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We develop a quantitative genetic model for conditional strategies that incorporates the ecological realism of previous strategic models. Similar to strategic models, the results show that environmental heterogeneity, cue reliability, and environment-dependent fitness trade-offs for the alternative tactics of the conditional strategy interact to determine when conditional strategies will be favored and that conditional strategies should be a common form of adaptive variation in nature. The results also show that conditional and unconditional development can be maintained in one of two ways: by frequency-dependent selection or by the maintenance of genetic variation that exceeds the threshold for induction. We then modified the model to take into account variance in exposures to the environmental cue as well as variance in response to the cue, which allows a derivation of a dose-response curve. Here the results showed that increasing the genetic variance for response both flattens and shifts the dose-response curve. Finally, we modify the model to derive the dose-response curve for a population polymorphic for a gene that blocks expression of the conditional strategy. We illustrate the utility of the model by application to predator-induced defense in an intertidal barnacle and compare the results with phenotypic models of selection.

The conditional expression of alternative phenotypes underlies the production of almost all life history decisions and many dichotomous traits, including male alternative reproductive morphs and behavioral tactics. Changes in tactic fitness should lead to evolutionary shifts in developmental switch points that underlie tactic expression. We used experimental evolution to directly test this hypothesis by rearing ten generations of the male-dimorphic mite Rhizoglyphus echinopus in either simple or three-dimensionally complex habitats that differed in their effects on morph fitness. In R. echinopus, fighter males develop weapons used for killing rivals, whereas scrambler males do not. Populations evolving in complex 3D habitats, where fighters had reduced fitness, produced fewer fighters because the switch point for fighter development evolved to a larger critical body size. Both the reduced mobility of fighter males and the altered spatial distribution of potential mates and rivals in the complex habitat were implicated in the evolutionary divergence of switch point between the habitats. Our results demonstrate how abiotic factors like habitat complexity can have a profound effect on evolution through sexual selection.

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JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range of content in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new forms of scholarship. For more information about JSTOR, please contact support@jstor.org.. Ecological Society of America is collaborating with JSTOR to digitize, preserve and extend access to Ecology.Abstract. Phenotypic plasticity is a widespread and often adaptive feature of organisms living in heterogeneous environments. The advantages of plasticity seem particularly clear in organisms that show environmentally cued switches between alternative morphs. Information concerning the presence and nature of variation underlying the induction of these morphs, especially under field conditions, would be valuable. Here we examined the basis for variation underlying a predator-induced defense in an intertidal barnacle (Chthamalus anisopoma). In a previous experiment, juvenile barnacles were exposed to a predatory gastropod (Acanthina angelica). Some of these individuals were induced to develop as a predation-resistant form, but other individuals developed as the default, undefended morph. Here we tested two alternative explanations for this observation. One, the "continuoussensitivity" model, holds that there is normally distributed genetic variation for sensitivity to the cue. This model predicts that, given sufficient exposure to the predator, all individuals would develop as the induced form; it suggests that the previous findings resulted from an insufficient dose of the cue. The second model, the "discontinuous-sensitivity" model, asserts that there is a genetic polymorphism for inducibility such that some individuals are not able to respond to the cue. This model suggests that, with repeated exposures to the predator, the resulting dose-response curve would reach an asymptote at <100%. We conducted a dose-response experiment in order to contrast these two alternatives, and to examine an expectation generated by life-history theory, namely, that repeated exposure to the predator would induce maturity at a younger age. With respect to the life-history prediction, we found no evidence to suggest that repeated exposure of juvenile barnacles to Acanthina affected the age at maturity, even though we found strong evidence for sizeselective attack by this predator. With respect to variation underlying induction to the defended morph, we obtained a dose-response curve showing a significant asymptote at about 22% induction, which is inconsistent with the continuous-sensitivity model. Hence the results indicate the possibility of a developmental polymorphism in the barnacle, but no indication of life-history shifts in response to the predator.

Seasonal polyphenism, in which different forms of a species are produced at different times of the year, is a common form of phenotypic plasticity among insects. Here I show that the production of dark fifth-instar caterpillars of the eastern black swallowtail butterfly, Papilio polyxenes, is a seasonal polyphenism, with larvae reared on autumnal conditions being significantly darker than larvae reared on midsummer conditions. Both rearing photoperiod and temperature were found to have individual and synergistic effects on larval darkness. Genetic analysis of variation among full-sibling families reared on combinations of two different temperatures and photoperiods is consistent with the hypothesis that variation in darkness is heritable. In addition, the genetic correlation in larval darkness across midsummer and autumnal environments is not different from zero, suggesting that differential gene expression is responsible for the increase in larval darkness in the autumn. The relatively dark autumnal form was found to have a higher body temperature in sunlight than did the lighter midsummer form, and small differences in temperature were found to increase larval growth rate. These results suggest that this genetically based seasonal polyphenism in larval color has evolved in part to increase larval growth rates in the autumn.

The tick Ixodes ricinus finds its hosts by climbing vegetation and adopting a sit-and-wait tactic. This “questing” behaviour is known to be temperature-dependent, such that questing increases with temperature up to a point where the vapor pressure deficit (drying effect) forces ticks down to rehydrate in the soil or mat layer. Little if any attention has been paid to understanding the questing of ticks from an evolutionary perspective. Here we ask whether populations from colder climatic conditions respond differently in terms of the threshold temperature for questing and the rate of response to a fixed temperature. We find significant variation between populations in the temperature sensitivity of questing, with populations from cooler climates starting questing at lower temperatures than populations from warmer temperatures. Cool climate populations also quest sooner when the temperature is held constant. These patterns are consistent with local adaptation to temperature either through direct selection or acclimation and challenge the use of fixed thresholds for questing in modeling the spread of tick populations. Our results also show how both time and temperature play a role in questing, but we are unable to explain the relationship in terms of degree-time used to model Arthropod development. We find that questing in response to temperature fits well with a quantitative genetic model of the conditional strategy, which reveals how selection on questing may operate and hence may be of value in understanding the evolutionary ecology of questing.

Associations between microhabitat, shell banding and apex colour were examined in the land snail Theba pisana. Snails on their summer aestivation sites were sampled from a transect that included a relatively sheltered Acacia habitat and a more exposed open habitat. The frequencies of fully banded snails and snails with dark apexes, as well as the intensity of banding in fully banded snails, were lower in the open habitat than in the Acacia habitat. No differences in microhabitat relative to shell phenotypes were found in samples from the open habitat. However, significant differences in microhabitat were found between effectively unbanded and fully banded snails in the Acacia habitat, with effectively unbanded snails more common in the exposed Acacia canopy and fully banded snails more common beneath the canopy. Air temperatures in the Acacia canopy were consistently higher than below the canopy, while body temperatures of living fully banded and unbanded snails in sunlight indicated that fully banded snails heat more rapidly than do unbanded snails. These results suggest a potential role for both climatic selection and adaptive plasticity in microhabitat choice in the maintenance of variation in shell banding.