The evolution of passive dispersal versus habitat selection have differing emergent consequences in metacommunities.

Dispersal among local communities is fundamental to the metacommunity concept but is only important to the metacommunity structure if dispersal causes distortions of species abundances away from what local ecological conditions favour. We know from much previous work that dispersal can cause such abundance distortions. However, almost all previous theoretical studies have only considered one species alone or two interacting species (e.g. competitors or predator and prey). Moreover, a systematic analysis is needed of whether different dispersal strategies (e.g. passive dispersal versus demographic habitat selection) result in different abundance distortion patterns, how these distortion patterns change with local food web structure, and how the dispersal propensities of the interacting species might evolve in response to one another. In this article, we show using computer simulations and analytical models that abundance distortions occur in simple food webs with both passive dispersal and habitat selection, but habitat selection causes larger distortions. Additionally, patterns in the evolution of dispersal propensity in interacting species are very different for these two dispersal strategies. This study identifies that the dispersal strategies employed by interacting species critically shape how dispersal will influence metacommunity structure. This article is part of the theme issue 'Diversity-dependence of dispersal: interspecific interactions determine spatial dynamics'.

Partial Clonality Expands the Opportunity for Spatial Adaptation.

AbstractReproductive mode may strongly impact adaptation in spatially varying populations linked by dispersal, especially when sexual and clonal offspring differ in dispersal. We determined how spatial structure affects adaptation in populations with mixed clonal and sexual reproduction. In a source-sink quantitative genetic deterministic model (with stabilizing selection around different optima), greater clonal reproduction or parent-offspring association (a measure of the part of the parent's phenotype other than the additive genetic component inherited by clonal offspring) increased the selective difference (difference between phenotypic optima) allowing sink populations to adapt. Given dispersal differences between clonally and sexually produced juveniles, adaptation increased with an increasing fraction of clonal dispersers. When considering migrational meltdown, partially clonal reproduction reduced cases where dispersal caused habitat loss. Stochastic individual-based simulations support these results, although the effect of differential dispersal was reversed, with decreased clonal dispersal allowing greater adaptation. These results parallel earlier findings that for an instantaneous shift in phenotypic optimum, increasing clonality allowed population persistence for a greater shift; here, selective change is spatial rather than temporal. These results may help explain the success of many partially clonal organisms in invading new habitats, complementing traditional explanations based on avoiding Allee effects.

Feeding the fever: Complex host-pathogen dynamics along continuous resource gradients.

Food has long been known to perform dual functions of nutrition and medicine, but mounting evidence suggests that complex host-pathogen dynamics can emerge along continuous resource gradients. Empirical examples of nonmonotonic responses of infection with increasing host resources (e.g., low prevalence at low and high resource supply but high prevalence at intermediate resources) have been documented across the tree of life, but these dynamics, when observed, often are interpreted as nonintuitive, idiosyncratic features of pathogen and host biology. Here, by developing generalized versions of existing models of resource dependence for within- and among-host infection dynamics, we provide a synthetic view of nonmonotonic infection dynamics. We demonstrate that where resources jointly impact two (or more) processes (e.g., growth, defense, transmission, mortality, predation), nonmonotonic infection dynamics, including alternative states, can emerge across a continuous resource supply gradient. We review the few empirical examples that concurrently measured resource effects on multiple rates and pair this with a wide range of examples in which resource dependence of multiple rates could generate nonmonotonic infection outcomes under realistic conditions. This review and generalized framework highlight the likely generality of such resource effects in natural systems and point to opportunities ripe for future empirical and theoretical work.

Landscape experiments unlock relationships among habitat loss, fragmentation, and patch-size effects.

