scispace - formally typeset
Search or ask a question

Showing papers in "The American Naturalist in 2017"


Journal ArticleDOI
TL;DR: It is argued that speciation research, both empirical and theoretical, needs to consider both the origin of barrier effects and the ways in which they are coupled, and for an extended view of reinforcement that includes coupling processes involving enhancement of any type of additional barrier effect as a result of an existing barrier.
Abstract: During the process of speciation, populations may diverge for traits and at their underlying loci that contribute barriers to gene flow. These barrier traits and barrier loci underlie individual barrier effects, by which we mean the contribution that a barrier locus or trait—or some combination of barrier loci or traits—makes to overall isolation. The evolution of strong reproductive isolation typically requires the origin of multiple barrier effects. Critically, it also requires the coincidence of barrier effects; for example, two barrier effects, one due to assortative mating and the other due to hybrid inviability, create a stronger overall barrier to gene flow if they coincide than if they distinguish independent pairs of populations. Here, we define “coupling” as any process that generates coincidence of barrier effects, resulting in a stronger overall barrier to gene flow. We argue that speciation research, both empirical and theoretical, needs to consider both the origin of barrier effects ...

151 citations


Journal ArticleDOI
TL;DR: Both theoretical and empirical studies of adaptation to rapid environmental change are reviewed, focusing on areas where genomic data are poised to contribute to the authors' ability to estimate species and population persistence and adaptation.
Abstract: Rapid environmental change currently presents a major threat to global biodiversity and ecosystem functions, and understanding impacts on individual populations is critical to creating reliable predictions and mitigation plans. One emerging tool for this goal is high-throughput sequencing technology, which can now be used to scan the genome for signs of environmental selection in any species and any system. This explosion of data provides a powerful new window into the molecular mechanisms of adaptation, and although there has been some success in using genomic data to predict responses to selection in fields such as agriculture, thus far genomic data are rarely integrated into predictive frameworks of future adaptation in natural populations. Here, we review both theoretical and empirical studies of adaptation to rapid environmental change, focusing on areas where genomic data are poised to contribute to our ability to estimate species and population persistence and adaptation. We advocate for the need to study and model evolutionary response architectures, which integrate spatial information, fitness estimates, and plasticity with genetic architecture. Understanding how these factors contribute to adaptive responses is essential in efforts to predict the responses of species and ecosystems to future environmental change.

124 citations


Journal ArticleDOI
TL;DR: A novel phylogenetic comparative method is developed to investigate how often and in which clades the macroevolutionary dynamics of the metabolic scaling have changed and whether traits such as ecto-/endothermy, genome size, and quadratic curvature with body mass could explain the observed pattern of shifts.
Abstract: Metabolism is the link between ecology and physiology—it dictates the flow of energy through individuals and across trophic levels. Much of the predictive power of metabolic theories of ecology derives from the scaling relationship between organismal size and metabolic rate. There is growing evidence that this scaling relationship is not universal, but we have little knowledge of how it has evolved over macroevolutionary time. Here we develop a novel phylogenetic comparative method to investigate how often and in which clades the macroevolutionary dynamics of the metabolic scaling have changed. We find strong evidence that the metabolic scaling relationship has shifted multiple times across the vertebrate phylogeny. However, shifts are rare and otherwise strongly constrained. Importantly, both the estimated slope and intercept values vary widely across regimes, with slopes that spanned across theoretically predicted values such as 2/3 or 3/4. We further tested whether traits such as ecto-/endother...

101 citations


Journal ArticleDOI
TL;DR: A comparative analysis of studies of fishes shows high variation in the extent of parallelism—from very high to very low, along with everything in between—highlighting the importance of quantifying parallelism rather than just asserting its presence.
Abstract: Evidence of phenotypic parallelism is often used to infer the deterministic role played by natural selection. However, variation in the extent or direction of divergence is often evident among independent evolutionary replicates, raising the following question: just how parallel, overall, is parallel evolution? We answer this question through a comparative analysis of studies of fishes, a taxon where parallel evolution has been much discussed. We first ask how much of the among-population variance in phenotypic traits can be explained by different "environment" types, such as high predation versus low predation or benthic versus limnetic. We then use phenotypic change vector analysis to quantify variation in the direction (vector angles) and magnitude (vector lengths) of environment-associated divergence. All analyses show high variation in the extent of parallelism-from very high to very low, along with everything in between-highlighting the importance of quantifying parallelism rather than just asserting its presence. Interestingly, instances of low extents of parallelism represent important components of divergence in many cases, promising considerable opportunities for inferences about the factors shaping phenotypic divergence.

