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Showing papers in "The American Naturalist in 2011"


Journal ArticleDOI
TL;DR: It is shown that in squamate ectotherms, two fitness-influencing components of performance, the critical thermal maximum and the thermal optimum, are more closely related to temperature variation and to precipitation, respectively, than they are to mean thermal conditions.
Abstract: Determining organismal responses to climate change is one of biology's greatest challenges. Recent forecasts for future climates emphasize altered temperature variation and precipitation, but most studies of animals have largely focused on forecasting the outcome of changes in mean temperature. Theory suggests that extreme thermal variation and precipitation will influence species performance and hence affect their response to changes in climate. Using an information-theoretic approach, we show that in squamate ectotherms (mostly lizards and snakes), two fitness-influencing components of performance, the critical thermal maximum and the thermal optimum, are more closely related to temperature variation and to precipitation, respectively, than they are to mean thermal conditions. By contrast, critical thermal minimum is related to mean annual temperature. Our results suggest that temperature variation and precipitation regimes have had a strong influence on the evolution of ectotherm performance, so that forecasts for animal responses to climate change will have to incorporate these factors and not only changes in average temperature.

405 citations


Journal ArticleDOI
TL;DR: A unified theoretical framework based on the Price equation is proposed that can be used to model evolution under an expanded inheritance concept that combines the effects of genetic and nongenetic inheritance and can be applied to a variety of scenarios.
Abstract: Inheritance—the influence of ancestors on the pheno- types of their descendants—translates natural selection into evolu- tionary change. For the past century, inheritance has been concep- tualized almost exclusively as the transmission of DNA sequence variation from parents to offspring in accordance with Mendelian rules, but advances in cell and developmental biology have now revealed a rich array of inheritance mechanisms. This empirical evi- dence calls for a unified conception of inheritance that combines genetic and nongenetic mechanisms and encompasses the known range of transgenerational effects, including the transmission of ge- netic and epigenetic variation, the transmission of plastic phenotypes (acquired traits), and the effects of parental environment and ge- notype on offspring phenotype. We propose a unified theoretical framework based on the Price equation that can be used to model evolution under an expanded inheritance concept that combines the effects of genetic and nongenetic inheritance. To illustrate the utility and generality of this framework, we show how it can be applied to a variety of scenarios, including nontransmissible environmental noise, maternal effects, indirect genetic effects, transgenerational epi- genetic inheritance, RNA-mediated inheritance, and cultural

269 citations


Journal ArticleDOI
TL;DR: The best way to fully understand the past is to integrate phylogenies with other types of historical data as well as with direct studies of evolutionary process, because phylogenies are much more informative about pattern than they are about process.
Abstract: The past 30 years have seen a revolution in comparative biology. Before that time, systematics was not at the forefront of the biological sciences, and few scientists considered phylogenetic relationships when investigating evolutionary questions. By contrast, systematic biology is now one of the most vigorous disciplines in biology, and the use of phylogenies not only is requisite in macroevolutionary studies but also has been applied to a wide range of topics and fields that no one could possibly have envisioned 30 years ago. My message is simple: phylogenies are fundamental to comparative biology, but they are not the be-all and end-all. Phylogenies are powerful tools for understanding the past, but like any tool, they have their limitations. In addition, phylogenies are much more informative about pattern than they are about process. The best way to fully understand the past—both pattern and process—is to integrate phylogenies with other types of historical data as well as with direct studies ...