Habitat loss is often considered the greatest near-term threat to biodiversity, while the impact of habitat fragmentation remains intensely debated. A key issue of this debate centers on the problem of scale-landscape or patch-at which to assess the consequences of fragmentation. Yet patterns are often confounded across scales, and experimental designs that could solve this scaling problem remain scarce. We conducted two field experiments in 30 experimental landscapes in which we manipulated habitat loss, fragmentation, and patch size for a community of four insect herbivores that specialize on the cactus Opuntia. In the first experiment, we destroyed 2088 Opuntia patches in either aggregated or random patterns and compared the relative effects of landscape-scale loss and fragmentation to those of local patch size on species occurrence. This experiment focused on manipulating the relative separation of remaining patches, where we hypothesized that aggregated loss would disrupt dispersal more than random loss, leading to lower occurrence. In the second experiment, we destroyed 759 Opuntia patches to generate landscapes that varied in patch number and size for a given amount of habitat loss and assessed species occurrence. This experiment focused on manipulating the subdivision of remaining habitat, where we hypothesized that an increase in the number of patches for a given amount of loss would lead to negative effects on occurrence. For both, we expected that occurrence would increase with patch size. We find strong evidence for landscape-scale effects of habitat fragmentation, with aggregated loss and a larger number of patches for a given amount of habitat loss leading to a lower frequency of patches occupied in landscapes. In both experiments, occurrence increased with patch size, yet interactions of patch size and landscape-scale loss and fragmentation drove species occurrence in patches. Importantly, the direction of effects were consistent across scales and effects of patch size were sufficient to predict the effects of habitat loss and fragmentation across entire landscapes. Our experimental results suggest that changes at both the patch and landscape scales can impact populations, but that a long-standing pattern-the patch-size effect-captures much of the key variation shaping patterns of species occurrence.

Litter, Plant Competition, and Ecosystem Dynamics: A Theoretical Perspective.

AbstractCommunity structure depends jointly on species' responses to, and effects on, environmental factors. Many such factors, including detritus, are studied in ecosystem ecology. Detritus in terrestrial ecosystems is dominated by plant litter (nonliving organic material), which, in addition to its role in material cycling, can act as a niche factor modulating interactions among plants. Litter thus links traditional community and ecosystem processes, which are often studied separately. We explore this connection using population dynamics models of two plant species and a litter pool. We first find conditions determining the outcome of interactions between these species, highlighting the role that litter plays and the role of broader ecosystem parameters, such as decomposition rate. Species trade-offs in tolerance to direct competition and litter-based interference competition allow for coexistence, provided the litter-tolerant species produces more litter at the population level; otherwise, priority effects may result. When species coexist, litter-mediated interactions between plants disrupt the traditional relationship between biomass accumulation and decomposition. Increasing decomposition rate may have no effect on standing litter density and, in some cases, may even increase litter load. These results illustrate how ecosystem variables can influence community outcomes that then feed back to influence the ecosystem.

A rodent herbivore reduces its predation risk through ecosystem engineering.

Predator-prey interactions are ubiquitous and powerful forces that structure ecological communities.1-3 Habitat complexity has been shown to be particularly important in regulating the strength of predator-prey interactions.4-6 While it is well established that changes in habitat structure can alter the efficacy of predatory and anti-predatory behaviors,7-9 little is known about the consequences of engineering activity by prey species who modify the external environment to reduce their own predation risk. Using field surveys and manipulative experiments, we evaluated how habitat modification by Brandt's voles (Lasiopodomys brandtii) influences predation risk from a principal avian predator (shrike; Lanius spp.) in a steppe grassland, located in Inner Mongolia, China. We found that voles actively modify habitat structure by cutting down a large, unpalatable bunchgrass species (Achnatherum splendens) in the presence of shrikes, a behavior that disappeared when these avian predators were excluded experimentally. The damage activities of these voless dramatically decreased the volume of unpalatable grasses, which in turn reduced visitations by shrikes and thus mortality rates. Our study shows that herbivorous prey that act as ecosystem engineers can directly reduce their own predation risk by modifying habitat structure. Given the ubiquity of predation risks faced by consumers, and the likely ability of many consumers to alter the habitat structure in which they live, the interplay between predation risk and ecosystem engineering may be an important but unappreciated mechanism at play in natural communities.

The factors that favor adaptive habitat construction versus non-adaptive environmental conditioning.

