Coevolution of species colonisation rates controls food-chain length in spatially structured food webs.

How the complexity of food webs depends on environmental variables is a long-standing ecological question. It is unclear though how food-chain length should vary with adaptive evolution of the constitutive species. Here we model the evolution of species colonisation rates and its consequences on occupancies and food-chain length in metacommunities. When colonisation rates can evolve, longer food-chains can persist. Extinction, perturbation and habitat loss all affect evolutionarily stable colonisation rates, but the strength of the competition-colonisation trade-off has a major role: weaker trade-offs yield longer chains. Although such eco-evo dynamics partly alleviates the spatial constraint on food-chain length, it is no magic bullet: the highest, most vulnerable, trophic levels are also those that least benefit from evolution. We provide qualitative predictions regarding how trait evolution affects the response of communities to disturbance and habitat loss. This highlights the importance of eco-evolutionary dynamics at metacommunity level in determining food-chain length.

When complex movement yields simple dispersal: behavioural heterogeneity, spatial spread and parasitism in groups of micro-wasps.

BACKGROUND: Understanding how behavioural dynamics, inter-individual variability and individual interactions scale-up to shape the spatial spread and dispersal of animal populations is a major challenge in ecology. For biocontrol agents, such as the microscopic Trichogramma parasitic wasps, an understanding of movement strategies is also critical to predict pest-suppression performance in the field. METHODS: We experimentally studied the spatial propagation of groups of parasitoids and their patterns of parasitism. We investigated whether population spread is density-dependent, how it is affected by the presence of hosts, and whether the spatial distribution of parasitism (dispersal kernel) can be predicted from the observed spread of individuals. Using a novel experimental device and high-throughput imaging techniques, we continuously tracked the spatial spread of groups of parasitoids over large temporal and spatial scales (8 h; and 6 m, ca. 12,000 body lengths). We could thus study how population density, the presence of hosts and their spatial distribution impacted the rate of population spread, the spatial distribution of individuals during population expansion, the overall rate of parasitism and the dispersal kernel (position of parasitism events). RESULTS: Higher population density accelerated population spread, but only transiently: the rate of spread reverted to low values after 4 h, in a "tortoise-hare" effect. Interestingly, the presence of hosts suppressed this transiency and permitted a sustained high rate of population spread. Importantly, we found that population spread did not obey classical diffusion, but involved dynamical switches between resident and explorer movement modes. Population distribution was therefore not Gaussian, though surprisingly the distribution of parasitism (dispersal kernel) was. CONCLUSIONS: Even homogenous asexual groups of insects develop behavioural heterogeneities over a few hours, and the latter control patterns of population spread. Behavioural switching between resident and explorer states determined population distribution, density-dependence and dispersal. A simple Gaussian dispersal kernel did not reflect classical diffusion, but rather the interplay of several non-linearities at individual level. These results highlight the need to take into account behaviour and inter-individual heterogeneity to understand population spread in animals.

Migration pulsedness alters patterns of allele fixation and local adaptation in a mainland-island model.

Geneflow across populations is a critical determinant of population genetic structure, divergence, and local adaptation. While evolutionary theory typically envisions geneflow as a continuous connection among populations, many processes make it fluctuating and intermittent. We analyze a mainland-island model where migration occurs as recurrent "pulses." We derive mathematical predictions regarding how the level of migration pulsedness affects the effective migration rate, for neutral and selected mainland alleles. We find that migration pulsedness can either decrease or increase geneflow, depending on the selection regime. Pulsedness increases geneflow for sufficiently (counter)selected alleles (s

Consistent variations in personality traits and their potential for genetic improvement in biocontrol agents: Trichogramma evanescens as a case study.

Improvements in the biological control of agricultural pests require improvements in the phenotyping methods used by practitioners to select efficient biological control agent (BCA) populations in industrial rearing or field conditions. Consistent inter-individual variations in behaviour (i.e. animal personality) probably affect BCA efficiency, but have never been taken into account in the development of phenotyping methods, despite having characteristics useful for phenotyping: repeatable (by definition), often heritable, etc. We developed a video-tracking method targeting animal personality traits and evaluated the feasibility of its use for genetic improvement in the BCA Trichogramma evanescens, by phenotyping 1,049 individuals from 24 isogenic lines. We found consistent individual variations in boldness, activity and exploration. Personality differences between the 24 isogenic lines suggested a genetic origin of the variations in activity and exploration (broad-sense heritability estimates of 0.06 to 0.11) and revealed a trade-off between exploration and fecundity.

Multi-seasonal modelling of plant-nematode interactions reveals efficient plant resistance deployment strategies.

