Male‐like females, mimicry and transvestism in butterflies (Lepidoptera: Papilionidae)

Abstract

Abstract. When a butterfly species has a polymorphic female, with one of the forms closely resembling the male, it is customary to suppose that this form is ancestral, and that the ‘odd’ forms have arisen later. R. I. Vane‐Wright, on the other hand, has suggested that in some species the male‐like form may be a ‘transvestite’ female, the ancestral form of the female having been strikingly unlike the male. As later‐derived forms are usually, but not always, genetically dominant to ancestral forms, we can make some choice between these hypotheses by discovering the dominance relations of the male‐like and the ‘odd’ forms of the female. In the mimetic Papilio aegeus the male‐like form is shown to be recessive to the ‘odd’ (mimetic) form, as has essentially been the case in all other butterflies so far investigated. Papilio phorcas is now shown to be the exception: the ‘odd’ (non‐mimetic) form is recessive to the male‐like form. We conclude that usually the male‐like form is ancestral, but that P.phorcas may be an authentic example of ‘transvestism’, or the ‘transfer’ of male epigamic colour to the female of the species. The yellow, male‐like pattern of the mimetic Papilio dardanus may be dominant or recessive to the mimetic forms according to the genetic background: largely recessive in Madagascar, and southern and western Africa, dominant to most forms in Ethiopia, and probably dominant to one mimetic form but recessive to the others in Kenya. All female dardanus patterns, both mimetic and yellow, are strongly dominant to both female phorcas patterns in P. dardanus × P. phorcas hybrids (P. ‘nandina’). The simplest explanation of this situation is that the male‐like pattern of dardanus is ancestral, and that dominance has become locally reversed in Ethiopia. The dominance relations, and the sex‐ or autosomal‐linkage of two forms can be determined without pedigree‐breeding, simply by observing a few offspring each from a large number of wild‐caught females.

Genetics of the mimetic African butterfly Hypolimnas misippus: hindwing polymorphism

“…A conventional interpretation of the dominance order at the A locus would suggest that the ancestral female hindwing was orange and A is the more recent allele (following the reasoning of Haldane, 1924, explored further by Turner, 1977 andClarke et a!. 1985).…”

“…1, specimen 11) does it spread to the forewing; this was seen in only 47 per cent of 910 melanie butterflies subjected to scrutiny. In many cases melanism was associated with the blue iridescence that surrounds the white patches in the male, giving a decidedly male-like or transvestite (Clarke et a!., 1985) appearance. Iridescence was also apparent in the rare melanie forewings.…”

“…The dominance order of alleles is particularly relevant as it provides a guide to their mutation To whom correspondence should be addressed. sequence and spread in evolutionary time (Clarke, Clarke, Gill, Turner and Collins, 1985). Moreover, field evidence exists for strong, though intermittent, selection on female colour patterns at four different sites on five different occasions: Newton, Sierra Leone, 1966(Unamba, 1968; Legon, Ghana, 1965, 1966(Edmunds, 1969; Dar-es- Salaam, Tanzania, 1974(Smith, 1976Cape Coast, Ghana, 1977 (Gordon, 1982(Gordon, , 1987.…”

Polymorphism ofPhilaenus spumariusL. (Hemiptera: Cercopidae) in the Samsun (Mid-Black Sea Region) populations of Turkey

“…Numerous cases of transient and stable polymorphism of genes affecting pigmentation have been demonstrated in diverse groups (Kettlewell 1973;Lees 1981) and to a large extent the examples included, the snails (see Jones et al 1977), spiders (Oxford & Gillespie 1996), butterflies (Clarke et al 1985), moths (Lees & Creed 1977;Mikkola 1984), and beetles (Brakefield & Lees 1987). The common and widespread meadow spittlebug P. spumarius has been recognised for being a remarkable example because its heritable polymorphism can serve to explore interesting population variations ranging from molecular levels (Loukas & Drosopoulos 1992;Nor 1994;Darasi 1996) to visible polymorphisms (Halkka et al 1973;West 1990;Stewart & Lees 1996;Yurtsever 2000).…”

A review of deviant phenotypes in bees in relation to brood parasitism, and a gynandromorph of Megalopta genalis (Hymenoptera: Halictidae)

“…A multivariate analysis of structures supports this 'transsexual' hypothesis for a Lasioglossum (Paralictus) and its Lasioglossum (Dialictus) host (Wcislo, 1999). The expression of male-like characters in females is one developmental mechanism to generate novel phenotypes (WestEberhard, 2003), such as mimicry patterns (Clarke et al, 1985). The phenetic similarity between male character states and those of brood parasitic females by coincidence means that an asymmetric cross-sex expression of traits can readily produce structures that frequently are associated with obligate parasitic behaviour in insects (Wcislo, 1987(Wcislo, , 1999.…”