Adaptive habitat construction is a process by which individuals alter their environment so as to increase their (inclusive) fitness. Such alterations are a subset of the myriad ways that individuals condition their environment. We present an individual-based model of habitat construction to explore what factors might favor selection when the benefits of environmental alterations are shared by individuals of the same species. Our results confirm the predictions of inclusive fitness and group selection theory and expectations based on previous models that construction will be more favored when its benefits are more likely to be directed to self or near kin. We found that temporal variation had no effect on the evolution of construction. For spatial heterogeneity, construction was disfavored when the spatial pattern of movement did not match the spatial pattern of environmental heterogeneity, especially when there was spatial heterogeneity in the optimal amount of construction. Under those conditions, very strong selection was necessary to favor genetic differentiation of construction propensity among demes. We put forth a constitutive theory for the evolution of adaptive habitat construction that unifies our model with previous verbal and quantitative models into a formal conceptual framework.

Temporal variation may have diverse impacts on range limits.

Environmental fluctuations are pervasive in nature, but the influence of non-directional temporal variation on range limits has received scant attention. We synthesize insights from the literature and use simple models to make conceptual points about the potentially wide range of ecological and evolutionary effects of temporal variation on range limits. Because organisms respond nonlinearly to environmental conditions, temporal variation can directionally alter long-term growth rates, either to shrink or to expand ranges. We illustrate this diversity of outcomes with a model of competition along a mortality gradient. Temporal variation can permit transitions between alternative states, potentially facilitating range expansion. We show this for variation in dispersal, using simple source-sink population models (with strong Allee effects, or with gene flow hampering local adaptation). Temporal variation enhances extinction risk owing to demographic stochasticity, rare events, and loss of genetic variation, all tending to shrink ranges. However, specific adaptations to exploit variation (including dispersal) may permit larger ranges than in similar but constant environments. Grappling with temporal variation is essential both to understand eco-evolutionary dynamics at range limits and to guide conservation and management strategies. This article is part of the theme issue 'Species' ranges in the face of changing environments (Part II)'.

The Effect of Predator Population Dynamics on Batesian Mimicry Complexes.

AbstractUnderstanding Batesian mimicry is a classic problem in evolutionary biology. In Batesian mimicry, a defended species (the model) is mimicked by an undefended species (the mimic). Prior theories have emphasized the role of predator behavior and learning as well as evolution in model-mimic complexes but have not examined the role of population dynamics in potentially governing the relative abundances and even persistence of model-mimic systems. Here, we examined the effect of the population dynamics of predators and alternative prey on the prevalence of warning-signaling prey composed of models and mimics. Using optimal foraging theory and signal detection theory, we found that the inclusion of predator and alternative prey population dynamics could reverse traditional theoretical predictions: as alternative prey increase in numbers, mimics suffer because larger populations of predators are maintained, resulting in apparent competition. Under some circumstances, apparent competition affects model populations as well, although not as severely as it affects mimics. Our results bear on the intriguing puzzle that in nature warning signals are relatively scarce, yet experiments suggest that such signals can be highly advantageous. The availability of alternative prey and numerical responses by predators can overwhelm advantages observed in experiments to keep warning signals in model-mimic systems relatively scarce.

Do I build or do I move? Adaptation by habitat construction versus habitat choice.

Trait adaptation to a heterogeneous environment can occur through six modes: genetic differentiation of those traits, a jack-of-all-trades phenotypic uniformity, diversified bet-hedging, phenotypic plasticity, habitat choice, and habitat construction. A key question is what circumstances favor one mode over another, and how they might interact if a system can express more than one mode at a time. We examined the joint evolution of habitat choice and habitat construction using individual-based simulations. We manipulated when during the life cycle construction occurred and the fitness value of construction. We found that for our model habitat construction was nearly always favored over habitat choice, especially if construction happened after dispersal. Because of the ways that the various modes of adaptation interact with each other, there is no simple answer as to which will be favored; it depends on details of the biology and ecology of a given system.

Metapopulation capacity determines food chain length in fragmented landscapes.