97 citations


Journal ArticleDOI
TL;DR: It is argued that many of the most dramatic biotic patterns, past and present, are likely to have been generated by diverse, mutually reinforcing drivers.
Abstract: An impediment to understanding the origin and dynamics of the latitudinal diversity gradient (LDG)-the most pervasive large-scale biotic pattern on Earth-has been the tendency to focus narrowly on a single causal factor when a more synthetic, integrative approach is needed. Using marine bivalves as a model system and drawing on other systems where possible, we review paleobiologic and biogeographic support for two supposedly opposing views, that the LDG is shaped primarily by (a) local environmental factors that determine the number of species and higher taxa at a given latitude (in situ hypotheses) or (b) the entry of lineages arising elsewhere into a focal region (spatial dynamics hypotheses). Support for in situ hypotheses includes the fit of present-day diversity trends in many clades to such environmental factors as temperature and the correlation of extinction intensities in Pliocene bivalve faunas with net regional temperature changes. Support for spatial dynamics hypotheses includes the age-frequency distribution of bivalve genera across latitudes, which is consistent with an out-of-the-tropics dynamic, as are the higher species diversities in temperate southeastern Australia and southeastern Japan than in the tropical Caribbean. Thus, both in situ and spatial dynamics processes must shape the bivalve LDG and are likely to operate in other groups as well. The relative strengths of the two processes may differ among groups showing similar LDGs, but dissecting their effects will require improved methods of integrating fossil data with molecular phylogenies. We highlight several potential research directions and argue that many of the most dramatic biotic patterns, past and present, are likely to have been generated by diverse, mutually reinforcing drivers.

94 citations


Journal ArticleDOI
TL;DR: It is suggested that the most fundamental theorem of evolution is the Price equation, both because of its simplicity and broad scope and because it can be used to derive four other familiar results that are similarly fundamental: Fisher’s average-excess equation, Robertson's secondary theorem of natural selection, the breeder's equation, and Fisher's fundamental theorem.
Abstract: Evolutionary biology is undergirded by an extensive and impressive set of mathematical models. Yet only one result, Fisher's theorem about selection and fitness, is generally accorded the status of a fundamental theorem. I argue that although its fundamental status is justified by its simplicity and scope, there are additional results that seem similarly fundamental. I suggest that the most fundamental theorem of evolution is the Price equation, both because of its simplicity and broad scope and because it can be used to derive four other familiar results that are similarly fundamental: Fisher's average-excess equation, Robertson's secondary theorem of natural selection, the breeder's equation, and Fisher's fundamental theorem. These derivations clarify both the relationships behind these results and their assumptions. Slightly less fundamental results include those for multivariate evolution and social selection. A key feature of fundamental theorems is that they have great simplicity and scope, which are often achieved by sacrificing perfect accuracy. Quantitative genetics has been more productive of fundamental theorems than population genetics, probably because its empirical focus on unknown genotypes freed it from the tyranny of detail and allowed it to focus on general issues.

90 citations


Journal ArticleDOI
TL;DR: Results from spatially explicit simulations indicate that the presence of an environmental gradient during range expansion reduces expansion load; conversely, increasing expansion load allows only locally adapted populations to persist at the range edge.
Abstract: The biotic and abiotic factors that facilitate or hinder species range expansions are many and complex. We examine the impact of two genetic processes and their interaction on fitness at expanding range edges: local maladaptation resulting from the presence of an environmental gradient and expansion load resulting from increased genetic drift at the range edge. Results from spatially explicit simulations indicate that the presence of an environmental gradient during range expansion reduces expansion load; conversely, increasing expansion load allows only locally adapted populations to persist at the range edge. Increased maladaptation reduces the speed of range expansion, resulting in less genetic drift at the expanding front and more immigration from the range center, therefore reducing expansion load at the range edge. These results may have ramifications for species being forced to shift their ranges because of climate change or other anthropogenic changes. If rapidly changing climate leads to ...

88 citations


Journal ArticleDOI
TL;DR: It is shown that colonization times are important for shaping elevational richness patterns within the Andes, with more species found in habitats colonized earlier, and it is suggested that these scale-dependent patterns might apply to many other richness gradients.
Abstract: The Tropical Andes make up Earth’s most species-rich biodiversity hotspot for both animals and plants. Nevertheless, the ecological and evolutionary processes underlying this extraordinary richness remain uncertain. Here, we examine the processes that generate high richness in the Tropical Andes relative to other regions in South America and across different elevations within the Andes, using frogs as a model system. We combine distributional data, a newly generated time-calibrated phylogeny for 2,318 frog species, and phylogenetic comparative methods to test the relative importance of diversification rates and colonization times for explaining Andean diversity at different scales. At larger scales (among regions and families), we find that faster diversification rates in Andean clades most likely explain high Andean richness. In contrast, at smaller temporal and spatial scales (within family-level clades within the Andes), diversification rates rarely explain richness patterns. Instead, we show t...