244 citations


Journal ArticleDOI
TL;DR: Overall, predictable behavioral changes occur in response to OSR, although the nature of the change is dependent on the type of mating behavior, and considerable flexibility of mating system structure within species can be predicted by OSR and likely results in variation in the strength of sexual selection.
Abstract: The evolution and maintenance of secondary sexual characteristics and behavior are heavily influenced by the variance in mating success among individuals in a population. The operational sex ratio (OSR) is often used as a predictor of the intensity of competition for mates, as it describes the relative number of males and females who are ready to mate. We investigate changes in aggression, courtship, mate guarding, and sperm release as a function of changes in the OSR using meta-analytic techniques. As the OSR becomes increasingly biased, aggression increases as competitors attempt to defend mates, but this aggression begins to decrease at an OSR of 1.99, presumably due to the increased costs of competition as rivals become more numerous. Sperm release follows a similar but not significant trend. By contrast, courtship rate decreases as the OSR becomes increasingly biased, whereas mate guarding and copulation duration increase. Overall, predictable behavioral changes occur in response to OSR, although the nature of the change is dependent on the type of mating behavior. These results suggest considerable flexibility of mating system structure within species, which can be predicted by OSR and likely results in variation in the strength of sexual selection.

236 citations


Journal ArticleDOI
TL;DR: The results do not support the hypothesis that widely distributed species have larger phenotypic plasticity for thermalolerance limits, and Drosophila species distributions are therefore more closely linked to differences in innate thermal tolerance limits.
Abstract: The distribution of insects can often be related to variation in their response to thermal extremes, which in turn may reflect differences in plastic responses or innate variation in resistance. Species with widespread distributions are expected to have evolved higher levels of plasticity than those from restricted tropical areas. This study compares adult thermal limits across five widespread species and five restricted tropical species of Drosophila from eastern Australia and investigates how these limits are affected by developmental acclimation and hardening after controlling for environmental variation and phylogeny. Irrespective of acclimation, cold resistance was higher in the widespread species. Developmental cold acclimation simulating temperate conditions extended cold limits by 2°–4°C, whereas developmental heat acclimation under simulated tropical conditions increased upper thermal limits by <1°C. The response to adult heat-hardening was weak, whereas widespread species tended to have ...

233 citations


Journal ArticleDOI
TL;DR: The authors' analyses provide little evidence that fitness trade-offs, correlated selection, or stabilizing selection strongly constrains the directional selection reported for most quantitative traits.
Abstract: Studies of phenotypic selection document directional selection in many natural populations. What factors reduce total directional selection and the cumulative evolutionary responses to selection? We combine two data sets for phenotypic selection, representing more than 4,600 distinct estimates of selection from 143 studies, to evaluate the potential roles of fitness trade-offs, indirect (correlated) selection, temporally varying selection, and stabilizing selection for reducing net directional selection and cumulative responses to selection. We detected little evidence that trade-offs among different fitness components reduced total directional selection in most study systems. Comparisons of selection gradients and selection differentials suggest that correlated selection frequently reduced total selection on size but not on other types of traits. The direction of selection on a trait often changes over time in many temporally replicated studies, but these fluctuations have limited impact in reduc...

229 citations


Journal ArticleDOI
TL;DR: The results illustrate that while critical slowing down may be a universal phenomenon at critical transitions, its detection through indirect indicators may have limitations in particular systems.
Abstract: Predicting the risk of critical transitions, such as the collapse of a population, is important in order to direct management efforts. In any system that is close to a critical transition, recovery upon small perturbations becomes slow, a phenomenon known as critical slowing down. It has been suggested that such slowing down may be detected indirectly through an increase in spatial and temporal correlation and variance. Here, we tested this idea in arid ecosystems, where vegetation may collapse to desert as a result of increasing water limitation. We used three models that describe desertification but differ in the spatial vegetation patterns they produce. In all models, recovery rate upon perturbation decreased before vegetation collapsed. However, in one of the models, slowing down failed to translate into rising variance and correlation. This is caused by the regular self-organized vegetation patterns produced by this model. This finding implies an important limitation of variance and correlation as indicators of critical transitions. However, changes in such self-organized patterns themselves are a reliable indicator of an upcoming transition. Our results illustrate that while critical slowing down may be a universal phenomenon at critical transitions, its detection through indirect indicators may have limitations in particular systems.