Root-knot nematodes, Meloidogyne spp., are soil-borne polyphagous pests with major impact on crop yield worldwide. Resistant crops efficiently control avirulent root-knot nematodes, but favour the emergence of virulent forms. Since virulence is associated with fitness costs, susceptible crops counter-select virulent root-knot nematodes. In this study, we identify optimal rotation strategies between susceptible and resistant crops to control root-knot nematodes and maximize crop yield. We developed an epidemiological model describing the within-season dynamics of avirulent and virulent root-knot nematodes on susceptible or resistant plant root-systems, and their between-season survival. The model was fitted to experimental data and used to predict yield-maximizing rotation strategies, with special attention to the impact of epidemic severity and genetic parameters. Crop rotations were found to be efficient under realistic parameter ranges. They were characterized by low ratios of resistant plants and were robust to parameter uncertainty. Rotations provide significant gain over resistant-only strategies, especially under intermediate fitness costs and severe epidemic contexts. Switching from the current general deployment of resistant crops to custom rotation strategies could not only maintain or increase crop yield, but also preserve the few and valuable R-genes available.

How community adaptation affects biodiversity-ecosystem functioning relationships.

Evidence is growing that evolutionary dynamics can impact biodiversity-ecosystem functioning (BEF) relationships. However the nature of such impacts remains poorly understood. Here we use a modelling approach to compare random communities, with no trait evolutionary fine-tuning, and co-adapted communities, where traits have co-evolved, in terms of emerging biodiversity-productivity, biodiversity-stability and biodiversity-invasion relationships. Community adaptation impacted most BEF relationships, sometimes inverting the slope of the relationship compared to random communities. Biodiversity-productivity relationships were generally less positive among co-adapted communities, with reduced contribution of sampling effects. The effect of community-adaptation, though modest regarding invasion resistance, was striking regarding invasion tolerance: co-adapted communities could remain very tolerant to invasions even at high diversity. BEF relationships are thus contingent on the history of ecosystems and their degree of community adaptation. Short-term experiments and observations following recent changes may not be safely extrapolated into the future, once eco-evolutionary feedbacks have taken place.

Parallel evolution of behaviour during independent host-shifts following maize introduction into Asia and Europe.

Maize was introduced into opposite sides of Eurasia 500 years ago, in Western Europe and in Asia. This caused two host-shifts in the phytophagous genus Ostrinia; O. nubilalis (the European corn borer; ECB) and O. furnacalis (the Asian corn borer; ACB) are now major pests of maize worldwide. They originated independently from Dicot-feeding ancestors, similar to O. scapulalis (the Adzuki bean borer; ABB). Unlike other host-plants, maize is yearly harvested, and harvesting practices impose severe mortality on larvae found above the cut-off line. Positive geotaxis in the ECB has been proposed as a behavioural adaptation to harvesting practices, allowing larvae to move below the cut-off line and thus escape harvest mortality. Here, we test whether the same behavioural adaptation evolved independently in Europe and in Asia. We sampled eight genetically differentiated ECB, ACB and ABB populations in France and China and monitored geotaxis through the entire larval development in artificial stacks mimicking maize stems. We find that all ECB and ACB populations show a similar tendency to move down during the latest larval stages, a behaviour not observed in any European or Asian ABB population. The behaviour is robustly expressed regardless of larval density, development mode or environmental conditions. Our results indicate that maize introduction triggered parallel behavioural adaptations in Europe and Asia, harvest selection presumably being the main driver.

Diversity spurs diversification in ecological communities.

Diversity is a fundamental, yet threatened, property of ecological systems. The idea that diversity can itself favour diversification, in an autocatalytic process, is very appealing but remains controversial. Here, we study a generalized model of ecological communities and investigate how the level of initial diversity influences the possibility of evolutionary diversification. We show that even simple models of intra- and inter-specific ecological interactions can predict a positive effect of diversity on diversification: adaptive radiations may require a threshold number of species before kicking-off. We call this phenomenon DDAR (diversity-dependent adaptive radiations) and identify mathematically two distinct pathways connecting diversity to diversification, involving character displacement and the positive diversity-productivity relationship. Our results may explain observed delays in adaptive radiations at the macroscale and diversification patterns reported in experimental microbial communities, and shed new light on the dynamics of ecological diversity, the diversity-dependence of diversification rates, and the consequences of biodiversity loss.

Metacommunity speciation models and their implications for diversification theory.

The emergence of new frameworks combining evolutionary and ecological dynamics in communities opens new perspectives on the study of speciation. By acknowledging the relative contribution of local and regional dynamics in shaping the complexity of ecological communities, metacommunity theory sheds a new light on the mechanisms underlying the emergence of species. Three integrative frameworks have been proposed, involving neutral dynamics, niche theory, and life history trade-offs respectively. Here, we review these frameworks of metacommunity theory to emphasise that: (1) studies on speciation and community ecology have converged towards similar general principles by acknowledging the central role of dispersal in metacommunities dynamics, (2) considering the conditions of emergence and maintenance of new species in communities has given rise to new models of speciation embedded in the metacommunity theory, (3) studies of diversification have shifted from relating phylogenetic patterns to landscapes spatial and ecological characteristics towards integrative approaches that explicitly consider speciation in a mechanistic ecological framework. We highlight several challenges, in particular the need for a better integration of the eco-evolutionary consequences of dispersal and the need to increase our understanding on the relative rates of evolutionary and ecological changes in communities.

