THE EVOLUTION OF DOMINANCE UNDER DISRUPTIVE SELECTION C. A. CLARKE and P. Ni. SHEPPARD Department of Medicine and Department of Zoology, University of Liverpool Received6.iii.59 1.INTRODUCTION INa paper on the effects of disruptive selection, Mather (1955) pointed out that if there are two optimum values for a character and all others are less advantageous or disadvantageous there will be disruptive selection which can lead to the evolution of a polymorphism. Sheppard (1958) argued that where such selection is effective and the change from one optimum value to the other is switched by a single pair of allelomorphs there will be three genotypes but only two advantageous phenotypes. Consequently if dominance were absent initially it would be evolved as a result of the disruptive selection, the heterozygote and one of the homozygotes both coming to resemble one of the two optimum phenotypes (see Ford, 1955, on Tripharna comes). Thoday (1959) has shown by means of an artificial selection experiment that, even when a character is, at the beginning, controlled polygenically (sternopleural chaeta-number in Drosophila) and there is 50 per cent. gene exchange between the "high" and "low" selected sub-popu- lations, a polymorphism can evolve. The most fully understood examples of disruptive selection (other than sex) are provided by instances of Batesian Mimicry, where there are a number of distinct warningly coloured species, acting as models, which are mimicked by the polymorphic forms of a single more edible species. Fisher and Ford (see Ford, 1953) have argued that a suffi- ciently good resemblance between mimic and model is not likely to arise by chance as the result of mutation. They therefore suggest that a mutant arises which gives a fair resemblance and that there is subsequent selection for modifiers which improve the resemblance controlled by the new mutant. However, if as Fisher (1928) believes, the effects of a new mutant are neither fully dominant nor fully recessive from the beginning, the modification of the phenotype towards more perfect mimicry would also involve the evolution of dominance since normally there will be only two optimum patterns, the mimetic and the original one. 2. MATERIAL Inthe course of an investigation into the genetics of the highly polymorphic mimetic butterfly Papilio dardanus Brown, we have obtained information which gives data on the evolution of dominance in a polymorphic species. P. dardanus is found in the Ethiopian region and occurs in several distinct races whose names 73 74 C. A. CLARKE AND P. M. SHEPPARD and geographical distribution are given in fig. i (for a detailed review see the excellent paper of Ford, 1936).Theraces which particularly concern us in this paper are (i) the South African race cenea (not to be confused with the mimetic form of the same name, which is found commonly in South Africa and mimics Amauris albimaculata and Amauris echeria), (2) the West African race dardanus, (3) the transitional race meseres, and ()theKenya race polytrophus. All these will be collectively known as the main African stock in this paper, since there is some gene exchange between them. Besides these races we shall also be considering two isolated ones, meriones from Madagascar and antinorii from Abyssinia. FIG. I. In P. dardanus the males are all rather similar in appearance having a black and yellow pattern which is non-mimetic and possessing" tails "on their hind wings, a character typical of the genus Papilio. In the main African stock the females are usually mimetic, always tailless, and, with the exception of some individuals in the polytrophus race, never have the yellow fluorescent pigment characteristic of the males. In Madagascar, as well as in race humbloti from Grand Comoro, the females are similar to the males in pattern, in possessing yellow fluorescent pigment and in being tailed. In Abyssinia all the females are tailed and the majority are like the males, but a minority have mimetic patterns. DOMINANCE AND DISRUPTIVE SELECTION 75 3.METHODS Themain variation of wing pattern in the forms to be considered occurs in three areas on the forewing and two on the hindwing. For convenience of description these areas have been given numbers in fig. 2.Twoother areas of interest are the cells of fore and hind wings which are also marked in the figure. Besides the wing pattern, there is also variation in tail length and in order to measure this it is necessary to determine fixed points of reference on the hindwing. The three points, given in fig. 2are,A at the apex of the cell nearest its point of attachment to the body, C the point where vein 4 reaches the margin of the