Brya, Prodego, McGeddon, Shoy, Pgk,
C.Fred, Speight, WilyD, EncycloPetey, KVDP, Jrockley, Hardyplants, Abbeyvet, M c, Edonovan, Edgar181, Gilliam, Ohnoitsjamie,
Nativeborncal, RDBrown, Marsoult, MalafayaBot, SchftyThree, Poison iva, Baa, DHN-bot, Paalexan, Mike hayes, Can't sleep, clown
will eat me, Abyssal, Chlewbot, Snowmanradio, Yidisheryid, Phexxa, GeorgeMoney, Addshore, Stevenmitchell, Flyguy649, Kingdon,
Dreadstar, Richard001, Greg.collver, Candorwien, SashatoBot, Aldaniel, Euchiasmus, Mgiganteus1, Scetoaux, IronGargoyle, Beetstra,
Avs5221, Mr Stephen, Waggers, Sasata, Atakdoug, Levineps, KlaudiuMihaila, Dead3y3, Iridescent, Lakers, Tmangray, Paul Foxworthy,
Twas Now, Daniel5127, The Letter J, JForget, Jptdrake, CRGreathouse, John Wilkins, Ruslik0, Sax Russell, WeggeBot, Fordmadoxfraud,
Besidesamiracle, DonMacneill, Flowerpotman, Alvesgaspar, Pascal.Tesson, Michael C Price, Christian75, Tom David, Dyanega, Altaileopard, Mattisse, Thijs!bot, Epbr123, Giantsshoulders, Pjvpjv, Sobreira, Marek69, John254, Chunminghan, Bgold, Escarbot, Hmrox, AntiVandalBot, Zoologyteacher, Jj137, TimVickers, Chill doubt, Spencer, Mdz, Figma, Res2216restar, Sluzzelin, JAnDbot, Sophie means
wisdom, Jimmy, PhilKnight, Steveprutz, Yahel Guhan, Magioladitis, Bongwarrior, VoABot II, Redaktor, Wikiality123, PoisonedQuill,
KConWiki, Catgut, Indon, Thibbs, DerHexer, JaGa, Philg88, Khalid Mahmood, Telesiphe, FlopTopSet, Peter coxhead, Laura1822, Atulsnischal, MartinBot, Karebh, Reguiieee, Bookinhand, Petter Bckman, Tgeairn, J.delanoy, Philcha, Tony1212, Ginsengbomb, Adzyaye,
Glump, Dawright12, Katalaveno, C8755, Sauria, Vanished User 4517, Dagtho, Aervanath, Nadiatalent, Sunderland06, Jnp2109, Treisijs,
Inter16, CardinalDan, Idioma-bot, Speciate, Wikieditor06, VolkovBot, CWii, Hersfold, Gnostrat, AlnoktaBOT, Vlmastra, Philip Trueman,
TXiKiBoT, Technopat, Trans Arctica, Anna Lincoln, Steven J. Anderson, SuperSpeller22, Abdullais4u, LeaveSleaves, Scrantonian, Quindraco, Katimawan2005, Madhero88, Imacolt, Yes four, Synthebot, We.eter, IMNOTAVANDALK9, Temporaluser, Why Not A Duck,
Artat, Monty845, Busaccsb, Nagy, EmxBot, Rstafursky, SieBot, Dawn Bard, Caltas, God Emperor, Backwalker, Nathan, Sylverfysh,
Wing gundam, EcoNerd1986, JetLover, Glenmin, Lawerjax, Byrialbot, Eiddie123, Billy55566666, Enok Walker, Jplyer, Eshstevebaugh,
Ks0stm, Jruderman, BenoniBot, Stfg, Anchor Link Bot, Prkapoorvijay, Denisarona, Graminophile, Escape Orbit, TFCforever, Invertzoo, Twinsday, Martarius, ClueBot, The Thing That Should Not Be, Weiner2952, Surfspeczz, Rodhullandemu, Cygnis insignis, SuperHamster, Boing! said Zebedee, CounterVandalismBot, Blanchardb, LonelyBeacon, Peteruetz, Jethrocapone, Ammonight423, Mspraveen,
Aua, Alexbot, Jusdafax, Jeayman, Jerry Zhang, Sun Creator, Birdkid100, Promethean, Frozen4322, Tinymonty, BOTarate, Taranet,
Thingg, Np105034, Amaltheus, Burner0718, AC+79 3888, SoxBot III, Maky, Oldekop, Rror, Bradv, Nepenthes, Frood, MystBot, RyanCross, Addbot, Cac070489, DOI bot, Fyrael, AkhtaBot, Blethering Scot, TutterMouse, Fieldday-sunday, Devrit, MrOllie, Bernstein0275,
AnnaFrance, 5 albert square, Numbo3-bot, Wings Upon My Feet, Tide rolls, Teles, First Light, Ettrig, Legobot, Luckas-bot, Yobot, II
MusLiM HyBRiD II, THEN WHO WAS PHONE?, SwisterTwister, Chyeahdawg, AnomieBOT, Sucro, Eliorcohen, Standproud1, Materialscientist, Limideen, The High Fin Sperm Whale, Citation bot, OllieFury, Maxis ftw, Haidata, Xqbot, Timir2, St.nerol, Capricorn42,
4twenty42o, Ed.howland, Jmundo, Maddie!, Iliveincanadadry, Dr Oldekop, GrouchoBot, RibotBOT, CFH7, Scoot-Overload, Doulos
Christos, Jabrona, Chriss789, Josemanimala, A.amitkumar, , Moracamilo, FrescoBot, Pepper, Dendrid, Zhafts, Vallonio, VI, Ricardo Ferreira de Oliveira, Citation bot 1, Intelligentsium, Mba123, Pekayer11, Pinethicket, I dream of horses, HRoestBot, MJ94, MastiBot,
Ezhuttukari, Baby mami 101, SkyMachine, Elekhh, FoxBot, TobeBot, Animalparty, Xook1kai Choa6aur, ItsZippy, Garethball, SeoMac,
Antipastor, Jmelville17, JV Smithy, Acer123456, Tbhotch, Obsidian Soul, RjwilmsiBot, Ripchip Bot, PPdd, Ninukk, Aircorn, EmausBot,
John of Reading, Orphan Wiki, WikitanvirBot, Wiki gezza, Immunize, Look2See1, GoingBatty, The Mysterious El Willstro, Dmsdjing,
Tommy2010, Kamal11992288, Wikipelli, P. S. F. Freitas, Daonguyen95, Kryp, Access Denied, John KB, WeGoAndiamo, L Kensington, Rigel may, Peter M. Brown, Kranix, Lizardcuckoo, BengMog, Taztouzi1, ChuispastonBot, SemanticMantis, Sunshine4921, Teapeat,
Danielgdm, ClueBot NG, Satellizer, , PaleCloudedWhite, FaEu, Frietjes, Rezabot, Widr, Jimpscott,
, Titodutta, Goldenhotmail,
BG19bot, Mohamed CJ, M0rphzone, Trumpkinius, Quahog5News, ...hellohere..*, Piguy101, Jamesalbert1234, Awesomator1, Silvrous,
Wikih101, CitationCleanerBot, Wandytoo, NotWith, Scavenger-X, Ted Baenziger, Tottingshire, La marts boys, Coolman7777, Futurist110, 23mjbulls, Zeeyanwiki, Sminthopsis84, Webclient101, Tashcon, Pradvk, Ayaeau, Lugia2453, Frosty, Graphium, 77tons of coal,
OakRunner, Utilist, Leprof 7272, BitBus, Zorahia, Thelionsareontheprowl, Nayday, I am One of Many, Luke05tea13, Zenibus, Ginsuloft,
Coreyemotela, Hullingergr, Nestorin8, Ocialdrgamer, Tim Topolski, Wepo123h, Oggmus, Monkbot, Casster M, Correctwhatiswrong247,
Lionblayze, Upeepee, Trackteur, Placenage, Mason meg, Qwertyabc12398, HOLASOYRENE, Danymorrison, Antjr01 and Anonymous:
654