85 citations


Journal ArticleDOI
TL;DR: This work assessed which solutions evolved depending on degree of temporal variation, costs of multiple underlying traits, and order of dispersal and development to explain the apparent relative rarity of matching habitat choice in nature.
Abstract: It can be challenging for organisms to achieve a good match between their phenotypic characteristics and environmental requirements that vary in space and time. The evolution of adaptive phenotypes can result from genetic differentiation at the population level. Individuals, however, could also change their phenotype (adaptive plasticity) or select an environment because it matches with their phenotype (matching habitat choice). It is poorly known under which conditions these different solutions to environmental heterogeneity evolve and whether they operate together. Using an individual-based simulation model, we assessed which solutions evolved depending on degree of temporal variation, costs of multiple underlying traits, and order of dispersal and development. Population genetic divergence was superseded by plasticity or matching habitat choice as temporal variation increased. Plasticity and matching habitat choice were limited by their trait costs, even when this involved only a part of the un...

79 citations


Journal ArticleDOI
TL;DR: The results reveal that species ecology in the form of habitat association is an important predictor of genetic diversity and population divergence and suggest that differences in diversity between floodplain and upland avifaunas in the Amazon may be driven by differences in the demographic and evolutionary processes at work in the two habitats.
Abstract: The ecological traits of organisms may predict their genetic diversity and population genetic structure and mediate the action of evolutionary processes important for speciation and adaptation. Making these ecological-evolutionary links is difficult because it requires comparable genetic estimates from many species with differing ecologies. In Amazonian birds, habitat association is an important component of ecological diversity. Here, we examine the link between habitat association and genetic parameters using 20 pairs of closely related Amazonian bird species in which one member of the pair occurs primarily in forest edge and floodplains and the other occurs in upland forest interior. We use standardized geographic sampling and data from 2,416 genomic markers to estimate genetic diversity, population genetic structure, and statistics reflecting demographic and evolutionary processes. We find that species of upland forest have greater genetic diversity and divergence across the landscape as well as signatures of older histories and less gene flow than floodplain species. Our results reveal that species ecology in the form of habitat association is an important predictor of genetic diversity and population divergence and suggest that differences in diversity between floodplain and upland avifaunas in the Amazon may be driven by differences in the demographic and evolutionary processes at work in the two habitats.

78 citations


Journal ArticleDOI
TL;DR: Perhaps the most important emerging conclusion is that the genetic mechanisms of convergent evolution are phylogenetically conserved; that is, more closely related species tend to share the same genetic basis of traits, even when independently evolved.
Abstract: A charm of biology as a scientific discipline is the diversity of life. Although this diversity can make laws of biology challenging to discover, several repeated patterns and general principles govern evolutionary diversification. Convergent evolution, the independent evolution of similar phenotypes, has been at the heart of one approach to understand generality in the evolutionary process. Yet understanding when and why organismal traits and strategies repeatedly evolve has been a central challenge. These issues were the focus of the American Society of Naturalists Vice Presidential Symposium in 2016 and are the subject of this collection of articles. Although naturalists have long made inferences about convergent evolution and its importance, there has been confusion in the interpretation of the pattern of convergence. Does convergence primarily indicate adaptation or constraint? How often should convergence be expected? Are there general principles that would allow us to predict where and when and by what mechanisms convergent evolution should occur? What role does natural history play in advancing our understanding of general evolutionary principles? In this introductory article, I address these questions, review several generalizations about convergent evolution that have emerged over the past 15 years, and present a framework for advancing the study and interpretation of convergence. Perhaps the most important emerging conclusion is that the genetic mechanisms of convergent evolution are phylogenetically conserved; that is, more closely related species tend to share the same genetic basis of traits, even when independently evolved. Finally, I highlight how the articles in this special issue further develop concepts, methodologies, and case studies at the frontier of our understanding of the causes and consequences of convergent evolution.