226 citations


Journal ArticleDOI
TL;DR: Several new results are presented, including robust sufficient conditions to initiate (and stop) spread, using a one-parameter cubic approximation applicable to several models and which has both basic and applied implications.
Abstract: Unlike unconditionally advantageous “Fisherian” variants that tend to spread throughout a species range once introduced anywhere, “bistable” variants, such as chromosome translocations, have two alternative stable frequencies, absence and (near) fixation. Analogous to populations with Allee effects, bistable variants tend to increase locally only once they become sufficiently common, and their spread depends on their rate of increase averaged over all frequencies. Several proposed manipulations of insect populations, such as using Wolbachia or “engineered underdominance” to suppress vector-borne diseases, produce bistable rather than Fisherian dynamics. We synthesize and extend theoretical analyses concerning three features of their spatial behavior: rate of spread, conditions to initiate spread from a localized introduction, and wave stopping caused by variation in population densities or dispersal rates. Unlike Fisherian variants, bistable variants tend to spread spatially only for particular pa...

214 citations


Journal ArticleDOI
TL;DR: A model that scales from the physiological and structural traits of individual trees competing for light and nitrogen across a gradient of soil nitrogen to their community-level consequences predicts that forests are limited by both nitrogen and light, or nearly so.
Abstract: We present a model that scales from the physiological and structural traits of individual trees competing for light and ni- trogen across a gradient of soil nitrogen to their community-level consequences. The model predicts the most competitive (i.e., the evolutionarily stable strategy (ESS)) allocations to foliage, wood, and fine roots for canopy and understory stages of trees growing in old- growth forests. The ESS allocations, revealed as analytical functions of commonly measured physiological parameters, depend not on simple root-shoot relations but rather on diminishing returns of carbon investment that ensure any alternate strategy will underper- form an ESS in monoculture because of the competitive environment that the ESS creates. As such, ESS allocations do not maximize nitrogen-limited growth rates in monoculture, highlighting the un- derappreciated idea that the most competitive strategy is not nec- essarily the "best," but rather that which creates conditions in which all others are "worse." Data from 152 stands support the model's surprising prediction that the dominant structural trade-off is be- tween fine roots and wood, not foliage, suggesting the "root-shoot" trade-off is more precisely a "root-stem" trade-off for long-lived trees. Assuming other resources are abundant, the model predicts that forests are limited by both nitrogen and light, or nearly so.

205 citations


Journal ArticleDOI
TL;DR: Analysis of parental care reaction norms in a bird that exhibits biparental care revealed personality and plasticity are not opposing concepts, and stimulated new hypotheses about sexual conflict over care and provisioning as a life-history trait.
Abstract: Personality (consistent differences between individuals in behavior) and plasticity (changes within individuals in behavior) are often viewed as separate and opposing phenomena. We tested this idea by analyzing parental care reaction norms in a bird that exhibits biparental care. Personality in provisioning behavior existed () and persisted despite being reduced after accounting for individual differences in environment. Plasticity was also evident and differed between the sexes. Male visit rate was associated with changes in brood size and time of day, but female visit rate was associated with changes in nestling age and date. In both sexes changes in visit rate were positively correlated with changes in their partner’s visit rate. Both sexes also exhibited multidimensional reaction norms; interaction terms revealed that within-individual visit rates increased more steeply with brood size when nestlings were older, and the effect of the partner’s visit rate was sensitive to variation in date, pre...

192 citations


Journal ArticleDOI
TL;DR: The results demonstrate that the two dominant mutualisms in terrestrial ecosystems can play major but contrasting roles in plant community assembly and speciation.
Abstract: Both pollination by animals and mycorrhizal symbioses with fungi are believed to have been important for the diversification of flowering plants. However, the mechanisms by which these above- and belowground mutualisms affect plant speciation and coexistence remain obscure. We provide evidence that shifts in pollination traits are important for both speciation and coexistence in a diverse group of orchids, whereas shifts in fungal partner are important for coexistence but not for speciation. Phylogenetic analyses show that recently diverged orchid species tend either to use different pollinator species or to place pollen on different body parts of the same species, consistent with the role of pollination-mode shifts in speciation. Field experiments provide support for the hypothesis that colonization of new geographical areas requires adaptation to new pollinator species, whereas co-occurring orchid species share pollinator species by placing pollen on different body parts. In contrast to pollinat...