11.2

Images

File:Carl_von_Linn.jpg Source: http://upload.wikimedia.org/wikipedia/commons/6/68/Carl_von_Linn%C3%A9.jpg License: Public


domain Contributors: Nationalmuseum press photo, cropped with colors slightly adjusted Original artist: Alexander Roslin
File:Edit-clear.svg Source: http://upload.wikimedia.org/wikipedia/en/f/f2/Edit-clear.svg License: Public domain Contributors: The
Tango! Desktop Project. Original artist:

11.3

Content license

13

The people from the Tango! project. And according to the meta-data in the le, specically: Andreas Nilsson, and Jakub Steiner (although
minimally).
File:Issoria_lathonia.jpg Source: http://upload.wikimedia.org/wikipedia/commons/2/2d/Issoria_lathonia.jpg License: CC-BY-SA-3.0
Contributors: ? Original artist: ?
File:John_Ray_from_NPG.jpg Source: http://upload.wikimedia.org/wikipedia/commons/b/bc/John_Ray_from_NPG.jpg License:
Public domain Contributors: National Portrait Gallery, London: NPG 563 Original artist: Unknown
File:Question_book-new.svg Source: http://upload.wikimedia.org/wikipedia/en/9/99/Question_book-new.svg License: Cc-by-sa-3.0
Contributors:
Created from scratch in Adobe Illustrator. Based on Image:Question book.png created by User:Equazcion Original artist:
Tkgd2007
File:Tree_of_life.svg Source: http://upload.wikimedia.org/wikipedia/commons/0/09/Tree_of_life.svg License: CC-BY-SA-3.0 Contributors: ? Original artist: ?
File:Undiscovered_species_chart.png Source: http://upload.wikimedia.org/wikipedia/commons/7/78/Undiscovered_species_chart.png
License: Public domain Contributors: Own work Original artist: KVDP
File:Wikispecies-logo.svg Source: http://upload.wikimedia.org/wikipedia/commons/d/df/Wikispecies-logo.svg License: CC BY-SA 3.0
Contributors: Image:Wikispecies-logo.jpg Original artist: (of code) cs:User:-xfi File:Wiktionary-logo-en.svg Source: http://upload.wikimedia.org/wikipedia/commons/f/f8/Wiktionary-logo-en.svg License: Public domain Contributors: Vector version of Image:Wiktionary-logo-en.png. Original artist: Vectorized by Fvasconcellos (talk contribs), based
on original logo tossed together by Brion Vibber

11.3

Content license

Creative Commons Attribution-Share Alike 3.0

También podría gustarte