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Of Pieris Napi (Pieridae)

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Of Pieris Napi (Pieridae) 1964 Jaltrnal of the Lepidopterists' Saciely 91 THE MAINTENANCE FOR EXPERIMENTAL PURPOSES OF FORM "SULPHUREA" OF PIERIS NAPI (PIERIDAE) by S. R. BOWDEN 53 Crouch Hall Lane, Redbourne, Herts., E.'J'CLAND It may be guessed that when in 1909 Mr. H. W. Head (as he related 30 years later) received his "female specimen of P. napi of a pale yellow colour" from the teacher at Tullybeg School in Donegal, its primary importance to him was as a possible source of income; since he was a commercial breeder, he cannot be reproached for that. The bright yellow variety, previously almost unknown, that he ultimately bred from this Irish female, became known as "ab. 'citronea' Frohawk". It has given pleasure to European collectors ever since and for a long time this was the chief function that it fulfilled. However, more recently this form, properly known as "sulphurea" Schoyen, has provided a means of attacking a number of problems, both those that are special to the Pieris napi - bryoniae group of butterflies and those of a more general character that can be studied in that group. Its merits for these purposes arise primarily from the certainty with which it can be determined, its simple recessive inheritance and its nevertheless robust constitution. A particular advantage is its genetic control from the locus which also governs the characteristic parti­ coloured wild-type pattern of napi and related butterflies. EXPERIMENTAL USES OF FORM "SULPHUREA" (1) It may seem obvious that experiments involving the breeding of any species will be carried out most conveniently where that species is native. This is far from being the case. Though the food-plant will certainly be available, and fresh stock can be obtained with little difficulty, in default of quite exceptional precautions there will always be some risk of waifs' entering the experimental stock with gathered food. In some cases this may vitiate the experiment or, if it is unsuspected, even lead to incorrect conclusions. If, however, the whole of the ex­ perimental material can be marked with a bisexual recessive variation, this risk is eliminated: if the stock is homozygous, the intruder is recog­ nised when the adult emerges; if it must be heterozygous, the suspected intruder is detectable only by mating with the corresponding homozygote. ( 2) Successful pairing of a homozygous rece:;sive female with a like or an unlike male is demonstrated by raising her eggs to the ima­ ginal state. This was the means of disproving a statement that a female Pieris napi would pair successfully only once, and of showing that after the second pairing the first became ineffective (Bowden, 1951). Even 92 BOWDEN: Pieris napi VaLlS: no.2 the forced-pairing techniques of Lorkovic (1952) fall short of artificial insemination, but if the latter is ever applied to Pieris we may be sure that "sulphurea" will again prove useful. (3) The present author has used it to supply a "visible" napi gene which could be traced in hybrids with P. bryoniae Ochsenheimer for many generations (Bowden, 1962). Although there are many ban-iers, unsuspected at first, between napi and bryoniae, it appears to be possible to backcross the hybrids indefinitely to bryoniae. As long as the presence of "sulphurea" can be demonstrated, it remains clear that napi has not been completely "bred out" by any automatic selective process. ( 4) Petersen and Tenow (1954) described experiments in which they investigated the supposed sexual preference ban-iers between Pier is napi and bryoniae. They found, rather surprisingly, that P. hryoniae males were more attracted to napi females than to their own, the white colour being apparently the deciding factor. We do not presume to suggest what the result might have been, but we should have liked to see these experi­ ments extended to include females of form "sulphurea". (5) The question of selective pairing may arise, of course, within a species. It has impOltance as one of the mechanisms which may con­ tribute to the maintenance of a polymorphjsm (Williamson, 1958). Pair­ ing selection, if proved to exist, is difficult to interpret: it may be the result of differing visual stimulus, or it may depend on some apparently unrelated invisible character which has become con-elated with the visible one. Assuming that the basis is visual, "sulphurea" would provide good experimental material. ( 6 ) It is quite possihly true that, as long as good "resting" camouflage is retained, white colour is advantageous to Pieris. The disadvantage that we are inclined to associate with recessive inheritance probably lies, in the case of "sulphurea", in the extended yellow pigmentation. The breeder would judge that any invisible correlated disadvantage must be slight indeed. He might be of a very different opinion about the "albino" variety (with decolorized "black" markings) which turns up rather frequently in the Pieridae: here there may well be an accompanying weakness. Quasi-"natural" selection experiments using these two re­ cessives and wild-type might give interesting results. (7) A project that we have toyed with for some time is an attempt to detect "sulphurea" alleles in natural populations of Pieris napi. The heterozygotes of any rare recessive will be velY much less rare than the visible homozygotes; it is the heterozygote therefore that we may hope to find. Though there are several "sulphurea" alleles (Bowden, 1961), Head's "citronea" is perhaps the ultimate recessive. Captive virgin females of this form, mated with wild males collected at random, would 1964 Journal of the LepidoT'terists' Society 93 show by their offspring if any of these "type" males was in fact the heterozygote of a "sulphurea" allele. A further generation would have to be bred only to characterize fully all' allele already detected in this way. Although the best-known British "sulphurea" specimen came from Nor­ folk (Barrett, 1881), and Thompson's pale yellow had apparently a Flintshire origin, it is still the general opinion that Ireland would be the best place to search. Nevertheless, if "sulphurea" is maintained by recurrent mutation, it must surely exist (even if at very low frequency) in Asiatic and American subspecies. ( 8) The dominant white-underside morph "subtalba" Schima which is maintained in balanced polymorphism in Pieris bryoniae neobryoniae Shelj. has proved to be very closely linked (if not indeed allelic) with the recessive "sulphurea" (Bowden, 1963). "Subtallba"-like forms which occur in such distinct subspecies or species as dubiosa Rober or virgin­ iensis Edwards are probably also dominant, but there is no obvious way of determining experimentally whether the responsible genes are identi­ calor homologous. However, successful hybridisation and back-croSSing with napi f. "sulphurea" should allow a decision whether or not they are linked. (9 ) Pieris napi provides exceptionally favourable material for the study of pterin pigments and their genetic control, apparently surpassing in this respect any species of Colias. The sex-limited ochre-yellow of the various bryoniae subspecies is from many points of view of outstanding interest; this is increased by its relation to the lemon-yellow and white pigments controlled by the "sulphurea"-"subtalba" alleles, some of which (but not all) operate independently of sex. BREEDING FORM "SULPHUREA" A few notes on the maintenance of stocks of this useful and attractive variety of napi may be of some assistance, in spite of their European bias. First, we offer some remarks which apply to any subspecies of P. napi or bryoniae. The larvae may be reared by the usual techniques, which it is not necessary to describe in detail. They are less exacting than most species, and can sometimes even be raised successfully in the dark, with little ventilation, provided that paper or the like is supplied to absorb moisture. Crowding also seems to be harmless in itself, as long as it does not lead to excessive condensation, but of course if disease does appear congestion results in disaster. Pupating insects abo may be more fre­ quently disturbed by wandering larvae, and the chance of cannibalism is probably rather increased. If it is intended to breed napi continuously, it is unsafe to rely entirely on wild food. In the late summer clean leaves in good condition may 94 BOWDEN: P'ieris napi VoLlS: no.2 not be plentiful on wild plants, just when stocks of larvae are at their highest. In England, the most satisfactory food to grow is Dame's Violet, Hesperis matronalis L.; it has the advantage, too, of possessing decorative, sweet-scented flowers. Very "sappy" leaves are bcst avoided, but the plant has a very long productive season, certainly well into December in normal years, and being cut down will spring up again. The flowering shoots have strong stems and as they last well in water can be put in the laying cages for eggs. Some individual females lay on the flower petals (which drop before the eggs hatch); if this is found to be happening, the flowers and flower-buds must be removed from the shoot for that female. Subject to this same contingency, the flowers when available are a good source of nectar for the caged butterflies. Eggs and larvae of wild P. napi, as well as occasional EuchlOe cardamines L., may be found on H. matronalis; it should therefore be searched before it is put in the breeding cage or used as fodder. More troublesome on H esperis are microlepidoptera, particularly PluteZZa porrectella L. When an infestation is discovered, affected leaves should be destroyed and the plants sprayed twice with derris at a fairly high concentration. After thorough washing by rain or hosing with water, to remove insecticide residues, the plants are soon available again and usually remain free for some time.
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