Journal ArticleDOI
TL;DR: It is found that herbivore and plant abundances change with temperature in proportion to the ratio of autotrophic to heterotrophic metabolic temperature dependences, and metabolic theory provides a foundation for understanding the effects of temperature change on multitrophic ecological communities.
Abstract: Concern about climate change has spurred experimental tests of how warming affects species’ abundance and performance. As this body of research grows, interpretation and extrapolation to other species and systems have been limited by a lack of theory. To address the need for theory for how warming affects species interactions, we used consumer-prey models and the metabolic theory of ecology to develop quantitative predictions for how systematic differences between the temperature dependence of heterotrophic and autotrophic population growth lead to temperature-dependent herbivory. We found that herbivore and plant abundances change with temperature in proportion to the ratio of autotrophic to heterotrophic metabolic temperature dependences. This result is consistent across five different formulations of consumer-prey models and over varying resource supply rates. Two models predict that temperature-dependent herbivory causes primary producer abundance to be independent of temperature. This finding...

Journal ArticleDOI
TL;DR: It is suggested that nest microclimate can influence nestling phenotype, but whether observed differences carry over to later life-history stages remains unknown.
Abstract: Because the maintenance of proper developmental temperatures during avian incubation is costly to parents, embryos of many species experience pronounced variation in incubation temperature. However, the effects of such temperature variation on nestling development remain relatively unexplored. To investigate this, we artificially incubated wild blue tit (Cyanistes caeruleus L.) clutches at 35.0°, 36.5°, or 38.0°C for two-thirds of the incubation period. We returned clutches to their original nests before hatching and subsequently recorded nestling growth and resting metabolic rate. The length of the incubation period decreased with temperature, whereas hatching success increased. Nestlings from the lowest incubation temperature group had shorter tarsus lengths at 2 weeks of age, but body mass and wing length were not affected by temperature. In addition, nestlings from the lowest temperature group had a significantly higher resting metabolic rate compared with mid- and high-temperature nestlings, ...

Journal ArticleDOI
TL;DR: This work examined patterns of spatial environmental variation within a serpentine mosaic grassland and selection on an annual plant within that landscape to find the first empirical example of differential selection for phenotypic plasticity in the field as a result of strong differences in the grain of environmental heterogeneity within habitats.
Abstract: Adaptive phenotypic plasticity and adaptive genetic differentiation enable plant lineages to maximize their fitness in response to environmental heterogeneity. The spatial scale of environmental variation relative to the average dispersal distance of a species determines whether selection will favor plasticity, local adaptation, or an intermediate strategy. Habitats where the spatial scale of environmental variation is less than the dispersal distance of a species are fine grained and should favor the expression of adaptive plasticity, while coarse-grained habitats, where environmental variation occurs on spatial scales greater than dispersal, should favor adaptive genetic differentiation. However, there is relatively little information available characterizing the link between the spatial scale of environmental variation and patterns of selection on plasticity measured in the field. I examined patterns of spatial environmental variation within a serpentine mosaic grassland and selection on an ann...

Journal ArticleDOI
TL;DR: A simulation model that integrates insights from life-history theory, animal personalities, network theory, and spatial ecology is used to derive a new mechanism for explaining variation in animal invasion success, showing that spread occurs most rapidly when a species includes a mix of life- history or personality types that differ in density-dependent performance and dispersal tendencies.
Abstract: Ecological invasions are a major worldwide problem exacting tremendous economic and ecological costs. Efforts to explain variability in invasion speed and impact by searching for combinations of ecological conditions and species traits associated with invasions have met with mixed success. We use a simulation model that integrates insights from life-history theory, animal personalities, network theory, and spatial ecology to derive a new mechanism for explaining variation in animal invasion success. We show that spread occurs most rapidly when (1) a species includes a mix of life-history or personality types that differ in density-dependent performance and dispersal tendencies, (2) the differences between types are of intermediate magnitude, and (3) patch connections are intermediate in number and widely spread. Within-species polymorphism in phenotype (e.g., life-history strategies or personality), a feature not included in previous models, is important for overcoming the fact that different trai...