Journal ArticleDOI
TL;DR: This article developed an approach that links quantitative genetic theory with data-driven structured models to allow prediction of population responses to environmental change via plasticity and adaptive evolution, and constructed a number of example models to demonstrate that evolutionary responses to short-term environmental change over the shortterm will be considerably slower than plastic responses and that the rate of adaptive evolution to a new environment depends on whether plastic responses are adaptive or nonadaptive.
Abstract: Understanding how the natural world will be impacted by environmental change over the coming decades is one of the most pressing challenges facing humanity. Addressing this challenge is difficult because environmental change can generate both population-level plastic and evolutionary responses, with plastic responses being either adaptive or nonadaptive. We develop an approach that links quantitative genetic theory with data-driven structured models to allow prediction of population responses to environmental change via plasticity and adaptive evolution. After introducing general new theory, we construct a number of example models to demonstrate that evolutionary responses to environmental change over the short-term will be considerably slower than plastic responses and that the rate of adaptive evolution to a new environment depends on whether plastic responses are adaptive or nonadaptive. Parameterization of the models we develop requires information on genetic and phenotypic variation and demography that will not always be available, meaning that simpler models will often be required to predict responses to environmental change. We consequently develop a method to examine whether the full machinery of the evolutionarily explicit models we develop will be needed to predict responses to environmental change or whether simpler nonevolutionary models that are now widely constructed may be sufficient.

Journal ArticleDOI
TL;DR: In this paper, the authors used multivariate statistical approaches to develop the beginnings for a periodic table of niches for lizards to examine patterns of global and regional niche diversification.
Abstract: Widespread niche convergence suggests that species can be organized according to functional trait combinations to create a framework analogous to a periodic table. We compiled ecological data for lizards to examine patterns of global and regional niche diversification, and we used multivariate statistical approaches to develop the beginnings for a periodic table of niches. Data (50+ variables) for five major niche dimensions (habitat, diet, life history, metabolism, defense) were compiled for 134 species of lizards representing 24 of the 38 extant families. Principal coordinates analyses were performed on niche dimensional data sets, and species scores for the first three axes were used as input for a principal components analysis to ordinate species in continuous niche space and for a regression tree analysis to separate species into discrete niche categories. Three-dimensional models facilitate exploration of species positions in relation to major gradients within the niche hypervolume. The firs...

Journal ArticleDOI
TL;DR: It is found that selection varied more among treatments in experimental studies than among spatial and temporal replicates in observational Studies, suggesting that experimental studies can detect relationships between environmental factors and selection that would not be apparent in observational studies.
Abstract: Although many selection estimates have been published, the environmental factors that cause selection to vary in space and time have rarely been identified. One way to identify these factors is by experimentally manipulating the environment and measuring selection in each treatment. We compiled and analyzed selection estimates from experimental studies. First, we tested whether the effect of manipulating the environment on selection gradients depends on taxon, trait type, or fitness component. We found that the effect of manipulating the environment was larger when selection was measured on life-history traits or via survival. Second, we tested two predictions about the environmental factors that cause variation in selection. We found support for the prediction that variation in selection is more likely to be caused by environmental factors that have a large effect on mean fitness but not for the prediction that variation is more likely to be caused by biotic factors. Third, we compared selection gradients from experimental and observational studies. We found that selection varied more among treatments in experimental studies than among spatial and temporal replicates in observational studies, suggesting that experimental studies can detect relationships between environmental factors and selection that would not be apparent in observational studies.

Journal ArticleDOI
TL;DR: It is shown that microhabitat can have a much stronger influence on diversification than Climatic niche position or rates of climatic niche change, and it is found that the best-fitting phylogenetic multiple regression model includes all three types of variables.
Abstract: A major goal of ecology and evolutionary biology is to explain patterns of species richness among clades. Differences in rates of net diversification (speciation minus extinction over time) may often explain these patterns, but the factors that drive variation in diversification rates remain uncertain. Three important candidates are climatic niche position (e.g., whether clades are primarily temperate or tropical), rates of climatic niche change among species within clades, and microhabitat (e.g., aquatic, terrestrial, arboreal). The first two factors have been tested separately in several studies, but the relative importance of all three is largely unknown. Here we explore the correlates of diversification among families of frogs, which collectively represent ∼88% of amphibian species. We assemble and analyze data on phylogeny, climate, and microhabitat for thousands of species. We find that the best-fitting phylogenetic multiple regression model includes all three types of variables: microhabitat, rates of climatic niche change, and climatic niche position. This model explains 67% of the variation in diversification rates among frog families, with arboreal microhabitat explaining ∼31%, niche rates ∼25%, and climatic niche position ∼11%. Surprisingly, we show that microhabitat can have a much stronger influence on diversification than climatic niche position or rates of climatic niche change.

