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What’s in a name: Species concepts just for plants, or a common concept for all of life?

What’s in a name: Species concepts just for plants, or a common concept for all of life?

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An essay for the 2012 Undergraduate Awards (International Programme) Competition by Audrey Chia. Originally submitted for Evolution and Ecology of Plants at University of Edinburgh, with lecturer Richard Milne in the category of Life Sciences
An essay for the 2012 Undergraduate Awards (International Programme) Competition by Audrey Chia. Originally submitted for Evolution and Ecology of Plants at University of Edinburgh, with lecturer Richard Milne in the category of Life Sciences

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Published by: Undergraduate Awards on Aug 31, 2012
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What’s in a name: Species concepts just for plants, or a common concept for all of life?
Charles Darwin wrote in
The Origin of Species
published more than 200 years ago
No one definition hasas yet satisfied all naturalists; yet every naturalist knows vaguely what he means when he speaks of aspecies
Despite being a fundamental concept to evolutionary biology, that ambiguity has lasted tilltoday and there has yet to be an agreement on the
definition of ‘species’. The many differences that
exist between major groups of organisms have led to different basis of comparison used to study themand the various species concepts that are used today. The most frequently used Biological SpeciesConcept, for example, cannot be applied to asexually reproducing organisms or hybrid organisms. Otherexisting species concepts, such as the Morphological Species Concept and Ecological Species Concepthave also been criticised for the ambiguity of their definition and their inapplicability to certain groupsof organisms.Several groups of plants undergo apomixis, hybridisation, and other biological processes that affectspeciation, and common knowledge is that plants undergo these processes more often than animals do.These properties make the use of a common species concept difficult, and some biologists have even
suggested that plant scientists ‘go it alone’ with a species concept that
caters to plants. Moreover, manyother species concepts have been formulated with the aim of tackling the limitations of existing speciesconcepts and accommodate the specific plant groups. It would indeed be understandable that groups of plant which undergo special processes should be studied using concepts most suitable for them.However, the emergence of the many new concepts for plants may distract biologists from realising that
those ‘special properties’ may not be significant enough to warrant a whole new species conce
pt just forplants, neither are they unique to plants alone.A common species concept may allow consistency in taxonomic practice, and would also be useful forthe storage and retrieval of information. It may also be practical for the use in policy-making andcommunication between scientists and the masses. However, while there are concepts that seem to fitthe bill of being applicable to all organisms, they tend to be only theoretical, and not suitable forpractical scientific use of delimiting species. With the diverse types of organisms that exist and thevariations produced by constant evolution of new life forms, a single species concept aimed toencompass all living things may still be inconceivable. It is vital to understand how each species conceptis very relevant and important to the study of the organisms and biological processes they are targetedat. The approach more practical and relevant to scientific study will be to employ different speciesconcepts to different groups of organisms, depending on which of them are most appropriate for theunderstanding of the organisms in question at the time of study.
It is these ‘species’ that will allow the
continual pursuit of a deeper understanding of them.
Species, Species concepts, Species delimitation, Plants, Taxonomy
A species concept that can be applied to all organisms has eluded biologists for decades. Mayden (1997)listed 22 different species concepts that existed, many of them variants of each other, and some rarelyapplied (Spooner et al., 2005). While many different species concepts are still adopted today, it has beenrecognised that different concepts are often incompatible to each other. Many of the species conceptshave been criticised for their inapplicability to certain groups of living organisms, such as asexual ormorphologically cryptic organisms (Niklas, 1997). Newer concepts have hence been formulated with theaim of tackling those limitations. The most widely adopted species concept so far, the
Biological SpeciesConcept 
(Donoghue, 1985; de Queiroz, 2005), has been rejected by plant scientists due to its failure toaccount for groups of plants that undergo processes such as asexual reproduction and hybridization. Ithas been suggested that plants should have a different species concept from animals due to thewidespread nature of these biological processes in plants (Grant, 1981). This essay will discuss thespecies concepts that have been applied to the study of plants, and the issues that are thought tochallenge the suitability of these species concepts. It will also examine the feasibility of a commonspecies concept for all living things.
