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Principios de Taxonomia

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94j 5 Diversity within the Species: Polymorphisms and the Polytypic Species<br />

One should also not confuse species-specific phenotypic traits with the genome.<br />

Many DNA sequences of the genome are neutral and thus, are largely not controlled by<br />

natural selection. They mutate and diverge between separated populations by chance<br />

and, hence, indicate evolutionary distances, but not species-specificity. Traits that are<br />

controlled by natural selection can be very similar among the individuals of a species,<br />

even if the individuals of the species have evolutionary diverged due to geographical<br />

distance. In turn, selection- controlled traits can be similar between the individuals of<br />

different species, even if these individuals have evolutionary largely diverged.<br />

Although genetic differences are a benchmark for evolutionary closeness or distance,<br />

some populations that are genetically very similar could belong to different species,<br />

whereas, in turn, other species that are evolutionarily old and geographically wi<strong>de</strong>ly<br />

distributed and whose individuals differ genetically could belong to the same species.<br />

Theforcesofnaturalselectioncanresultinphenotypicdivergenceswithinaspecies,<br />

even in the presence of appreciable gene flow. This phenomenon is called intraspecific<br />

polymorphism. Intraspecific polymorphism is a serious problem for trait-oriented<br />

taxonomy (Ford, 1954). Different traits do not necessarily mean different species.<br />

Because most traits have nothing to do with maintaining species differences, two<br />

individuals of the same species can be significantly more different than two individuals<br />

of two different species (P€a€abo, 2001; Avise, Walker, and Johns, 1998). Intraspecific<br />

divergences, in some cases, can be greater than the differences that are found<br />

between congeneric, but distinct, species (Smith, Schnei<strong>de</strong>r, and Hol<strong>de</strong>r, 2001).<br />

5.2<br />

Differences in Traits do not Necessarily Mean Species Differences<br />

On page 424 of the first edition of The Origin of Species, Darwin writes (cited in<br />

Ghiselin, 1997) the following: With species in a state of nature, every naturalist has<br />

in fact brought <strong>de</strong>scent into his classification; for he inclu<strong>de</strong>s in his lowest gra<strong>de</strong>, or<br />

that of a species, the two sexes; and how enormously these sometimes differ in the<br />

most important traits, is known to every naturalist: scarcely a single fact can be<br />

predicated in common of the males and hermaphrodites of certain barnacles<br />

(Cirripedia), when adult, and yet no one dreams of separating them. There is no<br />

more accurate expression of how fraught with problems it is to represent the position,<br />

this organism looks different, so it must belong to a different species.<br />

Darwin s adversary, and friend, Alfred Russel Wallace, titled a publication released<br />

in 1858: On the ten<strong>de</strong>ncy of varieties to <strong>de</strong>part in<strong>de</strong>finitely from the original type.<br />

(Wallace, 1858). With this title, he expressed that species are everything but<br />

homogeneous. Species tend to vary in<strong>de</strong>finitely in their traits. How can it then<br />

be acceptable to make the phenomenon of trait differences into a species criterion? It<br />

is a difficult problem to compare the nature of the trait differences among the<br />

organisms within a species with the nature of the trait differences between the<br />

organisms of different species (see Chapter 4).<br />

Only the traits that keep populations separate from each other have something to<br />

do with the <strong>de</strong>finition of what a species is. Only these traits are worthy of being called

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