VARIABLE SELECTION ON EUROSTA'S GALL SIZE,I: THE EXTENT AND NATURE OF VARIATION IN PHENOTYPIC SELECTION

Natural fluctuations in environmental conditions are likely to induce variation in the intensity or direction of natural selection. A long-term study of the insect, Eurosta solidaginins Fitch (Diptera; Tephritidae), which induces stem galls on the perennial herb Solidago altissima (Asteraceae) was performed to explore the patterns of variation in phenotypic selection. The intensity of selection imposed by parasitoids and predators on gallmaking larvae, for gall size, was measured across 16 populations over the course of 4 generations, for a total of 64 population-generations. Directional selection was quantified by i, the selection intensity, and variance selection by j‘, a measure of the intensity of selection on phenotypic variance. Size-dependent attack by parasitoids caused upward directional selection (mean i_p = 0.42; SE = 0.023), while size-dependent bird attack favored larvae that induced smaller galls (mean i_b = -0.07; SE = 0.013. The mean net directional selection intensity was 0.35 (SE = 0.030), which indicates that insects inducing larger galls are generally favored by selection. The opposing patterns of size-dependent attack resulted in stabilizing selection in half the population generations, with an overall average. j‘ of -0.11 (SE = 0.078). The magnitude of directional selection was strongly influenced by the population mean gall size and weakly by the optimal gall size. The intensity of variance selection was strongly influenced by the shape of the fitness function, with sigmoidal and Gaussian-like shapes causing greater depletion of phenotypic variance.     

Apostatic selection by song thrushes (Turdus philomelos) feeding on the snail Cepaea hortensis

There is no direct evidence that predators exert apostatic selection (the systematic overpredation of commoner forms) on live, naturally polymorphic prey. This study tested whether captive song thrushes ( Turdus philomelos ) select apostatically when presented with dimorphic populations of yellow five-banded and yellow unhanded morphs of the snail Cepaea hortensis. Four thrushes were used. Two were presented with a 9: 1 ratio of five-bandeds to unbandeds and two were presented with 1:9 ratios. Each thrush was given four trials in succession. In each trial 30 snails were presented and the trial was stopped when 15 had been eaten. There were no differences in shell size between morphs or between eaten and uneaten snails of each morph. Three thrushes selected apostatically and one thrush exerted virtually no selection. Overall, there was statistically significant apostatic selection.     

Do pollen beetles need pollen? The effect of pollen on oviposition,survival, and development of a flower‐feeding herbivore

Abstract.  1. Pollen is considered to be an important dietary component for many species of flower-feeding herbivores. Its influence on oviposition site selection by the pollen beetle Meligethes aeneus , and on the development of its larvae was investigated.
2. The effects of pollen presence and absence on adult, egg, and larval incidence in the field, and on larval development in the laboratory were compared through the use of Synergy, a composite hybrid oilseed rape Brassica napus variety comprising male-fertile (with pollen) and male-sterile (without pollen) plants.
3. In the field, adult females were more abundant on male-fertile plants during flowering, and a greater proportion of male-fertile than male-sterile buds were accepted for oviposition. These data indicate a possible role of pollen in oviposition site selection by female pollen beetles.
4. The numbers of first instar larvae on the two plant lines did not differ; however, more second instars were found on male-fertile than on male-sterile flowers. This suggests a greater larval survival on male-fertile plants, possibly due to the more readily available food resources and better nutrition afforded by the presence of pollen.
5. Laboratory experiments confirmed that a diet which included pollen improved survival to adulthood and resulted in heavier pupae and adults; however, pollen was not obligatory for larval survival and development.
6. The pollen beetle, previously thought to be an obligate pollen feeder, is therefore more generalist in its requirements for development. These findings may relate to the nutritional and behavioural ecology of other flower-feeding herbivores.     

Parasitoid behaviour: predicting field from laboratory

Abstract.  1. Most of what is known about parasitoid behaviour comes from laboratory observations: field quantitative observations on searching parasitoids are extremely difficult to do and are rare. The basic components of Aphytis melinus 's response to California red scale ( Aonidiella aurantii ) were studied in the laboratory: encounter, rejection, drumming, probing, oviposition, and host-feeding. It was then asked whether these observations provided a reliable guide to behaviour in the field in a situation that was very different from the laboratory.
2. Field observations were carried out on bark on the trunk and interior branches of trees where live scale density is extremely high in patches, dead scale make up 90% of all scale, and could be expected to interfere with Aphytis search.
3. The laboratory observations predicted well the time taken in the field for each basic event (drumming or probing) and average times spent on a scale. Also well predicted were the distributions of times spent on drumming, probing, and total time on a scale. Rejection rates were much higher in the field. Thus, the laboratory studies predicted foraging behaviour in the field with variable success; potential explanations for observed mismatch between laboratory and field and its possible larger implications are discussed.     

Reproduction and the central project of evolutionary theory

In much of the discourse of evolutionary theory, reproduction is treated as an autonomous function of the individual organism — even in discussions of sexually reproducing organisms. In this paper, I examine some of the functions and consequences of such manifestly peculiar language. In particular, I suggest that it provides crucial support for the central project of evolutionary theory — namely that of locating causal efficacy in intrinsic properties of the individual organism. Furthermore, I argue that the language of individual reproduction is maintained by certain methodological conventions that both obscure many of the problems it generates and serve to actively impede attempts to redress those difficulties that can be identified. Finally, I suggest that inclusion of the complexities introduced by sexual reproduction — in both language and methodology — may radically undermine the individualist focus of evolutionary theory.I am indepted to the Rockefeller Foundation for a Humanities Fellowship that supported this research during the spring of 1986. I am also grateful to Richard Lewontin, Diane Paul, and Lisa Lloyd for many extremely helpful conversations.     

Evidence for the retention of genetic variation in Erodium seed dormancy by variable rainfall

Summary The periodic occurrence of summer/early autumn precipitation in the California annual grassland can result in the formation of early and late emerging cohorts of Erodium botrys and E. brachycarpum. The occurrence of early rainfall and the timing of such rainfall are highly variable from year to year. A series of field watering experiments in 1980–81 were used to simulate early emergence conditions that would result from significant rainfall (1 cm) occurring in mid-July, late August, and mid-September. Net reproduction was used to estimate fitness differentials between Erodium cohorts emerging in response to a watering treatment (early emerging cohorts) and Erodium cohorts emerging with the onset of winter rains in mid-October (late emerging cohorts). Survival was lower and gross reproduction was higher among early emerging cohorts than late emerging cohorts. For both species, net reproduction of the early cohort was lower than that of the late cohort under the July watering treatment and higher than that of the late cohort under the August watering treatment.Early cohorts, formed in response to rainfall in mid-September, 1982, were also compared demographically to later cohorts emerging in October. Compared to late cohorts, net reproduction, gross reproduction and survival were higher for the early cohorts.Common garden experiments indicate that differences in the duration of seed dormancy between the progenies of early and late emerging plants reflect a significant genetic component. Progency produced by early cohorts of E. brachycarpum from all three watering treatments possessed more extended seed dormancy than progeny of late cohorts. In E. botrys, progeny from early cohorts emerging in response to the July watering treatment were also more dormant than late progeny. In contrast, early cohorts of E. botrys emerging in response to the September watering treatment produced seed less dormant than seed produced by late cohorts. When combined with demographic data, indicating that fitness differentials between early and late cohorts varied with changes in the date of early emergence, genetic results suggest that year to year variation in early rainfall may act to retain genetic variation in the duration of seed dormancy.