In the Eyes of the Beholders: Female Choice and Avian Predation Risk Associated with an Exaggerated Male Butterfly Color

By Nathan I. Morehouse and Ronald L. Rutowski

The American Naturalist, Volume 176, Issue 6
DOI: 10.1086/657043

Color ornaments are often viewed as products of countervailing sexual and natural selection, because more colorful, more attractive individuals may also be more conspicuous to predators. However, while evidence for such countervailing selection exists for vertebrate color ornaments (e.g., Trinidadian guppies), similar studies have yet to be reported in invertebrates. Indeed, evidence for female mate choice based on extant variation in male coloration is limited in invertebrates, and researchers have not explicitly asked whether more attractive males are also more conspicuous to predators. Here we provide evidence that more chromatic male cabbage white butterflies (Pieris rapae) are more attractive to females but should also be more conspicuous to predators. Female P. rapae preferentially mate with more chromatic males when choosing from populations of males with naturally occurring or commensurate, experimentally induced color variation. Mathematical models of female color vision confirm that females should be able to discriminate color differences between prospective mates. Further, chromatic and luminance contrast scores from female visual system models better predicted male mating success than did measures of male color derived more directly from color spectra. Last, models of avian color vision suggest that preferred males should be more conspicuous to known avian predators.

Evolution of Nutrient Uptake Reveals a Trade-Off in the Ecological Stoichiometry of Plant-Herbivore Interactions

By Pedro Branco, Maayke Stomp, Martijn Egas, and Jef Huisman

The American Naturalist, Volume 176, Issue 6
DOI: 10.1086/657036

Nutrient limitation determines the primary production and species composition of many ecosystems. Here we apply an adaptive dynamics approach to investigate evolution of the ecological stoichiometry of primary producers and its implications for plant-herbivore interactions. The model predicts a trade-off between the competitive ability and grazing susceptibility of primary producers, driven by changes in their nutrient uptake rates. High nutrient uptake rates enhance the competitiveness of primary producers but also increase their nutritional quality for herbivores. This trade-off enables coexistence of nutrient exploiters and grazing avoiders. If herbivores are not selective, evolution favors runaway selection toward high nutrient uptake rates of the primary producers. However, if herbivores select nutritious food, the model predicts an evolutionarily stable strategy with lower nutrient uptake rates. When the model is parameterized for phytoplankton and zooplankton, the evolutionary dynamics result in plant-herbivore oscillations at ecological timescales, especially in environments with high nutrient availability and low selectivity of the herbivores. High herbivore selectivity stabilizes the community dynamics. These model predictions show that evolution permits nonequilibrium dynamics in plant-herbivore communities and shed new light on the evolutionary forces that shape the ecological stoichiometry of primary producers.

Understanding Rapid Evolution in Predator-Prey Interactions Using the Theory of Fast-Slow Dynamical Systems

By Michael H. Cortez and Stephen P. Ellner

The American Naturalist, Volume 176, Issue 5
DOI: 10.1086/656485

The accumulation of evidence that ecologically important traits often evolve at the same time and rate as ecological dynamics (e.g., changes in species' abundances or spatial distributions) has outpaced theory describing the interplay between ecological and evolutionary processes with comparable timescales. The disparity between experiment and theory is partially due to the high dimensionality of models that include both evolutionary and ecological dynamics. Here we show how the theory of fast-slow dynamical systems can be used to reduce model dimension, and we use that body of theory to study a general predator-prey system exhibiting fast evolution in either the predator or the prey. Our approach yields graphical methods with predictive power about when new and unique dynamics (e.g., completely out-of-phase oscillations and cryptic dynamics) can arise in ecological systems exhibiting fast evolution. In addition, we derive analytical expressions for determining when such behavior arises and how evolution affects qualitative properties of the ecological dynamics. Finally, while the theory requires a separation of timescales between the ecological and evolutionary processes, our approach yields insight into systems where the rates of those processes are comparable and thus is a step toward creating a general ecoevolutionary theory.

