Principios de Taxonomia
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180j 6 Biological Species as a Gene-Flow Community<br />
6.29<br />
Gene theft between two Species of Green Frogs (Pelophylax ridibunda<br />
and P. lessonae)<br />
The well-known European Water Frog (Pelophylax esculenta) is a taxonomically<br />
problematic case whose status as a species is disputed. The Water Frog is not a<br />
distinct species; it is not reproductively isolated from the two species to which it is<br />
related, the Marsh Frog (P. ridibunda) and the Pool Frog (P. lessonae) (Color Plate 8).<br />
The Water Frog is diagnostically distinguished from the Marsh Frog and the Pool<br />
Frog, but it does not form a separate gene-flow community that is isolated from the<br />
other two species.<br />
Pelophylax esculenta results from a hybridization of two species, the Marsh<br />
Frog and the Pool Frog. No doubt, the Water Frog is a hybrid. The Water Frog,<br />
however, exists only as an F1 product and not as an F2 or even an F3 product. The<br />
Water Frog, as an F1 hybrid with the genomes of the Marsh Frog and the Pool Frog,<br />
does not continue into a second generation. Instead, the Water Frog must be<br />
repeatedly recreated. The hybrid status of the Water Frog initially resembles the<br />
many known cases of species hybrids, which result from acci<strong>de</strong>nts without<br />
evolutionary importance, because they do not reproduce among themselves any<br />
further but die off again at the end of their individual lives (see above). The Water<br />
Frog, however, is not an exceptional transient species. Instead, it is found permanently<br />
in most areas.<br />
Water Frogs reproduce very well among themselves, as can be observed in the<br />
gar<strong>de</strong>n pond. They are vital and completely fertile, but if two Water Frogs reproduce<br />
with each other, then only part of the offspring of this reproduction are Water Frogs;<br />
other offspring are again pure Marsh or Pool Frogs. The offspring of the Water Frog<br />
do not continue a distinct line of Water Frogs. The remarkable hybridization between<br />
the Marsh Frog and the Pool Frog does not produce a new line of organisms that<br />
would then exist separate from Marsh and Pool Frogs because there is no autonomous<br />
Water Frog genome that exists separately from the genomes of the Marsh and<br />
Pool Frogs and could pursue a separate evolutionary line. The Water Frog is not a third<br />
species alongsi<strong>de</strong> Marsh and Pool Frogs that evolved hybridogenically. Instead, the<br />
Water Frog evolves again and again anew.<br />
How is this explained? It starts with a hybridization between the two species,<br />
Marsh Frog and Pool Frog. The Marsh and Pool Frog mate with each other unrestrictedly;<br />
there is no prezygotic mating barrier. In doing so, they do not forfeit their<br />
i<strong>de</strong>ntities. The hybrid, the Water Frog, is vital and fertile. The zygote resulting from<br />
the hybridization, as well as all the Water Frog s somatic cells, are allodiploid; they<br />
contain a genome of the Marsh Frog and a genome of the Pool Frog. Its germ line cells<br />
are allodiploid, but only for a certain time in the ontogenetic <strong>de</strong>velopment of an<br />
individual Water Frog. Before meiosis starts in the testes of the male or in the ovaries<br />
of the female, that is to say, before the preliminary germ cells start to differentiate into<br />
spermatogonia or oogonia, one of the two genomes is completely removed from the<br />
preliminary germ cells. Accordingly, the premeiotic germ cells only contain one of<br />
the two parental genomes. Only the somatic cells of the Water Frog are equipped with