How life‐history traits affect ecosystem properties: effects of dispersal in meta‐ecosystems

The concept of life‐history traits and the study of these traits are the hallmark of population biology. Acknowledging their variability and evolution has allowed us to understand how species adapt in response to their environment. The same traits are also involved in how species alter ecosystems and shape their dynamics and functioning. Some theories, such as the metabolic theory of ecology, ecological stoichiometry or pace‐of‐life theory, already recognize this junction, but only do so in an implicitly non‐spatial context. Meanwhile, for a decade now, it has been argued that ecosystem properties have to be understood at a larger scale using meta‐ecosystem theory because source–sink dynamics, community assembly and ecosystem stability are all modified by spatial structure. Here, we argue that some ecosystem properties can be linked to a single life‐history trait, dispersal, i.e. the tendency of organisms to live, compete and reproduce away from their birth place. By articulating recent theoretical and empirical studies linking ecosystem functioning and dynamics to species dispersal, we aim to highlight both the known connections between life‐history traits and ecosystem properties and the unknown areas, which deserve further empirical and theoretical developments.     

Microsatellite data show evidence for male-biased dispersal in the Caribbean lizard Anolis roquet

Dispersal is a key component of an organism's life history and differences in dispersal between sexes appear to be widespread in vertebrates. However, most predictions of sex-biased dispersal have been based on observations of social structure in birds and mammals and more data are needed on other taxa to test whether these predictions apply in other organisms. Caribbean anole lizards are important model organisms in various biological disciplines, including evolutionary biology. However, very little is known about their dispersal strategies despite the importance of dispersal for population structure and dynamics. Here we use nine microsatellite markers to assess signatures of sex-biased dispersal on two spatial sampling scales in Anolis roquet, an anole endemic to the island of Martinique. Significantly higher gene diversity (H(S)) and lower mean assignment value (mAIC) was found in males on the larger spatial sampling scale. Significant heterozygote deficit (F(IS)), lower population differentiation (F(ST)), mAIC and variance of assignment index (vAIC) was found in males on the smaller spatial scale. The observation of male biased dispersal conform with expectations based on the polygynous mating system of Anolis roquet, and contributes to an explanation of the contrasting patterns of genetic structure between maternal and biparental markers that have been reported previously in this, and other anoline, species.     

Egg banks in freshwater zooplankton: evolutionary and ecological archives in the sediment

Many representatives of freshwater zooplankton produce at some stage in their life cycle resting stages. A variable portion of the eggs of the previous growing period will hatch at the next occasion while the remaining ones are added to a persistent egg bank, where they can remain viable for decades or longer. The importance of the study of resting eggs and egg banks in general for such different disciplines as taxonomy, ecological biogeography, paleolimnology, nature conservation, evolutionary ecology and community and population ecology is generally appreciated. The major current and expected future developments in this rapidly expanding field of research are presented here. The structure and dynamics of the egg bank are determined by the life history characteristics of the species (or local population), the hatching phenology of their resting stages, and the characteristics of the habitat. The horizontal distribution of dormant stages is generally patchy, with a greater density in the deeper and/or windward parts of a pond or lake. In sediment cores, most viable (responsive) eggs occur in the upper centimeters, although vertical variation related to the history of fish predation or water quality occurs. The accumulation of resting stages of different species, generations and genotypes with variable regeneration niches results in a mixed egg bank with greater potential biodiversity than the active community sampled at any one moment. Through the benthic–pelagic coupling, this dormant reservoir may have considerable impact on the evolutionary potential of the organisms, the ecological dynamics of the community and the distribution of species. Egg banks can be considered the archive of the local habitat, since the pattern of changes in species assemblage and genotypes from the past up to the present reflect changes due to natural or anthropogenic impact that can be used to reconstruct evolutionary processes or even to restore the local habitat. Overlooking the egg bank as an important component of zooplankton communities may lead to erroneous interpretations in the analysis of community and population genetic structure. This review integrates technical and scientific information needed in the study of the structure and function of egg banks in zooplankton with special focus on the fascinating latest developments in the field.     