Journal ArticleDOI
TL;DR: A hybrid, eco-evolutionary model is proposed to explain the remarkable pattern of global host resistance and local parasite infectivity and is not wholly compatible with either of the common models of host-parasite coevolution (i.e., matching infection or targeted recognition).
Abstract: Parasite infections are a product of both ecological processes affecting host-parasite encounter rates and evolutionary dynamics affecting host susceptibility. However, few studies examine natural infection variation from both ecological and evolutionary perspectives. Here, we describe the ecological and evolutionary factors generating variation in infection rates by a tapeworm (Schistocephalus solidus) in a vertebrate host, the threespine stickleback (Gasterosteus aculeatus). To explore ecological aspects of infection, we measured tapeworm prevalence in Canadian stickleback inhabiting two distinct environments: marine and freshwater. Consistent with ecological control of infection, the tapeworm is very rare in marine environments, even though marine fish are highly susceptible. Conversely, commonly infected freshwater stickleback exhibit substantial resistance in controlled laboratory trials, suggesting that high exposure risk overwhelms their recently evolved resistance. We also tested for parasite adaptation to its host by performing transcontinental reciprocal infections, using stickleback and tapeworm populations from Europe and western Canada. More infections occurred in same-continent host-parasite combinations, indicating parasite "local" adaptation, at least on the scale of continents. However, the recently evolved immunity of freshwater hosts applies to both local and foreign parasites. The pattern of adaptation described here is not wholly compatible with either of the common models of host-parasite coevolution (i.e., matching infection or targeted recognition). Instead, we propose a hybrid, eco-evolutionary model to explain the remarkable pattern of global host resistance and local parasite infectivity.

Journal ArticleDOI
TL;DR: This work uses a continuous time/continuous space model in which it varies both random (diffusive) movement and resource-following (advective) movement to characterize the optimal perceptual ranges for foragers in dynamic landscapes.
Abstract: How organisms gather and utilize information about their landscapes is central to understanding land-use patterns and population distributions. When such information originates beyond an individual's immediate vicinity, movement decisions require integrating information out to some perceptual range. Such nonlocal information, whether obtained visually, acoustically, or via chemosensation, provides a field of stimuli that guides movement. Classically, however, models have assumed movement based on purely local information (e.g., chemotaxis, step-selection functions). Here we explore how foragers can exploit nonlocal information to improve their success in dynamic landscapes. Using a continuous time/continuous space model in which we vary both random (diffusive) movement and resource-following (advective) movement, we characterize the optimal perceptual ranges for foragers in dynamic landscapes. Nonlocal information can be highly beneficial, increasing the spatiotemporal concentration of foragers on their resources up to twofold compared with movement based on purely local information. However, nonlocal information is most useful when foragers possess both high advective movement (allowing them to react to transient resources) and low diffusive movement (preventing them from drifting away from resource peaks). Nonlocal information is particularly beneficial in landscapes with sharp (rather than gradual) patch edges and in landscapes with highly transient resources.

Journal ArticleDOI
TL;DR: The results support theory suggesting that fitness trade-offs across life stages may contribute to the maintenance of personality variation within populations, and demonstrate that boldness can affect fitness differently acrosslife stages.
Abstract: To date, most studies investigating the relationship between personality traits and fitness have focused on a single measure of fitness (such as survival) at a specific life stage. However, many personality traits likely have multiple effects on fitness, potentially operating across different functional contexts and stages of development. Here, we address the fitness consequences of boldness, under seminatural conditions, across life stages and functional contexts in largemouth bass (Micropterus salmoides). Specifically, we report the effect of boldness on (1) juvenile survivorship in an outdoor pond containing natural prey and predators and (2) adult reproductive success in three outdoor ponds across three reproductive seasons (years). Juvenile survival was negatively affected by boldness, with bolder juveniles having a lower probability of survival than shyer juveniles. In contrast, bolder adult male bass had greater reproductive success than their shyer male counterparts. Female reproductive success was not affected by boldness. These findings demonstrate that boldness can affect fitness differently across life stages. Further, boldness was highly consistent across years and significantly heritable, which suggests that boldness has a genetic component. Thus, our results support theory suggesting that fitness trade-offs across life stages may contribute to the maintenance of personality variation within populations.

Journal ArticleDOI
TL;DR: It is shown that the use of social information alone increases grouping behavior but constrains group size to limit competition, ultimately increasing individual fitness substantially across various ecological contexts, supporting recent theory on competition-information quality trade-offs.
Abstract: Empirical studies in select systems suggest that social information—the incidental or deliberate information produced by animals and available to other animals—can fundamentally shape animal grouping behavior. However, to understand the role of social information in animal behavior and fitness, we must establish general theory that quantifies effects of social information across ecological contexts and generates expectations that can be applied across systems. Here we used dynamic state variable modeling to isolate effects of social information about food and predators on grouping behavior and fitness. We characterized optimal behavior from a set of strategies that included grouping with different numbers of conspecifics or heterospecifics and the option to forage or be vigilant over the course of a day. We show that the use of social information alone increases grouping behavior but constrains group size to limit competition, ultimately increasing individual fitness substantially across various e...