On the evolution of dispersal via heterogeneity in spatial connectivity.

Dispersal has long been recognized as a mechanism that shapes many observed ecological and evolutionary processes. Thus, understanding the factors that promote its evolution remains a major goal in evolutionary ecology. Landscape connectivity may mediate the trade-off between the forces in favour of dispersal propensity (e.g. kin-competition, local extinction probability) and those against it (e.g. energetic or survival costs of dispersal). It remains, however, an open question how differing degrees of landscape connectivity may select for different dispersal strategies. We implemented an individual-based model to study the evolution of dispersal on landscapes that differed in the variance of connectivity across patches ranging from networks with all patches equally connected to highly heterogeneous networks. The parthenogenetic individuals dispersed based on a flexible logistic function of local abundance. Our results suggest, all else being equal, that landscapes differing in their connectivity patterns will select for different dispersal strategies and that these strategies confer a long-term fitness advantage to individuals at the regional scale. The strength of the selection will, however, vary across network types, being stronger on heterogeneous landscapes compared with the ones where all patches have equal connectivity. Our findings highlight how landscape connectivity can determine the evolution of dispersal strategies, which in turn affects how we think about important ecological dynamics such as metapopulation persistence and range expansion.

The functional response predicts the effect of resource distribution on the optimal movement rate of consumers.

Understanding how often individuals should move when foraging over patchy habitats is a central question in ecology. By combining optimality and functional response theories, we show analytically how the optimal movement rate varies with the average resource level (enrichment) and resource distribution (patch heterogeneity). We find that the type of functional response predicts the effect of enrichment in homogeneous habitats: enrichment should decrease movement for decelerating functional responses, but increase movement for accelerating responses. An intermediate resource level thus maximises movement for type-III responses. Counterintuitively, greater movement costs favour an increase in movement. In heterogeneous habitats predictions further depend on how enrichment alters the variance of resource distribution. Greater patch variance always increases the optimal rate of movement, except for type-IV functional responses. While the functional response is well established as a fundamental determinant of consumer-resource dynamics, our results indicate its importance extends to the understanding of individual movement strategies.

How optimal foragers should respond to habitat changes: a reanalysis of the Marginal Value Theorem.

The Marginal Value Theorem (MVT) is a cornerstone of biological theory. It connects the quality and distribution of patches in a fragmented habitat to the optimal time an individual should spend exploiting them, and thus its optimal rate of movement. However, predictions regarding how habitat alterations should impact optimal strategies have remained elusive, with heavy reliance on graphical arguments. Here we derive the sensitivity of realized fitness and optimal residence times to general habitat attributes, for homogeneous and heterogeneous habitats, retaining the level of generality of the MVT. We provide new predictions on how altering travel times, patch qualities and/or relative abundances should affect optimal strategies, and study the consequences of habitat heterogeneity. We show that knowledge of average characteristics is in general not sufficient to predict the change in the average rate of movement. We apply our results to examine the conditions under which the optimal strategies are invariant to scaling. We prove a previously conjectured form of invariance in homogeneous habitats, but show that invariances to scaling are not generic in heterogeneous habitats. We also consider the relative exploitation of patches that differ in quality, clarifying the conditions under which it is adaptive to stay longer on poorer patches.

Extending the concept of keystone species to communities and ecosystems.

Keystone species are defined as having disproportionate importance in their community. This concept has proved useful and is now often used in conservation ecology. Here, we introduce the concept of keystone communities (and ecosystems) within metacommunities (and metaecosystems). We define keystone and burden communities as communities with impacts disproportionately large (positive or negative respectively) relative to their weight in the metacommunity. We show how a simple metric, based on the effects of single-community removals, can characterise communities along a 'keystoneness' axis. We illustrate the usefulness of this approach with examples from two different theoretical models. We further distinguish environmental heterogeneity from species trait heterogeneity as determinants of keystoneness. We suggest that the concept of keystone communities/ecosystems will be highly beneficial, not only as a fundamental step towards understanding species interactions in a spatial context, but also as a tool for the management of disturbed landscapes.

Competition-colonization dynamics in experimental bacterial metacommunities.