Journal ArticleDOI
TL;DR: It is estimated that secondary sympatry takes on the order of millions of years following population splitting and hence could impose an important limit on the rate of range expansion, thereby limiting further rounds of species formation.
Abstract: Range expansions are critical to renewed bouts of allopatric or parapatric speciation. Limits on range expansions—and, by implication, speciation—include dispersal ability and permeability of geographical barriers. In addition, recently diverged taxa may interfere with each other, preventing mutual expansion of each other’s range into sympatry, because reproductive isolation is incomplete and/or ecological competition particularly strong. On the basis of geographical distributions and mitochondrial DNA phylogenetic information for 418 recently diverged species of New World birds, we estimate that secondary sympatry takes on the order of millions of years following population splitting and hence could impose an important limit on the rate of range expansion, thereby limiting further rounds of species formation. Average rates of achievement of sympatry have been faster in the temperate region (we estimate 1.7 million years to sympatry at 60°) than in the tropics (3.2 million years to sympatry at the...

Journal ArticleDOI
TL;DR: An analysis on growth and development rate data compiled from the literature for a well-studied group, marine pelagic copepods, and uses an information-theoretic approach to test which equations best describe these rates.
Abstract: Growth and development rates are fundamental to all living organisms. In a warming world, it is important to determine how these rates will respond to increasing temperatures. It is often assumed that the thermal responses of physiological rates are coupled to metabolic rate and thus have the same temperature dependence. However, the existence of the temperature-size rule suggests that intraspecific growth and development are decoupled. Decoupling of these rates would have important consequences for individual species and ecosystems, yet this has not been tested systematically across a range of species. We conducted an analysis on growth and development rate data compiled from the literature for a well-studied group, marine pelagic copepods, and use an information-theoretic approach to test which equations best describe these rates. Growth and development rates were best characterized by models with significantly different parameters: development has stronger temperature dependence than does growt...

Journal ArticleDOI
TL;DR: The main discernible pattern of mate choice was assortative mating and it was found that partner similarity was due to initial choice rather than convergence and also at least in part to phenotypic matching.
Abstract: Human mate choice is central to individuals’ lives and to the evolution of the species, but the basis of variation in mate choice is not well understood. Here we looked at a large community-based sample of twins and their partners and parents ( individuals) to test for genetic and family environmental influences on mate choice, while controlling for and not controlling for the effects of assortative mating. Key traits were analyzed, including height, body mass index, age, education, income, personality, social attitudes, and religiosity. This revealed near-zero genetic influences on male and female mate choice over all traits and no significant genetic influences on mate choice for any specific trait. A significant family environmental influence was found for the age and income of females’ mate choices, possibly reflecting parental influence over mating decisions. We also tested for evidence of sexual imprinting, where individuals acquire mate-choice criteria during development by using their oppo...

Journal ArticleDOI
TL;DR: A species border likely set by limited adaptation to declining environmental quality is revealed, with distribution models employing demographically important environmental variables predicted low habitat favorability beyond the eastern border.
Abstract: Potential causes of species’ geographic distribution limits fall into two broad classes: (1) limited adaptation across spatially variable environments and (2) limited opportunities to colonize unoccupied areas. Combining demographic studies, analyses of demographic responses to environmental variation, and species distribution models, we investigated the causes of range limits in a model system, the eastern border of the California annual plant Clarkia xantiana ssp. xantiana. Vital rates of 20 populations varied with growing season temperature and precipitation: fruit number and overwinter survival of 1-year-old seeds declined steeply, while current-year seed germination increased modestly along west-to-east gradients in decreasing temperature, decreasing mean precipitation, and increasing variation in precipitation. Long-term stochastic finite rate of increase, λs, exhibited a fourfold range and varied among geologic surface materials as well as with temperature and precipitation. Growth rate dec...