Variation in Resource Acquisition and Use among Host Clones Creates Key Epidemiological Trade-Offs

By Spencer R. Hall, Claes R. Becker, Meghan A. Duffy, and Carla E. Caceres

The American Naturalist, Volume 176, Issue 5
DOI: 10.1086/656523

Parasites can certainly harm host fitness. Given such virulence, hosts should evolve strategies to resist or tolerate infection. But what governs those strategies and the costs that they incur? This study illustrates how a fecundity-susceptibility trade-off among clonally reared genotypes of a zooplankton (Daphnia dentifera) infected by a fungal parasite (Metschnikowia) arises due to variation in resource acquisition and use by hosts. To make these connections, we used lab experiments and theoretical models that link feeding with susceptibility, energetics, and fecundity of hosts. These feeding-based mechanisms also produced a fecundity-survivorship trade-off. Meanwhile, a parasite spore yield-fecundity trade-off arose from variation in juvenile growth rate among host clones (another index of resource use), a result that was readily anticipated and explained by the models. Thus, several key epidemiological trade-offs stem from variation in resource acquisition and use among clones. This connection should catalyze the creation of new theory that integrates resource- and gene-based responses of hosts to disease.

Recycling-Mediated Facilitation and Coexistence Based on Plant Size

By Antonio J. Golubski, Katherine L. Gross, and Gary G. Mittelbach

The American Naturalist, Volume 176, Issue 5
DOI: 10.1086/656493

We introduce nutrient recycling into a model where competitors differ in the scale at which they perceive their environment. In a two-resource system with both external nutrient inputs and recycling, larger consumers ("integrators") often generate resource distributions that favor their smaller ("nonintegrator") competitors, and vice versa. This occurs because recycling of integrator biomass reduces between-patch resource heterogeneity, whereas recycling of nonintegrator biomass does not. Combined, recycling and throughput can allow coexistence when it is not possible with either alone. With recycling, the presence of an integrator also may facilitate higher biomass of a co-occurring nonintegrator. Our model provides a context where recycling can generate negative feedback between competitors that differ in size and so promote coexistence. This is opposite to the positive recycling-mediated feedback commonly expected on the basis of litter chemistry differences between competitors. Effects of recycling and homogenization on nonintegrators may also be negative in our model, depending on the conformation of the system's resource supply points and the species' relative resource requirements. Our model suggests that the effects of plant size on competitive outcomes may depend critically on the degree of resource recycling found in the system and, reciprocally, that the effects of recycling may depend on plant size.

Tactical Population Movements and Distributions for Ideally Motivated Competitors

By Jonathan T. Rowell

The American Naturalist, Volume 176, Issue 5
DOI: 10.1086/656494

The spatial distributions of populations are a reflection of underlying rules for movement behavior in the context of the environment encountered by individuals. Here I study how ideal directed movement--in which individuals travel in the direction offering the most immediate perceived improvement to their personal fitness--dictates the spatial position of two populations occupying the same relative niche and engaged in competition via interference to an individual's ability to gather resources. Drawing on the analytic derivation of equilibria, numerical simulations, and graphical assessments, I provide conditions under which sympatry, parapatry, or regional exclusion is expected during different phases of the community's development. I also demonstrate that specific competitive asymmetries produce distinguishable distributions and invasion patterns and identify which populations are found centrally or peripherally. Dynamic and dispersal equilibria were examined for differences in the sensitivity to spatial variations in fitness, per capita mortality, metabolic efficiency, the strength of interspecific interference, resource collection speed, and the optimal location of each population along an environmental cline. These asymmetries were studied both in isolation and pairwise in fitness trade-off scenarios.

Ecology of Sexual Dimorphism and Clinal Variation of Coloration in a Damselfly

By Idelle A. Cooper

The American Naturalist, Volume 176, Issue 5
DOI: 10.1086/656491

Sexual selection, more so than natural selection, is posited as the major cause of sex differences. Here I show ecological correlations between solar radiation levels and sexual dimorphism in body color of a Hawaiian damselfly. Megalagrion calliphya exhibits sexual monomorphism at high elevations, where both sexes are red in color; sexual dimorphism at low elevations, where females are green; and female-limited dimorphism at midelevations, where both red and green females exist. Within a midelevation population, red females are also more prevalent during high daily levels of solar radiation. I found that red pigmentation is correlated with superior antioxidant ability that may protect from UV damage and confer a benefit to damselflies in exposed habitats, including males, which defend exposed mating habitats at all elevations, and females, which are in shaded habitats except at high elevation. This study characterizes the ecology of sexual dimorphism and provides a new, ecological hypothesis for the evolution of female-limited dimorphism.