Journal ArticleDOI
TL;DR: It is concluded that human vision can detect much of the variation in coloration in the visible range, providing fodder for subsequent studies in ecology, evolution, behavior, and visual ecology.
Abstract: Assessing variation in animal coloration is difficult, as animals differ in their visual system properties. This has led some to propose that human vision can never be used to evaluate coloration, yet many studies have a long history of relying on human vision. To reconcile these views, we compared the reflectance spectra of preserved avian plumage elements with two measures that are human biased: RGB values from digital photographs and the corresponding reflectance spectra from a field guide. We measured 73 plumage elements across 14 bird species. The field guide reflectance spectra were drastically different from that of the actual birds, particularly for blue elements. However, principal component analyses on all three data sets indicated remarkably similar data structure. We conclude that human vision can detect much of the variation in coloration in the visible range, providing fodder for subsequent studies in ecology, evolution, behavior, and visual ecology.

Journal ArticleDOI
TL;DR: A dynamic state variable model is developed that captures the interplay among behavioral decisions, energy, reproductive needs, and the environment in baleen whales and provides a general framework to investigate the ecology of migration and assess the population consequences of disturbance, while identifying critical knowledge gaps.
Abstract: Integrating behavior and physiology is critical to formulating new hypotheses on the evolution of animal life-history strategies. Migratory capital breeders acquire most of the energy they need to sustain migration, gestation, and lactation before parturition. Therefore, when predicting the impact of environmental variation on such species, a mechanistic understanding of the physiology of their migratory behavior is required. Using baleen whales as a model system, we developed a dynamic state variable model that captures the interplay among behavioral decisions, energy, reproductive needs, and the environment. We applied the framework to blue whales (Balaenoptera musculus) in the eastern North Pacific Ocean and explored the effects of environmental and anthropogenic perturbations on female reproductive success. We demonstrate the emergence of migration to track prey resources, enabling us to quantify the trade-offs among capital breeding, body condition, and metabolic expenses. We predict that per...

Journal ArticleDOI
TL;DR: This work uses standard models of consumer-resource interactions to explore the ecological dynamics of natural selection and how various trait types influence these dynamics and the resulting structure of a community of coevolving species.
Abstract: Natural selection has both genetic and ecological dynamics. The fitnesses of individuals change with their ecological context, and so the form and strength of selective agents change with abiotic factors and the phenotypes and abundances of interacting species. I use standard models of consumer-resource interactions to explore the ecological dynamics of natural selection and how various trait types influence these dynamics and the resulting structure of a community of coevolving species. Evolutionary optima favored by natural selection depend critically on the abundances of interacting species, and the traits of species can undergo dynamic cycling in limited areas of parameter space. The ecological dynamics of natural selection can also drive shifts from one adaptive peak to another, and these ecologically driven adaptive peak shifts are fundamental to the dynamics of niche differentiation. Moreover, this ecological differentiation is fostered in more productive and more benign environments where species interactions are stronger and where the selection gradients generated by species interactions are stronger. Finally, community structure resulting from coevolution depends fundamentally on the types of traits that underlie species interactions. The ecological dynamics of the process cannot be simplified, neglected, or ignored if we are to build a predictive theory of natural selection.

Journal ArticleDOI
TL;DR: Survey data was analyzed using an integrative occupancy and structural equation modeling framework to quantify the strengths of hypothesized interaction pathways, and fine-scale spatiotemporal responses of nonapex predators to wolf activity clusters identified from radio-collar data showed a near guild-wide, positive response of nonapes predators to localized wolf activity.
Abstract: Competition and suppression are recognized as dominant forces that structure predator communities. Facilitation via carrion provisioning, however, is a ubiquitous interaction among predators that could offset the strength of suppression. Understanding the relative importance of these positive and negative interactions is necessary to anticipate community-wide responses to apex predator declines and recoveries worldwide. Using state-sponsored wolf (Canis lupus) control in Alaska as a quasi experiment, we conducted snow track surveys of apex, meso-, and small predators to test for evidence of carnivore cascades (e.g., mesopredator release). We analyzed survey data using an integrative occupancy and structural equation modeling framework to quantify the strengths of hypothesized interaction pathways, and we evaluated fine-scale spatiotemporal responses of nonapex predators to wolf activity clusters identified from radio-collar data. Contrary to the carnivore cascade hypothesis, both meso- and small predator occupancy patterns indicated guild-wide, negative responses of nonapex predators to wolf abundance variations at the landscape scale. At the local scale, however, we observed a near guild-wide, positive response of nonapex predators to localized wolf activity. Local-scale association with apex predators due to scavenging could lead to landscape patterns of mesopredator suppression, suggesting a key link between occupancy patterns and the structure of predator communities at different spatial scales.

