Saudi Journal of Biological Sciences xxx (2017) xxxxxx

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Saudi Journal of Biological Sciences

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Original article

Species concept and speciation

Amal Y. Aldhebiani
Department of Biological Sciences, Faculty of Sciences, King Abdulaziz University, P.O. Box 80200, Jeddah 21589, Saudi Arabia

a r t i c l e i n f o a b s t r a c t

Article history: Defining and recognizing a species has been a controversial issue for a long time. To determine the vari-
Received 25 January 2017 ation and the limitation between species, many concepts have been proposed. When a taxonomist study
Revised 8 March 2017 a particular taxa, he/she must adopted a species concept and provide a species limitation to define this
Accepted 30 April 2017
taxa. In this paper some of species concepts are discussed starting from the typological species concepts
Available online xxxx
to the phylogenetic concept. Positive and negative aspects of these concepts are represented in addition
to their application.
Keywords:
2017 Production and hosting by Elsevier B.V. on behalf of King Saud University. This is an open access
Species concept
Species limitation
article under the CC BY-NC-ND license (http://creativecommons.org/licenses/by-nc-nd/4.0/).
Species
Taxonomy
Classification

1. Introduction but by defining what a species is, they also clarify what speciation
is. Therefore, research programs result on speciation with concen-
Since the early ages of the living world man has been using clas- trating on conditions and factors are relying on species concept.
sification in his normal live. Having used plants for different pur- Moreover, many biological studies depend on delimitation of spe-
poses, he divided them to edible, medicinal, fuel and etc. The cies. For instance, ecological and behavioral studies may examine
plant kingdom as other living kingdoms has a hierarchy structure the collaboration between species, including the interaction
ends mostly with species rank. The term species has been unde- between species abundance with biotic and abiotic factors. Most
fined precisely for more than three centuries. It has different mean- works in conservation biology emphasize on species. Even in legis-
ings for different botanists including both systematist and lation, species is one of the most used biological units, not only in
taxonomist. Species are one of the basic units to compare in almost conservation and biology but also in agricultural and medical dis-
all fields of biology, from anatomy to behavior, development, ecol- putes. Therefore, a precise clear definition of the term species is
ogy, evolution, genetics, molecular biology, paleontology, physiol- needed for a larger spectators than just the academic biologists
ogy, and systematics. Largely, the importance of species stems (Hausdorf, 2011). For a better understanding of the development
from its significance in systematics, which is a science all the of species concept, the history of major ones will be overviewed
branches of biology rely on (de Queiroz, 2005). A main issue in in the first section of this paper. Then, some of the modern species
the field of Biological systematics is developing a theory to lead concepts will be discussed. Finally, a conclusion of the application
the taxonomic practices in building a persistent biological classifi- of these concepts will be addressed.
cation (Velasco, 2008). In addition, taxonomic and systematic liter-
ature are always referring to issues about species such as,
speciation models, whether species even exist in nature is taxon- 2. History of defining the term species
omy, and delimitation of species which usually involve species
concept. Species concepts do not only define what a species is, The earliest systematist to achieve a species definition was the
English scholar John Ray (16271705). In his major work Historia
Plantarum (Ray, 1686) he stated: No matter what variations occur
Peer review under responsibility of King Saud University. in the individuals or the species, if they spring from the seed of one
and the same plant, they are accidental variations and not such as
distinguish a species permanently; one species never springs from
the seed of another nor vice versa. Therefore, Roy attempt to
Production and hosting by Elsevier define species as groups of plant truly breed within their limits
E-mail address: aaldhdebiani@kau.edu.sa of variation.

http://dx.doi.org/10.1016/j.sjbs.2017.04.013
1319-562X/ 2017 Production and hosting by Elsevier B.V. on behalf of King Saud University.
This is an open access article under the CC BY-NC-ND license (http://creativecommons.org/licenses/by-nc-nd/4.0/).

