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Principios de Taxonomia

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5.19 The Mimicry Morphs of the Female Swallowtails of the Genus Papilioj123<br />

advantage of being amenable to cities, which are rich in food and safety, has led to<br />

significant changes in allelic frequency distributions in several bird species in the<br />

course of only a few generations.<br />

5.19<br />

The Mimicry Morphs of the Female Swallowtails of the Genus Papilio<br />

Intraspecific morphs were of particular concern to taxonomists when it came to<br />

changing taxonomy from a purely trait-oriented grouping process to one that was<br />

more scientific. More than one hundred years ago, Edward Bagnall Poulton recognized<br />

the inconsistency and contradictory nature of a species concept based only on<br />

traits. The importance of his classical publications What is a species? and The<br />

conception of species as interbreeding communities, published in 1904 and 1938 in<br />

the Proceedings of the Entomological Society of London and the Proceedings of<br />

the Linnaean Society of London respectively (see Mallet, 2004), can hardly be<br />

un<strong>de</strong>restimated. These publications stripped the Linnaean typology from its scientific<br />

foundation and exposed the classification by traits to be a subjective grouping<br />

according to our own pragmatic objectives and trait assessments, that is, that species<br />

were, as Darwin said in 1859, ma<strong>de</strong> for our convenience (Darwin, 1859). Moreover,<br />

Poulton anticipated Ernst Mayr s essential lines of thought by <strong>de</strong>ca<strong>de</strong>s, proposing the<br />

fundamental concept of the species as a reproductive community.<br />

Poulton had been influenced by the observations by Alfred Russel Wallace and<br />

Henry Walter Bates of mimicry morphs in butterflies (Mallet, 2004). Bates primarily<br />

investigated South American Heliconiids, whereas Wallace discovered the fascinating<br />

mimicry morphs of the Southeast Asian Swallowtails Papilio memnon, P. polytes<br />

and P. aegeus. There are several different morphs among the females of these<br />

butterfly species. In contrast, there are no morphs among the males, which have<br />

uniform shapes. The different female morphs mimic butterfly species of entirely<br />

different families, which are inedible to birds. The caterpillars of the mimicked<br />

species consume poisonous substances that are still present in the body of the<br />

imaginal stage butterfly. The mimics do not contain any unpalatable substances, but<br />

due to their similar appearance, the imitators (just as the imitatees) are protected<br />

from predators. This has an immense selective advantage and is a classic case (and<br />

historically one of the first published examples) of Batesian mimicry, referring to the<br />

imitation of a dangerous or inedible animal by a harmless animal to mislead the<br />

latter s predators (Wickler, 1968).<br />

For Poulton, the importance of these butterflies lay most of all in the fact that<br />

erroneous conclusions could be drawn when phenotypes were used to <strong>de</strong>termine<br />

species affiliation. The females of swallowtail butterflies can differ much more<br />

strongly from members of their own species than they do from the members of other<br />

swallowtail species. The same phenomenon was later observed in the African<br />

Swallowtail Papilio dardanus, which is today counted among the best known<br />

examples of Batesian mimicry (Mallet, 1995). Here, too, the males are uniformly<br />

shaped, whereas the females are polymorphic. In both Madagascar and Ethiopia, only

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