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Principios de Taxonomia

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5.14 The Color Pattern Polymorphism of the Shells of the Brown-Lipped Snail Cepaea nemoralisj117<br />

from one to five bands. There are 20–30 different morphs within a single population.<br />

What are the reasons for this diversity? Why is the best-adapted type not dominant? It<br />

cannot be the case that all of the morphs, with their differing colorations and patterns,<br />

have precisely the same chances for survival and reproduction. Why does one<br />

genotype not replace the other? Visual selection by predators or climatic selection<br />

should certainly repeatedly prefer one or another phenotype. And if this represents a<br />

case of a variable adaptation to different ecological niches, why then does the differing<br />

niching not lead to assortative mating and thus to sympatric speciation (Chapter 6).<br />

Whether C. nemoralis exhibits bands may have something to do with the likely<br />

primary predators of the Brown-lipped Snail in several areas: song thrushes (Turdus<br />

philomelos) (Jones, Leith, and Rawlings, 1977). In this regard, many investigations<br />

have been performed to <strong>de</strong>termine whether the extent of banding or the coloration of<br />

the snail s shell has beneficial camouflage effects. In<strong>de</strong>ed, it was found that song<br />

thrushes have more difficulty spotting ban<strong>de</strong>d snails in the vegetation than they do<br />

spotting non-ban<strong>de</strong>d snails. This result does not, however, explain the polymorphisms<br />

observed in this species of snails. On the contrary, the polymorphism should<br />

eventually disappear, at least in those locations where the song thrush is the primary<br />

predator of the snail.<br />

Climate factors may exert a selective influence on the presence and color of bands<br />

in C. nemoralis. Dark shells absorb more thermal radiation, whereas yellow shells<br />

reflect light more strongly. The color of the shell therefore influences the temperature<br />

of the snails. Exposure to the sun is crucial. Too much sun leads to heat <strong>de</strong>ath, but too<br />

little sun leads to the cold-bloo<strong>de</strong>d snails being too stiff and having limited mobility,<br />

which in turn puts the animals at risk for starvation. However, this explanation is also<br />

dissatisfying. If exposure to sunlight is the primary factor in the persistence of the<br />

multiple morphs, why are so many different morphs found at the same location?<br />

C. nemoralis lives in a large geographical range in Europe and is found from<br />

Norway to Spain and from the coast up to an elevation of 1200 meters. One can<br />

observe C. nemoralis in habitats as diverse as dunes, meadows and forests. Admittedly,<br />

the morphs are not uniformly distributed. Instead, some regions are dominated<br />

by certain shell variations. However, it is not the case that every geographical<br />

location, every mountain peak or every habitat hosts its own specific morphs.<br />

Furthermore, it has been observed that different morphs remain in the sun for<br />

differing time periods (Jones, Leith, and Rawlings, 1977). Some morphs prefer the<br />

cool morning hours for their activities, others are active around noon, and still others<br />

are predominantly active in shadows. However, no conclusive connection between<br />

the ambient temperature and the snails coloration or <strong>de</strong>gree of banding has been<br />

observed. In most cases, scientists cannot assign the specific color patterns to the<br />

individual climatic factors of the morphs respective habitats; the situation is<br />

too complex.<br />

The driving force for the intraspecific diversity of Cepaea snails appears to be that<br />

increased species diversity leads to an increase in the number of potentially habitable<br />

environments. By increasing their own diversity, the snails expand their resources<br />

and thus increase their population size. This would without doubt be a selective<br />

advantage. However, the question of why snails living in different niches have not

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