Journal ArticleDOI
TL;DR: A single-species quantitative-genetics framework that predicts rates of environmental change beyond which populations go extinct is extended and finds that predators can indeed help prey persist if they sufficiently increase prey adaptedness (decrease phenotypic lag).
Abstract: To persist in a changing world, populations must adapt. The ability to adapt is influenced by interactions with other species, such as predators. Recent experiments and theory suggest that selective pressures arising from predation may help prey adapt phenotypically to changing environments, but how this influences persistence remains unclear. In particular, it has not yet been shown whether predator-induced adaptation can outweigh predator-imposed reductions in population size, allowing prey to persist when they would otherwise go extinct. Here we examine if (and if so, how) predation can enhance the ability of prey to persist in a directionally changing environment. To do so, we extend a single-species quantitative-genetics framework that predicts rates of environmental change beyond which populations go extinct. While we assume predation decreases prey density, we find that predators can indeed help prey persist if they sufficiently increase prey adaptedness (decrease phenotypic lag). We show two ways this can occur: (1) the selective push, in which predators consume maladapted individuals and thus add selection that pushes the mean prey trait toward its optimum; and (2) the evolutionary hydra effect, when predation reduces prey density and thereby increases prey birthrate, allowing more selective events per unit time and effectively reducing generation time. We also discuss how our results apply more broadly to sources of mortality beyond predation.

Journal ArticleDOI
TL;DR: An experiment with Escherichia coli is reviewed, in which 12 populations were founded from the same ancestral strain and have evolved for almost 30 years and more than 65,000 generations under the same conditions, to reveal convergence, divergence, and often a complicated mix thereof.
Abstract: Suitably designed experiments offer the possibility of quantifying evolutionary convergence because the fraction of replicate populations that converge is known. Here I review an experiment with Escherichia coli, in which 12 populations were founded from the same ancestral strain and have evolved for almost 30 years and more than 65,000 generations under the same conditions. The tension between divergence and convergence has been a major focus of this experiment. I summarize analyses of competitive fitness, correlated responses to different environments, cell morphology, the capacity to use a previously untapped resource, mutation rates, genomic changes, and within-population polymorphisms. These analyses reveal convergence, divergence, and often a complicated mix thereof. Complications include concordance in the direction of evolutionary change with sustained quantitative variation among populations, and the potential for a given trait to exhibit divergence on one timescale and convergence on another. Despite these complications, which also occur in nature, experiments provide a powerful way to study evolutionary convergence based on analyzing replicate lineages that experience the same environment.

Journal ArticleDOI
TL;DR: This work studies the stochastic population dynamics of a population exposed to random changes in the optimum phenotype for a quantitative trait that evolves in response to this moving optimum and reveals how the mean and variance of log population size change with the variance and autocorrelation of deviations of the evolving mean phenotype from the optimum.
Abstract: Many natural populations exhibit temporal fluctuations in abundance that are consistent with external forcing by a randomly changing environment As fitness emerges from an interaction between the phenotype and the environment, such demographic fluctuations probably include a substantial contribution from fluctuating phenotypic selection We study the stochastic population dynamics of a population exposed to random (plus possibly directional) changes in the optimum phenotype for a quantitative trait that evolves in response to this moving optimum We derive simple analytical predictions for the distribution of log population size over time both transiently and at stationarity under Gompertz density regulation These predictions are well matched by population- and individual-based simulations The log population size is approximately reverse gamma distributed, with a negative skew causing an excess of low relative to high population sizes, thus increasing extinction risk relative to a symmetric (eg, normal) distribution with the same mean and variance Our analysis reveals how the mean and variance of log population size change with the variance and autocorrelation of deviations of the evolving mean phenotype from the optimum We apply our results to the analysis of evolutionary rescue in a stochastic environment and show that random fluctuations in the optimum can substantially increase extinction risk by both reducing the expected growth rate and increasing the variance of population size by several orders of magnitude