The many species concepts used to study plants
Different species concepts employ different basis of comparison between organisms (e.g. reproductiveisolation, morphological differences or niche occupation) (De Queiroz, 2007). This can result in differentconclusions being made about the boundaries between species and the number of species that exists(Baum and Donoghue, 1995). The more frequently used species concepts relevant to the study of plantsinclude
Morphological Species Concept, Biological Species Concept, Ecological Species Concept 
Phylogenetic Species Concept 
. Each of them has faced criticisms, and several new concepts have beendeveloped to target the limitation of these species concepts.
The Morphological Species Concept 
The Morphological Species Concept (MSC)
defines a species as ‘a diagnosable
cluster of individualswithin w
hich there is a pattern of ancestry and descent, and beyond which there is not’
(Eldredge andCracraft, 1980). Because of its practicality for taxonomic purposes (Spooner et al., 2005), themorphological species concept has been frequently used. Cronquist (1978) found that most plantmonographers used morphological species concept as the criteria to delimit species, with some of themattributing this choice to the lack of other sources of data. Despite being widely used even today (Soltisand Soltis, 2009), it has long been recognized that classifying species based on morphology can besubjective and very misleading. For example, convergent evolution, simple morphology, and naturalvariation within species (such as sexual polymorphism or phenotypic plasticity), can confoundmorphological classification (Niklas, 1997). Due to strong local adaptations or introgressive hybridization,individuals that are actually genetically different and unrelated may be very similar morphologically(Baum and Donoghue, 1995). Different individuals of the same species which vary phenotypically may
also be incorrectly recognized as different species. Also, although separated by reproductive barriersgenetically,
sibling species
are not easily distinguishable based on morphology or anatomy (Mishler andDonoghue, 1982). When a barrier to reproduction occurs relatively recently, morphological differenceswill not have set in yet, although species are not able to cross the barrier. These
incipient species
,morphologically similar yet reproductively isolated,
are still considered as the same species (Mayr, 1996).Furthermore, plant species that differ radically in terms of morphology can produce viable hybrids thatare genetically complex (Tucker, 1953; Clausen et al., 1940), and this again challenges the validity of theMSC.Phylogenetic systematics has also led to biologists turning away from the MSC as it does not trackhistorical relationships (Baum and Donoghue, 1995). Hence, despite the wide use of the MSC today, it isgenerally recognised that it is not a good candidate for a species concept for plants, and many otherorganisms.
Ecological Species Concept 
Van Valen (1976) introduced the Ecological Species Concept (ESC) based on the observation that despitemany existing hybrids of oak (genus
), individual species are able to maintain their integrity intheir respective habitats. The ESC
describes a species as ‘a lineage which occupies an adaptive zone
minimally different from that of any other lineage in its range, and which evolves separately from otherlineages outsi
de its range’ (Van Valen, 1976), emphasizing the close relationship between ecological
factors and genetic differences. This concept has received criticism
, since ‘adaptive zones’ and ‘range’
are not easy to identify in nature, and this is even truer for plants (de Queiroz, 2007).
Phylogenetic Species Concept 
The Phylogenetic Species Concept (PSC)
defines species as ‘an irreducible cluster of organi
sms,diagnosably distinct from other such clusters, and within which there is parental pattern of ancestry anddescent
(Cracraft, 1983). This concept tracks evolutionary and historical relationships as well aspatterns of divergence (Davis and Manos, 1991), and these are not covered by the other speciesconcepts aforementioned. It also does not depend on specific kinds of biological processes, and hencecan be broadly applied to all organisms.However, it is difficult to define
a ‘diagnosable cluster’
, and it has been argued that different datasources will frequently lead to conflicting phylogenetic results (Wendel and Doyle, 1998). Also, Agapowet al. (2004) estimated that using the PSC leads to a 49% higher number of species, as compared to non-phylogenetic species concept, because it focuses on every discernible difference between organisms,even down to single molecular markers (Davis and Mannos, 1991). PSC will partition organisms into toomany separate biological entities which may become meaningless in the evolutionary sense. A case inpoint is the application of this concept to members of the species
Brassica oleracea.
Using the PSC,broccoli, Brussels sprout and cabbage will be unwarrantedly classified as different species (Niklas, 1997).

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