Metapopulation capacity provides an analytic tool to quantify the impact of landscape configuration on metapopulation persistence, which has proven powerful in biological conservation. Yet surprisingly few efforts have been made to apply this approach to multispecies systems. Here, we extend metapopulation capacity theory to predict the persistence of trophically interacting species. Our results demonstrate that metapopulation capacity could be used to predict the persistence of trophic systems such as prey-predator pairs and food chains in fragmented landscapes. In particular, we derive explicit predictions for food chain length as a function of metapopulation capacity, top-down control, and population dynamical parameters. Under certain assumptions, we show that the fraction of empty patches for the basal species provides a useful indicator to predict the length of food chains that a fragmented landscape can support and confirm this prediction for a host-parasitoid interaction. We further show that the impact of habitat changes on biodiversity can be predicted from changes in metapopulation capacity or approximately by changes in the fraction of empty patches. Our study provides an important step toward a spatially explicit theory of trophic metacommunities and a useful tool for predicting their responses to habitat changes.

Disturbance-induced emigration: an overlooked mechanism that reduces metapopulation extinction risk.

Emigration propensity (i.e., the tendency to leave undisturbed patches) is a key life-history trait of organisms in metapopulations with local extinctions and colonizations. Metapopulation models of dispersal evolution typically assume that patch disturbance kills all individuals within the patch, thus causing local extinction. However, individuals may instead be able to leave a patch when it is disturbed, either by fleeing before being killed or simply because the disturbance destroys the patch without causing mortality. This scenario may pertain to a wide range of organisms from horizontally transmitted symbionts, to aquatic insects inhabiting temporary ponds, to vertebrates living in fragmented forests. We generalized a Levins-type metapopulation model of dispersal evolution by adding a new parameter of disturbance escape probability, which incorporates a second source of dispersal into the model: disturbance-induced emigration. We show that disturbance escape expands the domain of metapopulation viability and selects for lower rates of emigration propensity when disturbance rates are high. The fitness gains from disturbance-induced emigration are generally moderate, suggesting that disturbance escape might act more as a complementary dispersal strategy rather than a replacement to emigration propensity, at least for metapopulations that meet the assumptions of the Levins-type model. Yet disturbance-induced emigration may in some circumstances rescue a metapopulation from long-term extinction when the combination of high disturbance rates and low local population growth rates compromises its viability. Further, a metapopulation could persist exclusively by disturbance escape if local carrying capacities are large enough to counterbalance two sources of mortality: mortality driven by disturbance and mortality during dispersal. This study opens two promising research lines: (1) the investigation of disturbance escape in metapopulations of ephemeral habitats with unsaturated populations and non-equilibrium dynamics and (2) the incorporation of information costs to investigate the joint evolution of disturbance escape and emigration propensity.

Why aren't warning signals everywhere? On the prevalence of aposematism and mimicry in communities.

Warning signals are a striking example of natural selection present in almost every ecological community - from Nordic meadows to tropical rainforests, defended prey species and their mimics ward off potential predators before they attack. Yet despite the wide distribution of warning signals, they are relatively scarce as a proportion of the total prey available, and more so in some biomes than others. Classically, warning signals are thought to be governed by positive density-dependent selection, i.e. they succeed better when they are more common. Therefore, after surmounting this initial barrier to their evolution, it is puzzling that they remain uncommon on the scale of the community. Here, we explore factors likely to determine the prevalence of warning signals in prey assemblages. These factors include the nature of prey defences and any constraints upon them, the behavioural interactions of predators with different prey defences, the numerical responses of predators governed by movement and reproduction, the diversity and abundance of undefended alternative prey and Batesian mimics in the community, and variability in other ecological circumstances. We also discuss the macroevolution of warning signals. Our review finds that we have a basic understanding of how many species in some taxonomic groups have warning signals, but very little information on the interrelationships among population abundances across prey communities, the diversity of signal phenotypes, and prey defences. We also have detailed knowledge of how a few generalist predator species forage in artificial laboratory environments, but we know much less about how predators forage in complex natural communities with variable prey defences. We describe how empirical work to address each of these knowledge gaps can test specific hypotheses for why warning signals exhibit their particular patterns of distribution. This will help us to understand how behavioural interactions shape ecological communities.