Journal ArticleDOI
TL;DR: The results suggest that sex-specific modifiers such as testosterone can orchestrate sex-biased gene expression to facilitate the phenotypic development of sexual dimorphism while simultaneously reducing genetic correlations that would otherwise constrain the independent evolution of the sexes.
Abstract: The evolution of sexual dimorphism is predicted to occur through reductions in between-sex genetic correlations (rmf) for shared traits, but the physiological and genetic mechanisms that facilitate these reductions remain largely speculative. Here, we use a paternal half-sibling breeding design in captive brown anole lizards (Anolis sagrei) to show that the development of sexual size dimorphism is mirrored by the ontogenetic breakdown of rmf for body size and growth rate. Using transcriptome data from the liver (which integrates growth and metabolism), we show that sex-biased gene expression also increases dramatically between ontogenetic stages bracketing this breakdown of rmf. Ontogenetic increases in sex-biased expression are particularly evident for genes involved in growth, metabolism, and cell proliferation, suggesting that they contribute to both the development of sexual dimorphism and the breakdown of rmf. Mechanistically, we show that treatment of females with testosterone stimulates the expression of male-biased genes while inhibiting the expression of female-biased genes, thereby inducing male-like phenotypes at both organismal and transcriptomic levels. Collectively, our results suggest that sex-specific modifiers such as testosterone can orchestrate sex-biased gene expression to facilitate the phenotypic development of sexual dimorphism while simultaneously reducing genetic correlations that would otherwise constrain the independent evolution of the sexes.

Journal ArticleDOI
TL;DR: In this article, the authors apply an established model of phytoplankton growth to explore a trade-off between the ability of cells to replenish their internal quota and their ability to synthesize new biomass.
Abstract: Rates of metabolism and population growth are often assumed to decrease universally with increasing organism size. Recent observations have shown, however, that maximum population growth rates among phytoplankton smaller than ∼6 μm in diameter tend to increase with organism size. Here we bring together observations and theory to demonstrate that the observed change in slope is attributable to a trade-off between nutrient uptake and the potential rate of internal metabolism. Specifically, we apply an established model of phytoplankton growth to explore a trade-off between the ability of cells to replenish their internal quota (which increases with size) and their ability to synthesize new biomass (which decreases with size). Contrary to the metabolic theory of ecology, these results demonstrate that rates of resource acquisition (rather than metabolism) provide the primary physiological constraint on the growth rates of some of the smallest and most numerically abundant photosynthetic organisms on Earth.

Journal ArticleDOI
TL;DR: Competition is modeled as a tendency of sympatric species to evolve toward difference from one another, producing trait overdispersion and high phylogenetic signal and the model predicts elevated trait variance across species and a slowdown in evolutionary rate both across the clade and within each branch.
Abstract: The incorporation of ecological processes into models of trait evolution is important for understanding past drivers of evolutionary change. Species interactions have long been thought to be key drivers of trait evolution. However, models for comparative data that account for interactions between species are lacking. One of the challenges is that such models are intractable and difficult to express analytically. Here we present phylogenetic models of trait evolution that include interspecific competition among chosen species. Competition is modeled as a tendency of sympatric species to evolve toward difference from one another, producing trait overdispersion and high phylogenetic signal. The model predicts elevated trait variance across species and a slowdown in evolutionary rate both across the clade and within each branch. The model also predicts a reduction in correlation between otherwise correlated traits. We use an approximate Bayesian computation approach to estimate model parameters. We find reasonable power to detect competition in sufficiently large (20+ species) trees compared with Brownian trait evolution and with Ornstein-Uhlenbeck and early burst models. We apply the model to examine the evolution of bill morphology of Darwin's finches and find evidence that competition affects the evolution of bill length.

Journal ArticleDOI
TL;DR: A consilience of independent lines of evidence supports a lockstep model in which deciduousness evolved in situ, in parallel, and concurrent with a gradual coolingClimate, which may explain the massive convergence of adaptive strategies that characterizes the world's biomes.
Abstract: The deciduous habit of northern temperate trees and shrubs provides one of the most obvious examples of convergent evolution, but how did it evolve? Hypotheses based on the fossil record posit that deciduousness evolved first in response to drought or darkness and preadapted certain lineages as cold climates spread. An alternative is that evergreens first established in freezing environments and later evolved the deciduous habit. We monitored phenological patterns of 20 species of Viburnum spanning tropical, lucidophyllous (subtropical montane and warm temperate), and cool temperate Asian forests. In lucidophyllous forests, all viburnums were evergreen plants that exhibited coordinated leaf flushes with the onset of the rainy season but varied greatly in the timing of leaf senescence. In contrast, deciduous species exhibited tight coordination of both flushing and senescence, and we found a perfect correlation between the deciduous habit and prolonged annual freezing. In contrast to previous stepwise hypotheses, a consilience of independent lines of evidence supports a lockstep model in which deciduousness evolved in situ, in parallel, and concurrent with a gradual cooling climate. A pervasive selective force combined with the elevated evolutionary accessibility of a particular response may explain the massive convergence of adaptive strategies that characterizes the world's biomes.