Journal ArticleDOI
TL;DR: This work develops a framework for neighbor-dependent selection that predicts the conditions under which selection can enable coexistence, as opposed to merely augmenting it, and elucidates the effects of heritability on the eco-evolutionary feedbacks that drive coexistence.
Abstract: Recent studies suggest that selection can allow coexis- tence in situations where ecological dynamics lead to competitive exclusion, provided that there is a trade-off between traits optimal for interacting with conspecifics and traits optimal for interacting with heterospecifics. Despite compelling empirical evidence, there is no general framework for elucidating how and when selection will allow coexistence in natural communities. Here we develop such a framework for a mechanism that we term "neighbor-dependent se- lection." We show that this mechanism can both augment coexistence when ecological conditions allow for niche partitioning and enable coexistence when ecological conditions lead to competitive exclusion. The novel insight is that when ecological conditions lead to exclusion, neighbor-dependent selection can allow coexistence via cycles driven by an intransitive loop; selection causes one species to be a superior interspecific competitor when it is rare and an inferior interspecific competitor when it is abundant. Our framework predicts the con- ditions under which selection can enable coexistence, as opposed to merely augmenting it, and elucidates the effects of heritability on the eco-evolutionary feedbacks that drive coexistence. Given increas- ing evidence that evolution operates on ecological timescales, our approach provides one means for evaluating the role of selection and trait evolution in species coexistence.

Journal ArticleDOI
TL;DR: It is demonstrated that sexual conflict can drive a population to extinction but also highlighted how simple mechanisms, such as harassment costs to males and females and the coevolution between harassment and resistance, can help avert a tragedy of the commons caused by sexual conflict.
Abstract: It is widely understood that the costs and benefits of mating can affect the fecundity and survival of individuals. Sexual conflict may have profound consequences for populations as a result of the negative effects it causes males and females to have on one another’s fitness. Here we present a model describing the evolution of sexual conflict, in which males inflict a direct cost on female fitness. We show that these costs can drive the entire population to extinction. To males, females are an essential but finite resource over which they have to compete. Population extinction owing to sexual conflict can therefore be seen as an evolutionary tragedy of the commons. Our model shows that a positive feedback between harassment and the operational sex ratio is responsible for the demise of females and, thus, for population extinction. We further show that the evolution of female resistance to counter harassment can prevent a tragedy of the commons. Our findings not only demonstrate that sexual conflic...

Journal ArticleDOI
TL;DR: This work derives a new expression for the threshold frequency that extends existing theory to incorporate important details of the insect’s life history and shows how the type of immigration and the temporal dynamics of the host population can strongly affect the spread of Wolbachia.
Abstract: Wolbachia are endosymbionts that are found in many insect species and can spread rapidly when introduced into a naive host population. Most Wolbachia spread when their infection frequency exceeds a threshold normally calculated using purely population genetic models. However, spread may also depend on the population dynamics of the insect host. We develop models to explore interactions between host population dynamics and Wolbachia infection frequency for an age-structured insect population regulated by larval density dependence. We first derive a new expression for the threshold frequency that extends existing theory to incorporate important details of the insect's life history. In the presence of immigration and emigration, the threshold also depends on the form of density-dependent regulation. We show how the type of immigration (constant or pulsed) and the temporal dynamics of the host population can strongly affect the spread of Wolbachia. The results help understand the natural dynamics of Wolbachia infections and aid the design of programs to introduce Wolbachia to control insects that are disease vectors or pests.

Journal ArticleDOI
TL;DR: It is found that an evolutionary response to thermal selection was capable of explaining up to two-thirds of the phenotypic trend, and this method augments the toolbox for predicting responses to environmental change.
Abstract: Environmental change can shift the phenotype of an organism through either evolutionary or nongenetic processes. Despite abundant evidence of phenotypic change in response to recent climate change, we typically lack sufficient genetic data to identify the role of evolution. We present a method of using phenotypic data to characterize the hypothesized role of natural selection and environmentally driven phenotypic shifts (plasticity). We modeled historical selection and environmental predictors of interannual variation in mean population phenotype using a multivariate state-space model framework. Through model comparisons, we assessed the extent to which an estimated selection differential explained observed variation better than environmental factors alone. We applied the method to a 60-year trend toward earlier migration in Columbia River sockeye salmon Oncorhynchus nerka, producing estimates of annual selection differentials, average realized heritability, and relative cumulative effects of sele...