The evolution of habitat construction with and without phenotypic plasticity.

Habitat construction and phenotypic plasticity are alternative responses to variable environments. We explored evolution along an environmental gradient of habitat construction alone and in combination with phenotypic plasticity using individual-based simulations that manipulated the fitness benefit of construction and whether construction maintained or eliminated that gradient. Construction was favored when its benefits were more likely to flow to the immediate offspring of the constructing individuals. Habitat construction and phenotypic plasticity traded off against each other or plasticity was selected against, depending on how the optimum environment varied and with the fitness value of construction. When selection favored differences in the amount of construction along the environmental gradient, genetic differentiation for habitat construction increased as the fitness value of construction increased. The degree to which each adaptive response was likely to evolve also depended on the precise ordering of life history events. Adaptive habitat construction does not always occur and may be selected against.

Environmental fluctuations dampen the effects of clonal reproduction on evolutionary rescue.

Evolutionary rescue occurs when genetic change allows a population to persist in response to an environmental change that would otherwise have led to extinction. Most studies of evolutionary rescue assume that species have either fully clonal or fully sexual reproduction; however, many species have partially clonal reproductive strategies in which they reproduce both clonally and sexually. Furthermore, the few evolutionary rescue studies that have evaluated partially clonal reproduction did not consider fluctuations in the environment, which are nearly ubiquitous in nature. Here, we use individual-based simulations to investigate how environmental fluctuations (either uncorrelated or positively autocorrelated) influence the effect of clonality on evolutionary rescue. We show that, for moderate magnitudes of environmental fluctuations, as was found in the absence of fluctuations, increasing the degree of clonality increases the probability of population persistence in response to an abrupt environmental change, but decreases persistence in response to a continuous, directional environmental change. However, with large magnitudes of fluctuations, both the benefits of clonality following a step change and the detrimental effects of clonality following a continuous, directional change are generally reduced; in fact, in the latter scenario, increasing clonality can even become beneficial if environmental fluctuations are autocorrelated. We also show that increased generational overlap dampens the effects of environmental fluctuations. Overall, we demonstrate that understanding the evolutionary rescue of partially clonal organisms requires not only knowledge of the species life history and the type of environmental change, but also an understanding of the magnitude and autocorrelation of environmental fluctuations.

Revisiting the Role of Hyperparasitism in the Evolution of Virulence.

AbstractHyperparasitism denotes the natural phenomenon where a parasite infecting a host is in turn infected by its own parasite. Hyperparasites can shape the dynamics of host-parasite interactions and often have a deleterious impact on pathogens, an important class of parasites, causing a reduction in their virulence and transmission rate. Hyperparasitism thus could be an important tool of biological control. However, host-parasite-hyperparasite systems have so far been outside the mainstream of modeling studies, especially those dealing with eco-evolutionary aspects of species interactions. Here, we theoretically explore the evolution of life-history traits in a generic host-parasite-hyperparasite system, focusing on parasite virulence and the positive impact that hyperparasitism has on the host population. We also explore the coevolution of life-history traits of the parasite and hyperparasite, using adaptive dynamics and quantitative genetics frameworks to identify evolutionarily singular strategies. We find that in the presence of hyperparasites, the evolutionarily optimal pathogen virulence generally shifts toward more virulent strains. However, even in this case the use of hyperparasites in biocontrol could be justified, since overall host mortality decreases. An intriguing possible outcome of the evolution of the hyperparasite can be its evolutionary suicide.

The interplay of movement and spatiotemporal variation in transmission degrades pandemic control.

Successful public health regimes for COVID-19 push below unity long-term regional Rt -the average number of secondary cases caused by an infectious individual. We use a susceptible-infectious-recovered (SIR) model for two coupled populations to make the conceptual point that asynchronous, variable local control, together with movement between populations, elevates long-term regional Rt , and cumulative cases, and may even prevent disease eradication that is otherwise possible. For effective pandemic mitigation strategies, it is critical that models encompass both spatiotemporal heterogeneity in transmission and movement.