Verbal and quantitative genetic models of sexually antagonistic coevolution suggest that coevolutionary arms races should be common. Sexual selection favors exaggeration of male persistence traits that are costly to females, and females, in turn, are selected to resist these traits. The heightened resistance by females is thought to then favor further exaggeration in the male trait, leading to an escalating coevolutionary arms race between persistence and resistance traits. Much of this theory, however, is based on an (implicit) assumption that there are tight constraints on how female resistance can evolve. We develop a theory that identifies and relaxes these constraints, allowing female resistance to evolve in a fashion that better reflects known empirical patterns in the evolution of female preference functions (the resistance trait). Our results suggest that evolutionary arms races that lead to the exaggeration of persistence and resistance will be much less common than formerly predicted. Females sometimes evolve indifference to male traits rather than resistance and can even evolve to discriminate against these traits. These alternative outcomes depend on the existence of genetic variance in the components of the female sensory system underlying female resistance and on the strength of natural selection acting on these components. Female indifference tends to evolve when natural selection on the sensory system is weak, and under these conditions, sexually antagonistic coevolution tends not to reduce female fitness significantly at equilibrium. When natural selection on the female sensory system is strong, however, then arms races are more likely, and female fitness is then sometimes significantly depressed at equilibrium. Sexually antagonistic coevolution is thus likely to have strong deleterious effects on population fitness only when female sensory traits are under strong natural selection to perform functions in addition to those involved with mating. Together, these results suggest that identifying the nature of genetic variation in and the strength of natural selection on female resistance should be a central goal of future studies of sexual conflict.
It is quite common in studies of life-history plasticity to find a negative relationship between the age at which various life-history transitions occur and the growth conditions under which individuals develop. In particular, high growth typically results in earlier transitions, often at a larger size. Here, we use a relatively general optimization model for age and size at life-history transitions to argue that current life-history theory cannot adequately explain these results. Specifically, most such theory requires key assumptions that are unlikely to be generally met. This suggests that some important component of the biology of many organisms must be missing from many of the models in life-history theory. We suggest that this missing component might be the phenomenon of developmental thresholds. There are at least two different types of developmental thresholds possible, and we incorporate these into our general optimality model to demonstrate how they can cause a negative relationship between growth conditions and age at a transition. If developmental thresholds are common throughout taxa, then this might explain the empirical results. Our model formulation and analysis also formalizes the popular Wilbur-Collins hypothesis for age and size at metamorphosis in amphibians. The results demonstrate that optimal combinations of age and size, and the slope of the reaction norm connecting them, depend on the existence and type of threshold assumed. Our results also provide an evolutionary framework that can be used to view the data and many of the proximate submodels derived from the Wilbur-Collins hypothesis.
The long‐term economic benefits of `patch' spraying are likely to be related to the initial spatial distribution of the target weeds, the demographic characteristics of the species and the weed control and crop husbandry practices to which they are subjected. This paper describes a stochastic simulation model developed to investigate the interaction between weed seed dispersal and patch spraying. Simulated weed plant and seed populations are generated and compared with data from field observations. Lloyd's Patchiness index is used to quantify the patchiness of the weed density distribution, and the parameter k of the negative binomial distribution is used as a measure of distribution shape. A method of assessing the spatial scale of weed aggregation is proposed, in which spatial weed density information is transformed into the frequency domain, using a discrete two‐dimensional Fourier transform. In this paper, we simulate `on/off' patch spraying (full or zero herbicide application rate). A quantitative analysis of the effects of sprayer resolution and weed seed dispersal range on the herbicide reduction and yield benefits from patch spraying is performed for three initial spatial seedbank distributions. The model is parameterized for the grass weed Alopecurus myosuroides Huds. Herbicide is applied in square areas (whose size is defined by the spatial resolution of the sprayer) in which mean weed density is greater than or equal to one plant m–2. For a system conforming to this specification we show that for the control of A. myosuroides, it is unlikely that patch spraying would be profitable in the long term if the control area is larger than 6 m × 6 m. In some circumstances higher resolution may be required.
A model is presented to explore how the form of selection arising from competition for resources is affected by spatial resource heterogeneity. The model consists of a single species occupying two patches connected by migration, where the two patches can differ in the type of resources that they contain. The main goal is to determine the conditions under which competition for resources results in disruptive selection (i.e., selection favoring a polymorphism) since it is this form of selection that will give rise to the evolutionary diversification of resource exploitation strategies. In particular, comparing the conditions giving rise to disruptive selection when the two patches are identical to the conditions when they contain different resources reveals the effect of spatial resource heterogeneity. Results show that when the patches are identical, the conditions giving rise to disruptive selection are identical to those that give rise to character displacement in previous models. When the patches are different, the conditions giving rise to disruptive selection can be either more or less stringent depending upon demographic parameters such as the intrinsic rate of increase and the migration rate. Surprisingly, spatial resource heterogeneity can actually make forms of evolutionary diversification such as character displacement less likely. It is also found that results are dependent on how the resource exploitation strategies and the spatial resource heterogeneity affect the population dynamics. One robust conclusion however, is that spatial resource heterogeneity always has a disruptive effect when the migration rate between patches is low.
Existing optimality models of propagule size and number are not appropriate for many organisms. First, existing models assume a monotonically increasing offspring fitness/propagule size relationship. However, offspring survival during certain stages may decrease with increasing propagule size, generating a peaked offspring fitness/propagule size function (e.g., egg size in oxygen-limited aquatic environments). Second, existing models typically do not consider maternal effects on total reproductive output and the expression of offspring survival/propagule size relationships. However, larger females often have greater total egg production and may provide better habitats for their offspring. We develop a specific optimality model that incorporates these effects and test its predictions using data from salmonid fishes. We then outline a general model without assuming specific functional forms and test its predictions using data from freshwater fishes. Our theoretical and empirical results illustrate that, when offspring survival is negatively correlated with propagule size, optimal propagule size is larger in better habitats. When larger females provide better habitats, their optimal propagule size is larger. Nevertheless, propagule number should increase more rapidly than propagule size for a given increase in maternal size. In the absence of density dependence, females with greater relative reproductive output (i.e., for a given body size) should produce more but not larger propagules.
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