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Principios de Taxonomia

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6.31 The Problem of Smooth Boundaries between two Gene-Flow Communitiesj183<br />

In addition, there is also the phenomenon of maternal inheritance. The homozygotically<br />

recessive l/l combination does not affect the individual itself that possesses<br />

this genotype, but only the structure of the cytoplasm of the eggs that are<br />

produced by this individual, if it is a female. The structural organization of the<br />

cytoplasm <strong>de</strong>termines the direction of coiling of the daughter snails that <strong>de</strong>velop<br />

from these eggs. Right- or left-han<strong>de</strong>dness consequently does not affect the generation<br />

that has the phenotype, but only the subsequent generation.<br />

What is important for the snail species problem is the fact that right- and lefthan<strong>de</strong>dness<br />

each result in mating barriers. A right-han<strong>de</strong>d snail cannot copulate with<br />

a left-han<strong>de</strong>d one. The copulation position is such that the penis and vagina do not fit<br />

if two mirror-inverted, differently han<strong>de</strong>d snails want to mate. This phenomenon<br />

alone offers the possibility of a rapid reproductive isolation and thus the formation of<br />

a new species, which would require the left-han<strong>de</strong>d snails pooling together.<br />

That these are favorable conditions for species formation is further supported by<br />

the fact that because of the maternal inheritance, the l allele can already preadaptively<br />

spread in a population, without the left-han<strong>de</strong>d snails phenotypically appearing in<br />

this population. This can statistically lead to a sud<strong>de</strong>n appearance of many lefthan<strong>de</strong>d<br />

snails, which can then build up a population of many individuals that is<br />

strong enough to compete with the right-han<strong>de</strong>d snails.<br />

The statistical chance that an autonomous left-han<strong>de</strong>d population will build up in<br />

an existing right-han<strong>de</strong>d population is still a matter of controversial discussion. Allen<br />

Orr (Orr, 1991), however, has posited via a computer simulation that an entire snail<br />

population can turn from a right-han<strong>de</strong>d population into a left-han<strong>de</strong>d population.<br />

Such a population would immediately become isolated from the other populations.<br />

This would then be a one-gene speciation. This mo<strong>de</strong>l is not purely speculative, for<br />

among fossil snails, examples can be found of snail species that became left-han<strong>de</strong>d<br />

in a relatively short evolutionary period.<br />

The capability of an immediate speciation in only one generation has already been<br />

mentioned for tetra- and polyploid organisms that can emerge in plants. If a plant<br />

doubles its chromosomal set and thus becomes tetraploid, gene flow is instantly<br />

disrupted because the tetraploid plant can no longer backcross with the parental<br />

plants. Tetraploidy is a remarkable form of speciation that happens without the<br />

alteration of even a single gene because no gene in the newly evolved species is<br />

different from those in the original species (Schilthuizen, 2001).<br />

These examples again make clear that the species that is <strong>de</strong>fined as a <strong>de</strong>limited<br />

gene-flow community is a fundamentally different concept from the species as a<br />

phylogenetically distinct group of organisms, which is the kind of species that is<br />

aimed at by the barcoding approach (Chapter 4).<br />

6.31<br />

The Problem of Smooth Boundaries between two Gene-Flow Communities<br />

A frequently observed misun<strong>de</strong>rstanding of the species concept of the gene-flow<br />

community lies in the problem of <strong>de</strong>limitation. Gene flow barriers are such

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