Plant life history and above–belowground interactions: missing links

The importance of above–belowground interactions for plant growth and community dynamics became clear in the last decades, whereas the numerous studies on plant life history improved our knowledge on eco‐evolutionary dynamics. However, surprisingly few studies have linked both research fields despite their potential to increase our mechanistic understanding of how above belowground interactions are governed. Here I briefly review studies on above–belowground interactions and plant life history and identify important research gaps. To advance our understanding of ecological strategies and eco‐evolutionary dynamics of plants and their associated organisms it is warranted to elucidate the interconnectivity and tradeoffs of plant life history traits of growth, defence, reproduction, nutrient cycling and the functional composition of above‐ and belowground heterotrophic communities. Using the concept of tradeoffs in growth, reproduction and defence we can postulate that plants in rich soil grow, reproduce and die fast whilst avoiding above‐ and belowground antagonists, whereas plants in poor soil grow slow, live and reproduce longer and invest in above‐ and belowground mutualists and defences. However, alternative scenarios are possible and depend on the selection pressure by above‐ and belowground mutualists and antagonists during plant ontogeny and via after‐life effects. To elucidate missing links between life history traits and above–belowground interactions, complementary modelling and empirical studies are needed that reveal the coupling between below‐ and aboveground plant traits of growth, defence and reproduction, their heritability and their cost/benefit relation. These cost/benefit analyses of defence should span from individuals to future generations, taking feedback effects via altered biotic communities and resource competition into account. The role of soil fertility in steering plant life history traits requires explicit testing of trans‐generational trait shifts in growth, defence, reproduction, cost/benefit of associations with mutualists and antagonists and soil feedbacks across plant genotypes/species with distinct life history traits, grown across soil fertility gradients.     

The danger within: the role of genetic,behavioural and ecological factors in population persistence of colour polymorphic species

Polymorphic species have been the focus of important work in evolutionary biology. It has been suggested that colour polymorphic species have specific evolutionary and population dynamics that enable them to persist through environmental changes better than less variable species. We suggest that recent empirical and theoretical work indicates that polymorphic species may be more vulnerable to extinction than previously thought. This vulnerability arises because these species often have a number of correlated sexual, behavioural, life history and ecological traits, which can have a simple genetic underpinning. When exacerbated by environmental change, these alternate strategies can lead to conflict between morphs at the genomic and population levels, which can directly or indirectly affect population and evolutionary dynamics. In this perspective, we identify a number of ways in which the nature of the correlated traits, their underpinning genetic architecture, and the inevitable interactions between colour morphs can result in a reduction in population fitness. The principles illustrated here apply to all kinds of discrete polymorphism (e.g. behavioural syndromes), but we focus primarily on colour polymorphism because they are well studied. We urge further empirical investigation of the genetic architecture and interactions in polymorphic species to elucidate the impact on population fitness.     

The role of fish life histories in allometrically scaled food‐web dynamics

相似文献   

The importance of effective sampling for exploring the population dynamics of haploid–diploid seaweeds

The mating system partitions genetic diversity within and among populations and the links between life history traits and mating systems have been extensively studied in diploid organisms. As such most evolutionary theory is focused on species for which sexual reproduction occurs between diploid male and diploid female individuals. However, there are many multicellular organisms with biphasic life cycles in which the haploid stage is prolonged and undergoes substantial somatic development. In particular, biphasic life cycles are found across green, brown and red macroalgae. Yet, few studies have addressed the population structure and genetic diversity in both the haploid and diploid stages in these life cycles. We have developed some broad guidelines with which to develop population genetic studies of haploid‐diploid macroalgae and to quantify the relationship between power and sampling strategy. We address three common goals for studying macroalgal population dynamics, including haploid‐diploid ratios, genetic structure and paternity analyses.     

Spatial scaling of population synchrony in marine fish depends on their life history

The synchrony of population dynamics in space has important implications for ecological processes, for example affecting the spread of diseases, spatial distributions and risk of extinction. Here, we studied the relationship between spatial scaling in population dynamics and species position along the slow‐fast continuum of life history variation. Specifically, we explored how generation time, growth rate and mortality rate predicted the spatial scaling of abundance and yearly changes in abundance of eight marine fish species. Our results show that population dynamics of species' with ‘slow’ life histories are synchronised over greater distances than those of species with ‘fast’ life histories. These findings provide evidence for a relationship between the position of the species along the life history continuum and population dynamics in space, showing that the spatial distribution of abundance may be related to life history characteristics.     

It's about time: divergence, demography, and the evolution of developmental modes in marine invertebrates

Differences in larval developmental mode are predicted to affect ecological and evolutionary processes ranging from gene flow and population bottlenecks to rates of population recovery from anthropogenic disturbance and capacity for local adaptation. The most powerful tests of these predictions use comparisons among species to ask how phylogeographic patterns are correlated with the evolution and loss of prolonged planktonic larval development. An important and largely untested assumption of these studies is that interspecific differences in population genetic structure are mainly caused by differences in dispersal and gene flow (rather than by differences in divergence times among populations or changes in effective population sizes), and that species with similar patterns of spatial genetic variation have similar underlying temporal demographic histories. Teasing apart these temporal and spatial patterns is important for understanding the causes and consequences of evolutionary changes in larval developmental mode. New analytical methods that use the coalescent history of allelic diversity can reveal these temporal patterns, test the strength of traditional population-genetic explanations for variation in spatial structure based on differences in dispersal, and identify strongly supported alternative explanations for spatial structure based on demographic history rather than on gene flow alone. We briefly review some of these recent analytical developments, and show their potential for refining ideas about the correspondence between the evolution of larval developmental mode, population demographic history, and spatial genetic variation.     