Please cite this article in press as: Aldhebiani, A.Y. Species concept and speciation. Saudi Journal of Biological Sciences (2017), http://dx.doi.org/10.1016/j.
sjbs.2017.04.013
2 A.Y. Aldhebiani / Saudi Journal of Biological Sciences xxx (2017) xxxxxx

Linnaeus (17071778), nearly 50 years later whose work was 4.1. Biological species concept
the most eminent and momentous in the taxonomy field, adopting
a broader concept gave a new definition of species. In his work Spe- In nineteen century the first who produced the most quoted
cies Plantarum (Linnaeus, 1753), using mainly the floral structure definition of what he called biological species was the zoologist
and sexual characters, Linnaeus described briefly and systemati- Mayr (1942) who defined species as: groups of actually or poten-
cally approximately 5900 species of plants known to man then. tially interbreeding natural populations which are reproductively
So, he used a sexual system natural system for defining species. isolated from other such groups. Thus groups of related plants
Linnaeus concept was simple, applicable and was accepted widely. which are distinct at the level of biological species do not inter-
Both Ray and Linnaeus approaches in defining species were breed when growing in the same area in nature. This explains sim-
typological; they believed that under natural intraspecific varia- ply what is called now the Biological Species Concept (BCS) which
tions exists a fixed unchangeable type of each species and this is a non-phylogenetic species concept because it is potentially
refute the Ancient Greek idea of transmission of species which interbreeding process with no references of ancestry. In another
was widely believed in those days (Briggs and Walters, 1984). words a species is a group of reproducing natural populations inca-
De Candolle (17781841) who was the first to introduce the pable to effectively mate or breed with other such groups, and
word taxonomy through his book Thorie lmentaire de la Botani- which inhabits a particular niche in nature (Mayr, 1982; Bisby
que (De Candole, 1813) defined species as a species is a collection and Coddington, 1995). Although this theory is so simple and obvi-
of all the individuals which resemble each other more than they ous, it has mainly two disadvantages. First it is inapplicable onto
resemble anything else, which can by natural fecundation produce asexual organisms. Secondly, it is impractical in instances of allo-
fertile individuals, and which reproduce themselves by generation, patric populations (geographically isolated) (Cronquist, 1978;
in such a manner that we may from analogy suppose them all to Stace, 1989).
have sprung from one single individual. He divided plants into Both isolating species concept and recognition species concept
two major groups, non-vascular and vascular plants. A book in can be part of biological concept or genetic concept because both
seven volumes covered all species of cotyledons in the world see the species as a field for gene recombination. While the isolat-
including 161 families and 58000 species was his main and most ing concept stresses on reproductive isolation as the mechanism
important work which is called Prodromus Systematis Naturalis responsible for discontinuity between species (each species is
Regni Vegetabilis (18231873). Most of taxonomist after him, reproductively isolated from all other species, precluding them
before the next phase of taxonomy, used De Candolles system from mixing their genes and their traits), recognition concept
sometimes with some modifications. stresses on reproductive coherence as the factor responsible for
Years later, in his theory of evolution with Wallace (Darwin and continuity within species. They also have the same disadvantages