One of the simplest hypotheses used to explain species coexistence is the competition-colonization trade-off, that is, species can stably coexist in a landscape if they show a trade-off between competitive and colonization abilities. Despite extensive theory, the dynamics predicted to result from competition-colonization trade-offs are largely untested. Landscape change, such as habitat destruction, is thought to greatly influence coexistence under competition-colonization dynamics, although there is no formal test of this prediction. Here we present the first illustration of competition-colonization dynamics that fully transposes theory into a controlled experimental metacommunity of two Pseudomonas bacterial strains. The competition-colonization dynamics were achieved by directly manipulating trade-off strength and colonization rates to generate the full range of coexistence conditions and responses to habitat destruction. Our study successfully generates competition-colonization dynamics matching theoretical predictions, and our results further reveal a negative relationship between diversity and productivity when scaling up to entire metacommunities.

From N400 to N300: variations in the timing of semantic processing with repetition.

The present study aimed to explore the variations of semantic processing according to the number of target words (i.e., 4, 12 and 24) and according to the number of repetitions (i.e, 1 to 15). The number of targets had no impact on the N400 brain potential, the index of semantic processing, nor on the late positive component (LPC), an index of episodic encoding and retrieval. Analyses of the effects of the number of repetitions showed that the duration of semantic processes--assessed by measuring N400 latency--was linearly shortened along repetitions while their extent--as indexed by N400 amplitude--remained constant after the second presentation. In contrast, the extent of episodic processes--as indexed by LPC amplitude--was found to increase linearly with repetition. By showing that N400 latency may be much less stable than previously thought, these results bring new constraints on the functional correlates of this key stage in the processing of semantic information. They also suggest that semantic processes can be studied at high repetition rates whatever the number of target stimuli. Finally, our findings show that each episode of prior presentation has an impact on the late processing of a stimulus despite the absence of an explicit memory task.

High plant diversity is needed to maintain ecosystem services.

Biodiversity is rapidly declining worldwide, and there is consensus that this can decrease ecosystem functioning and services. It remains unclear, though, whether few or many of the species in an ecosystem are needed to sustain the provisioning of ecosystem services. It has been hypothesized that most species would promote ecosystem services if many times, places, functions and environmental changes were considered; however, no previous study has considered all of these factors together. Here we show that 84% of the 147 grassland plant species studied in 17 biodiversity experiments promoted ecosystem functioning at least once. Different species promoted ecosystem functioning during different years, at different places, for different functions and under different environmental change scenarios. Furthermore, the species needed to provide one function during multiple years were not the same as those needed to provide multiple functions within one year. Our results indicate that even more species will be needed to maintain ecosystem functioning and services than previously suggested by studies that have either (1) considered only the number of species needed to promote one function under one set of environmental conditions, or (2) separately considered the importance of biodiversity for providing ecosystem functioning across multiple years, places, functions or environmental change scenarios. Therefore, although species may appear functionally redundant when one function is considered under one set of environmental conditions, many species are needed to maintain multiple functions at multiple times and places in a changing world.

Constraints on food chain length arising from regional metacommunity dynamics.

Classical ecological theory has proposed several determinants of food chain length, but the role of metacommunity dynamics has not yet been fully considered. By modelling patchy predator-prey metacommunities with extinction-colonization dynamics, we identify two distinct constraints on food chain length. First, finite colonization rates limit predator occupancy to a subset of prey-occupied sites. Second, intrinsic extinction rates accumulate along trophic chains. We show how both processes concur to decrease maximal and average food chain length in metacommunities. This decrease is mitigated if predators track their prey during colonization (habitat selection) and can be reinforced by top-down control of prey vital rates (especially extinction). Moreover, top-down control of colonization and habitat selection can interact to produce a counterintuitive positive relationship between perturbation rate and food chain length. Our results show how novel limits to food chain length emerge in spatially structured communities. We discuss the connections between these constraints and the ones commonly discussed, and suggest ways to test for metacommunity effects in food webs.

Rapid exploiter-victim coevolution: the race is not always to the swift.

The modeling of coevolutionary races has traditionally been dominated by methods invoking a timescale separation between ecological and evolutionary dynamics, the latter assumed to be much slower than the former. Yet it is becoming increasingly clear that in many cases the two processes occur on similar timescales and that such "rapid" evolution can have profound implications for the dynamics of communities and ecosystems. After briefly reviewing the timescale separations most common in coevolution theory, we use a general model of exploiter-victim coevolution to confront predictions from slow-evolution analysis with Monte Carlo simulations. We show how rapid evolution radically alters the dynamics and outcome of coevolutionary arms races. In particular, a fast-evolving exploiter can enable victim diversification and thereby lose a race it is expected to win. We explain simulation results, using mathematical analysis with relaxed timescale separations. Unusual mutation parameters are not required, since rapid evolution naturally emerges from slow competitive exclusion. Our results point to interesting consequences of exploiter rapid evolution and experimentally testable patterns, while indicating that more attention should be paid to rapid evolution in evolutionary theory.