Journal ArticleDOI
TL;DR: The results suggest that phages target common bacterial species, including an important plant pathogen, within plant host tissues; this result has important implications for therapeutic phage epidemiology and highlights that biotic environment can play a key role in shaping the spatial scale of parasite adaptation and influencing the outcome of coevolutionary interactions.
Abstract: The ecological, epidemiological, and evolutionary consequences of host-parasite interactions are critically shaped by the spatial scale at which parasites adapt to hosts. The scale of interaction between hyperparasites and their parasites is likely to be influenced by the host of the parasite and potentially likely to differ among within-host environments. Here we examine the scale at which bacteriophages adapt to their host bacteria by studying natural isolates from the surface or interior of horse chestnut leaves. We find that phages are more infective to bacteria from the same tree relative to those from other trees but do not differ in infectivity to bacteria from different leaves within the same tree. The results suggest that phages target common bacterial species, including an important plant pathogen, within plant host tissues; this result has important implications for therapeutic phage epidemiology. Furthermore, we show that phages from the leaf interior are more infective to their local ...

Journal ArticleDOI
TL;DR: In conclusion, repellency of 2PE is highly sensitive to dosage, giving it a key role in shaping ecological interactions between skypilot plants and their floral visitors, at no pollination cost.
Abstract: All volatile organic compounds (VOCs) vary quantitatively, yet how such variation affects their ecological roles is unknown. Because floral VOCs are cues for both pollinators and floral antagonists, variation in emission may have major consequences for costs and benefits in plant-pollinator interactions. In Polemonium viscosum, the emission rate for the floral VOC 2-phenylethanol (2PE) spans more than two orders of magnitude. We investigated the ecological and evolutionary impacts of this immense phenotypic variation. The emission rate of 2PE varies independently of nectar rewards and thus is uninformative of profitability. Emission is elevated in flowers that are morphologically vulnerable to ant larcenists, suggesting that chemical deterrence may compensate for weak physical barriers. In nature, plants emitting more 2PE than their neighbors escape ant damage. Flower-damaging ants die when exposed to 2PE in the laboratory, and they avoid high 2PE emitters in the field. High 2PE also reduces bumblebee visitation and pollination, suggesting an ecological cost of defense in pollinator service. However, at more moderate emission rates, 2PE enhances the amount of nectar left in flowers, at no pollination cost. In conclusion, repellency of 2PE is highly sensitive to dosage, giving it a key role in shaping ecological interactions between skypilot plants and their floral visitors.

Journal ArticleDOI
TL;DR: Support is found for the hypothesis that plant size and resource heterogeneity interact to determine species diversity, which figures prominently in resource-ratio models of competition.
Abstract: Spatial heterogeneity in soil resources is widely thought to promote plant species coexistence, and this mechanism figures prominently in resource-ratio models of competition. However, most experimental studies have found that nutrient enhancements depress diversity regardless of whether nutrients are uniformly or heterogeneously applied. This mismatch between theory and empirical pattern is potentially due to an interaction between plant size and the scale of resource heterogeneity. Clonal plants that spread vegetatively via rhizomes or stolons can grow large and may integrate across resource patches, thus reducing the positive effect of small-scale resource heterogeneity on plant species richness. Many rhizomatous clonal species respond strongly to increased soil fertility, and they have been hypothesized to drive the descending arm of the hump-shaped productivity-diversity relationship in grasslands. We tested whether clonals reduce species richness in a grassland community by manipulating nutr...

