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Principios de Taxonomia

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6.17 Sympatric and Allopatric Speciationj157<br />

First, it is based on the assumption of a coinci<strong>de</strong>ntal separation of two populations<br />

caused by external forces, such as, for example, geological forces or climatic events,<br />

and second, it is based on the assumption that, in the course of the allopatric<br />

lifetimes, mutations that lead to genetic incompatibility accumulate coinci<strong>de</strong>ntally.<br />

Neither of these two coinci<strong>de</strong>nces is foun<strong>de</strong>d on a quantifiable scientific theory.<br />

The allopatric paradigm is nothing but the putting-into-words of an ad hoc concept<br />

(Tautz, 2009).<br />

If allopatrically separated populations become different, then selection has no<br />

interest in erecting prezygotic crossing barriers because the populations, which<br />

have become different, do not encounter each other anyway. For this reason,<br />

allopatric speciation usually needs a long time (Orr, 2009), while sympatric speciation<br />

can be a matter of only a few generations because it is stimulated by positive selection.<br />

If the allopatrically separated populations were to meet again later, then the<br />

differences that have originated by chance in allopatry can be an obstacle to remating.<br />

However, these differences do not necessarily have to be an obstacle.<br />

Mayr has <strong>de</strong>fen<strong>de</strong>d allopatric speciation as the only way in which species can<br />

originate (Mayr, 1963). The theory that speciation must almost always be allopatric<br />

has consequences that cannot readily be brought into agreement with our current<br />

knowledge of speciation. For example, according to the allopatric paradigm of<br />

speciation, those populations that live sympatrically without an external separation<br />

cannot split and, hence, would have to remain in existence eternally. As long as<br />

speciation must almost always be allopatric, a population can never un<strong>de</strong>rgo<br />

speciation un<strong>de</strong>r sympatric conditions because, without geographical separation,<br />

there can be no split. (In this consi<strong>de</strong>ration, anagenetic alterations of traits are, of<br />

course, not counted as speciation; see Chapter 7).<br />

Furthermore, the theory of allopatric speciation does not explain why there is rapid<br />

speciation in some groups of animals but not in other groups of animals, even though<br />

the groups should, in principle, have the same chance of being torn apart into isolated<br />

groups by external factors. For example, over approximately ten to a hundred<br />

thousand years, the Cichli<strong>de</strong>s of the African lakes have produced, in every lake in<br />

which they occur, ten to several hundred species, while the members of other families<br />

of fish in the same lakes did not (Verheyen et al., 2003) (see below). From this<br />

scenario, it clearly follows that the ability of a population to speciate also <strong>de</strong>pends on<br />

the internal genetic factors of the organism itself, and these factors are specific to the<br />

respective groups of animals. In these cases, speciation is <strong>de</strong>finitively not based on a<br />

separation by external factors but instead is based on the genetic constitution that is<br />

present, to a varying extent, in the different groups of animals.<br />

Additionally, the paradigm of allopatric speciation does not explain why the<br />

number of species is so large in beetles even though most beetles are able to fly<br />

and, therefore, could easily overcome many external barriers. The (apparently) very<br />

large number of beetle species, however, must be acknowledged with some care.<br />

Currently, the or<strong>de</strong>r of Coleoptera (beetles) is divi<strong>de</strong>d into approximately 400 000<br />

species. This or<strong>de</strong>r would be the or<strong>de</strong>r with the highest number of species in the<br />

animal kingdom except that the majority of these species are <strong>de</strong>limited by typological<br />

trait differences, which have often been <strong>de</strong>termined in only a small number of

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