Poulton, Wallace and Jordan: how discoveries in Papilio butterflies led to a new species concept 100 years ago

"A hundred years ago, in January 1904, E.B. Poulton gave an address entitled 'What  is a species?' The resulting article, published in the Proceedings of the Entomological Society of London, is perhaps the first paper ever devoted entirely to a discussion of species concepts, and the first to elaborate what became known as the 'biological species concept'. Poulton argued that species were syngamic (i.e. formed reproductive communities), the individual members of which were united by synepigony (common descent). Poulton's species concept was informed by his knowledge of polymorphic mimicry in Papilio butterflies: male and female forms were members of the same species, in spite of being quite distinct morphologically, because they belonged to syngamic communities. It is almost certainly not a coincidence that Alfred Russel Wallace had just given Poulton a book on mimicry in December 1903. This volume contained key reprints from the 1860s including the first mimicry papers, by Henry Walter  Bates, Wallace himself and Roland Trimen. All these papers deal with species concepts and speciation  as well as mimicry, and the last two contain the initial discoveries about mimetic polymorphism in Papilio: strongly divergent female morphs must belong to the same species as non-mimetic males, because they can be observed in copula in nature. Poulton, together with his contemporaries Karl Jordan and Walter Rothschild, who had monographed world Papilionidae, were strongly influential on the evolutionary synthesis 40 years later. Ernst Mayr, in particular, had collected  birds and butterflies for Walter Rothschild, and had visited Tring, where Jordan worked, in the 1920s. The recognition of different kinds of reproductive and geographic isolation, the classification of isolating mechanisms, the use of the term sympatry, and the biological species concept all trace back to Poulton's 1904 paper.Poulton's paper, in turn, inherits much from Wallace's 1865 paper on Asian Papilio contained in the very book Wallace gave Poulton a month earlier. Wallace's gift, and Poulton's subsequent New Year address are thus key events in the history of species concepts, systematics and evolutionary biology."