Wallace, 1859) even before his famous book On the origin of species as the biological concept inapplicable on asexual organisms and
by means of natural selection Charles Darwin (18091882) consid- impractical on allopatric.
ered species as the fundamental units of evolution, starting a
new era of species definition. Darwin emphasized the fact that spe- 4.2. Morphological species concept (MSC)
cies could be produced rapidly if the conditions were appropriate
and in the absence of such conditions, species might remain Cronquist (1978) adopting this concept he defined species as
unchanged for a long time. All Darwin theories and studies were the smallest groups that are constantly and determinedly distinc-
carried out before the establishment of the genetic science. tive and distinguishable by average means. Thus, species are the
In the middle of the twenty century, exactly in 1920s, a new smallest natural populations permanently separated from each
science was born. The science of genetics which focus on the math- other by a distinct discontinuity in the series of biotype (Du
ematical expression of the properties of populations and the ways Rietz, 1930; Bisby and Coddington, 1995). In other words, morpho-
in which such properties could be maintained or altered has made logical species concept states that a species is a community, or a
powerful contribution in understanding the species evolution. number of related communities, whose distinctive morphological
Since then, many modern species concepts were developed. characters are, in the opinion of a competent systematist, suffi-
ciently definite to entitle it, or them, to a specific name (Regan,
3. Modern species concepts 1926). It can be applied to sexual and asexual organisms and it is
also useful for species concepts in the fossil record. However,
Based on the previous theories and with more studies, tax- sometimes morphological characteristics are subjective and
onomist proposed different approached of species concepts in depend on expert opinion for key traits. And in some cases the
modern science. To approach a satisfactory and acceptable classifi- species are sympatric (morphologically indistinguishable) but are
cation the relationship between individuals should be considered. clearly different lineages.
These relations could be phenetic or phylogenetic. The term phe-
netic is applied to classification system which relies on similarities 4.3. Ecological species concept (ESC)
between present properties of organisms with no consideration or
references in how they possess them. Morphology, cytology, phy- The ecological species concept is mainly about ecological com-
tochemistry, anatomy, embryology and even some generic features petition. Van Valen (1976) stated: A species is a lineage (or a clo-
are considered to be source data for phenetic way. On the other sely related set of lineages) which occupies an adaptive zone
hand, when the relationship describes the pathways of ancestry minimally different from that of any other lineage in its range
(how the characters of organisms arose in evolution regardless and which evolves separately from all lineages outside its range.
their present day state) it is called cladistic which is the same as Colinvaux (1986) also wrote: A species is a number of related pop-
phylogenetic (Heywood, 1976). ulations the members of which compete more with their own kind
than with members of other species. In other words, when two
4. Definition of species concepts organisms are similar to each other, their needs are more likely
to overlap, therefore, they are expected to contest, and conse-
In the following a discussion of several species concepts known: quently the more likely that they are of the same species.