Journal ArticleDOI
TL;DR: The classic cost-benefit model of ectothermic thermoregulation is extended to include the case where Te exceeds Tsel, and costs and benefits are defined in terms of fitness to include effects of body temperature (Tb) on performance and survival.
Abstract: The classic cost-benefit model of ectothermic thermoregulation compares energetic costs and benefits, providing a critical framework for understanding this process (Huey and Slatkin 1976). It considers the case where environmental temperature (Te) is less than the selected temperature of the organism (Tsel), and it predicts that, to minimize increasing energetic costs of thermoregulation as habitat thermal quality declines, thermoregulatory effort should decrease until the lizard thermoconforms. We extended this model to include the case where Te exceeds Tsel, and we redefine costs and benefits in terms of fitness to include effects of body temperature (Tb) on performance and survival. Our extended model predicts that lizards will increase thermoregulatory effort as habitat thermal quality declines, gaining the fitness benefits of optimal Tb and maximizing the net benefit of activity. Further, to offset the disproportionately high fitness costs of high Te compared with low Te, we predicted that li...

Journal ArticleDOI
TL;DR: These correlated patterns at the species and haplotype level are consistent with individual-based stochastic dispersal proposed by neutral theories of biodiversity and demonstrate the utility of haplotype data for exploring macroecological patterns in poorly known biota and predicting large-scale biodiversity patterns based on genetic inventories of local samples.
Abstract: A positive correlation between species diversity and genetic diversity has been proposed, consistent with neutral predictions in macroecology. We assessed the species–genetic diversity correlation in tenebrionid beetle communities of the Aegean archipelago on 15 islands of different sizes, distances to mainland, and ages of isolation. Alpha and beta diversity of species and haplotypes were assessed using sequences of >1,000 individuals (mitochondrial cytochrome oxidase 1 and nuclear muscular protein 20). We show that (i) there is a strong species-area and haplotype-area relationship; (ii) species richness in island communities is correlated with intraspecific genetic diversity in the constituent species except when island size or distance to mainland is factored out in partial correlations; (iii) community similarity declines exponentially at an increasing rate when calculated on the basis of species, nuclear, and mtDNA haplotypes; and (iv) distance decay of community similarity is slower in dispe...

Journal ArticleDOI
TL;DR: Different host and parasite genotypes would be selected under different climatic conditions, affecting the coevolutionary dynamics of the host‐parasite interaction and the course of chestnut blight epidemics.
Abstract: The outcome of host‐parasite interactions may depend not only on the genotypes of the species involved but also on environmental factors. We used the fungus Cryphonectria parasitica, the causal agent of chestnut blight, and its hyperparasitic virus, Cryphonectria hypovirus‐1 (CHV1), to test for genotype‐by‐genotype‐by‐environment interactions in a host‐parasite system. In C. parasitica, infection with CHV1 induces a hypovirulent phenotype with reduced virulence toward the chestnut tree (Castanea spp.) and thus controls chestnut blight in many European regions. In contrast, uninfected virulent C. parasitica have nearly eradicated the American chestnut in North America. We applied a full factorial design and assessed the fungal growth and sporulation of four C. parasitica strains, uninfected and infected with each of the four known CHV1 subtypes, at 12°, 18°, 24°, and 30°C. We found a significant ( \documentclass{aastex} \usepackage{amsbsy} \usepackage{amsfonts} \usepackage{amssymb} \usepackage{bm...

Journal ArticleDOI
TL;DR: It is found that environmental shifts that promote tolerance ultimately result in populations harboring more parasitized individuals, and that the opportunity for selection, as indicated by the variance around traits, varied considerably with the environmental treatment.
Abstract: Victims of infection are expected to suffer increasingly as parasite population growth increases. Yet, under some conditions, faster-growing parasites do not appear to cause more damage, and infections can be quite tolerable. We studied these conditions by assessing how the relationship between parasite population growth and host health is sensitive to environmental variation. In experimental infections of the crustacean Daphnia magna and its bacterial parasite Pasteuria ramosa, we show how easily an interaction can shift from a severe interaction, that is, when host fitness declines substantially with each unit of parasite growth, to a tolerable relationship by changing only simple environmental variables: temperature and food availability. We explored the evolutionary and epidemiological implications of such a shift by modeling pathogen evolution and disease spread under different levels of infection severity and found that environmental shifts that promote tolerance ultimately result in populat...