wing and B the point where a straight line joining the distal end of veins 3 and 5 cuts vein 4. The length of the tail is taken to be the distance BC and the length of the wing to be BA. Because the edge of the wing is scalloped even insects with no tail have a small protuberance which is measured by BC. 10_H 12 area 3 area 4 I CB A Fic. 2 Since the size of the hindwing varies from insect to insect, from sex to sex and from race to race and since there is a marked correlation between tail length, BC, and wing size, BA, it is necessary to adjust tail length to take into account differences in the size of the insects before comparisons can be made between them. This was done by calculating for each insect the factor which would adjust the length of BA so that it equalled the average (358o mm.) for all insects examined and correcting tail length BC by this same factor. For example if the length of BA for a particular insect were 385O mm. and the tail length were 9 mm. this value would be adjusted by the factor to give a length of 837 mm. The procedure would not be legitimate if there were allometric growth. However, such a situation would lead to a correlation between adjusted tail length and un- adjusted wing length and as no such correlation is apparent we can conclude that any allometric growth that there may be is too small to be detected in this material and that the procedure of adjusting tail length in accordance with the length of 76 C. A. CLARKE AND P. M. SHEPPARD the wing is a valid one. Not only does this adjustment reduce differences in mean tail length between the sexes and races but it also materially reduces the sample variance. 4.THE FEMALE FORMS Thefemale forms investigated by us, which occur together in each race (i.e. aresympatric) are given in table i together with some indication as to their frequency TABLE i Approximate Reference Race Female form frequency Model for in per cent, frequency cenea 40-85 Amauris albimaculata, F., C. A. echeria hippocoonides 9-50 A. niaeius F., C. trophonius 4-15 Danaus chrysippus F., C. cenea natalica 0-10 None F., C. leighi <2 None F., C. Imperfect-trophonius 1-0 ? D. chrysippus cal. salaami <02 None cal. hippocoonides (hzpocoon) 6o-ioo A. niavius F. planemoides 0-40 Beinatistes poggei F. Transitional niobe 0-7 B. tellus F. and trophonius (trophonissa) 0-5 D. chrysippus F. dardanus cenea 0-7 A. echeria, F. A. albimaculata red-brown cenea o-o4 None cal. cenea 20-40 A. echeria, F. A. albimaculata proto-cenea 5-15 A. echeria, F. A. albiinaculata hippocoonides 20-40 A. niavius F. yellow 3-10 ?A. niavius F. polytrophus (proto-hippocoonides) pal e-poultoni None F. bright-poulloni 5-15 trophonius <I D. chrysippus F. proto-trophonius 5-15 ? D. chrysippus F. planemoides B. poggei F. proto-planemoides 5-IS ? B. poggei F. (yellow (male-like) 8o-ioo None M. Ihippocoonides (niavioides) 0-20 A. niavius M. Jtrophonius (?ruspinae) 0-5 D. chrysippus M. an iflOTli wienholti o-s ? B. epaea and M. I ? Acraeajodutta .cenea 0-s A. echeria M. meriones yellow (male_like) soo None F. In the references C. stands for Clarke et al., cal.for calculated, F. for Ford and M. for Museum and Mansfield collections. and mimetic status. Where two colour patterns are controlled by the same main allelomorph but differ in appearance because of differences in the gene-complexes of the various races they have usually been given different names by entomologists. DOMINANCE AND DISRUPTIVE SELECTION 77 In the table we have used one name for these genetically similar forms but have put in brackets the varietal name used by entomologists. The patterns are described with reference to the areas marked on fig. 2. Owing to the difficulty of describing colours accurately, where necessary, the reference numbers of the colour of a typical specimen have been taken from Villalobos and Villalobos (1947) and are put in parentheses. The numbers are given in the order hue, value and degree respectively. cenea (plate 3, no. 6) has the pale area 4 of the hindwing yellowish-buff (8-i8-8) and the spots on the forewing are white to pale yellowish-buff. hippocoonides (plate 3, no. 4) has all the pale areas white. The width of area 5 varies from race to race in parallel with that of a similar area on the local sub-species of the model. trophonim has the same black pattern as hippocoonides except that area 5 is always narrow, as is a similar area on its model. Areas and 4 are reddish-brown (7-16-12) and area i is white. natalica has hindwings like those of cenea. The black pattern of the forewings is the same as that of hippocoonides but areas i and 3 are yellowish-buff to ochreous (8-19-9) like the hindwings and not white.