Please cite this article in press as: Aldhebiani, A.Y. Species concept and speciation. Saudi Journal of Biological Sciences (2017), http://dx.doi.org/10.1016/j.
sjbs.2017.04.013
 A.Y. Aldhebiani / Saudi Journal of Biological Sciences xxx (2017) xxxxxx 3

Nevertheless, the ecological species concept has some complications

4.7. Phylogenetic species concept (PSC)
subsequently it requires that the life histories for members of indi-
vidual species are the same which practically is not always true. It
With the presence of Darwin and Wallace theory of evolution,
also has a problem similar to the morphological species concept the rediscovery of G. Mendels laws of inheritance in 1900 and
which is: at what point does one stop the process of splitting the development of the modern theory of chromosome, all these
divergent forms into new species?. Finally, it is not always signif- led to the cladistic speciation. Simply it defines species as a group
icant to determine the degree to which two or more entities are of organisms that share an ancestor. In other words species are
competing ecologically. individuals show a high degree of resemblances in many unique
traits which give a monophyletic clusters based on discriminative
4.4. Evolutionary species concept phenotypes. This concept integrates character- based concepts that
emphasize the presence of an apparent organism attribute with
An evolutionary species is a single lineage of ancestor- history based concepts that emphasize the degree of relatedness
descendant populations of organisms which maintains its identity of a new isolate to previously characterized organism. Comparing
from other such lineages [in space and time] and which has its own with BCS this concept is applicable on both sexual and allopatric
evolutionary tendencies and historical fate (Wiley, 1981). This populations. However, it runs into two great practical problems,
concept was developed by Simpson (1951) in order to include it is rarely possible to reconstruct with certainty the past evolu-
asexual organisms and extinct species whom the biological species tionary pathway; and if so, it is hardly possible to devise a satisfac-
concept could not be applied to. The problem in this evolutionary tory method of designation a branching pattern by means of a
concept arose when the gaps in the fossil record levy prejudice lim- single linear sequence which is so important in flora and system-
its between species, especially those which experiencing regular atic treatment. However, many attempts have been made to pro-
size/shape evolution. duce such a system, the aim being to construct a sequence
starting with the most primitive and ending with the most
4.5. Cohesion species concept advanced; ensuring that each taxon recognized is a monophyletic
or polyphyletic (Stace, 1989; Agapow et al., 2004).
A cohesion species is an evolutionary lineage that serves as the
arena of action of basic micro evolutionary forces, such as gene 4.8. Pluralistic species concept
flow (when applicable), genetic drift and natural selection
(Templeton, 1994). Thus the cohesion concept is similar to the evo- When a given species concept is favored in a given conditions,
lutionary species concept in a way that a population genetic stress that does not mean it could be universally applicable. For under-
on the origins of phenotypic similarity within species. standing all species living at all times, a broader concept of species
should be applied. A comprehensive concept larger than any spe-
cies concept indicated above. The need to use more than one spe-
4.6. Phenetic species concept
cies concepts in order to be applicable arose the idea of a pluralistic
species concept. This recognizes, basically, that the factors that
Based on the idea that species concept shouldnt be bound to
are most important for the cohesion of individuals as a species
any precise theory Ridley (1993) gave this definition: A species
vary (Campbell and Reece, 2002).
is a set of organisms that look similar to each other and distinct
from other sets. Thus, it would clarify some particular degree of
phenetic resemblance, and similarity would be measured by a phe- 5. Conclusion
netic remoteness statistic.
Practically, the phenetic concept measures as many characters These species concepts mentioned above are some of the others
as possible in as many organisms as possible, and then identifies present in taxonomic world. There are many others (ex. composite,
phenetic clusters by multivariate statistics. The smallest unit in internodal, genetic and etc.). With this large number of concepts it
these clusters has sufficient similarity to be called a species. The is not an easy or simple decision to adapt one. Generally, it
theory of phenetic species concept can be opposed on the bases depends on the criteria and the aim of each project. For example,
of that, to a specific degree there is a resemblance between any biological, isolation and recognition concepts can be used if the
two objects in the universe. Moreover, members of the same spe- organisms were sexual breeding and from same community or
cies can be significantly different (especially in polytypic species) geographical area. In addition, if a study is concerned on the simi-
and individuals of various species may look more related to each larity of a group of plant with enough information of characters
other than members of the same species. Therefore, to achieve a (morphology, anatomy, cytology) with no need to a lineage, a mor-
better classification based on phenetic similarity some principals phological concept could be adapted. Sometimes more than one
should be followed (Stace, 1989): concept can be used, for example the ecological concept can be
used with the morphological one.
* A great content of information and more characters in the taxa Moreover, many concepts may be used with phenetic concept
concerned should be attained. too. Although some taxonomists believe that phenetic classification
* Each trait has an equal weight when forming natural taxa. represent a more practical solution than phylogenetic ones (Raven,
* Overall similarity between two taxa (units) is a result of their 1976; Heywood, 1978) phenetic or numerical taxonomy does not
individual resemblance among the many traits used to compare generate new data and is not a new system of classification. It is
them. only a new method of organizing data and obtaining classification
* Taxa can be identified and recognized because correspondences or special presentation forms from them. Therefore, it can combine
of characters differ in the groups of organisms under more than one concept by using their data in numerical forms.
investigation. On the other hand cladistic or phylogenetic basically view evo-
* Taxonomy is usually viewed and practiced as an empirical lution as an ordered and divergent transformation of characters.
science. So, it is trying to uncover populations genealogical relationships

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4 A.Y. Aldhebiani / Saudi Journal of Biological Sciences xxx (2017) xxxxxx

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Please cite this article in press as: Aldhebiani, A.Y. Species concept and speciation. Saudi Journal of Biological Sciences (2017), http://dx.doi.org/10.1016/j.
sjbs.2017.04.013