Host–parasite determinants of parasite population structure: lessons from bats and mites on the importance of time

By definition, parasitic organisms are strongly dependant on their hosts, and for a great majority, this dependence includes host-to-host transmission. Constraints imposed by the host's spatial distribution and demography, in combination with those of the parasite, can lead to a metapopulation structure, where parasite populations are highly stochastic (i.e. prone to frequent extinctions and re-colonizations) and where drift becomes a major force shaping standing genetic variation. This, in turn, will directly affect the observed population structure, along with the ability of the parasite to adapt (or co-adapt) to its host. However, only a specific consideration of temporal dynamics can reveal the extent to which drift shapes parasite population structure; this is rarely taken into account in population genetic studies of parasitic organisms. The study by Bruyndonckx et al. in this issue of Molecular Ecology does just this and, in doing so, illustrates how a comparison of host–parasite co-structures in light of temporal dynamics can be particularly informative for understanding the ecological and evolutionary constraints imposed by the host. More specifically, the authors examine spatial and temporal population genetic data of a parasitic mite Spinturnix bechsteini that exclusively exploits the Bechstein's bat Myotis bechsteinii and consider these data in relation to host–parasite life histories and the population structure of the host.     

Post‐reproductive life span and demographic stability

Recent field studies suggest that it is common in nature for animals to outlive their reproductive viability. Post‐reproductive life span has been observed in a broad range of vertebrate and invertebrate species. But post‐reproductive life span poses a paradox for traditional theories of life history evolution. The only commonly‐cited explanation is the ‘grandmother hypothesis’, which is limited to higher, social mammals. We propose that post‐reproductive life span evolves to stabilize population dynamics, avoiding local extinctions. Predator–prey and other ecosystem interactions tend to produce volatility that can create population crashes and local extinctions. Total fertility rates that exceed the ecosystem's recovery rate contribute to population overshoot, followed by collapse. These local extinctions may constitute a potent group selection mechanism, driving evolution toward controlled rates of population growth, even when there is a significant individual cost. In this paper, we consider the question: what life history characteristics support demographic homeostasis at the least cost to individual fitness? In individual‐based evolutionary simulations, we find that reduction in fertility is sufficient to avoid population instabilities leading to extinction, but that life histories that include senescence can accomplish the same thing at a lower cost to individual fitness. Furthermore, life histories that include the potential for a post‐reproductive period are yet more efficient at stabilizing population dynamics, while minimizing the impact on individual fitness.     

Phylogentic constraints,adaptive syndromes,and emergent properties: From individuals to population dynamics

The hypothesis is developed that there are causal linkages in evolved insect herbivore life histories and behaviors from phylogenetic constraints to adaptive syndromes to the emergent properties involving ecological interactions and population dynamics. Thus the argument is developed that the evolutionary biology of a species predetermines its current ecology.Phylogenetic Constraints refer to old characters in the phylogeny of a species and a group of species which set limits on the range of life history patterns and behaviors that can evolve. For example, a sawfly is commonly limited to oviposition in soft plant tissue, while plants are growing rapidly.Adaptive Syndromes are evolutionary responses to the phylogenetic constraints that minimize the limitations and maximize larval performance. Such syndromes commonly involve details of female ovipositional behavior and how individuals make choices for oviposition sites relative to plant quality variation which maximize larval survival. Syndromes also involve larval adaptations to the kinds of choices females make in oviposition. The evolutionary biology involved with phylogenetic constraints and adaptive syndromes commonly predetermines the ecological interactions of a species and its population dynamics. Therefore, these ecological interactions are calledEmergent Properties because they are natural consequences of evolved morphology, behavior, and physiology. They commonly strongly influence the three-trophic-level interactions among host plants, insect herbivores, and carnivores, and the relative forces of bottom-up and top-down influences in food webs. The arguments are supported using such examples as galling sawflies and other gallers, shoot-boring moths and beetles, budworms, and forest Macrolepidoptera. The contrasts between outbreak or eruptive species and uncommon and rare species with latent population dynamics are emphasized.     

Phoretic dispersal influences parasite population genetic structure

Dispersal is a fundamental component of the life history of most species. Dispersal influences fitness, population dynamics, gene flow, genetic drift and population genetic structure. Even small differences in dispersal can alter ecological interactions and trigger an evolutionary cascade. Linking such ecological processes with evolutionary patterns is difficult, but can be carried out in the proper comparative context. Here, we investigate how differences in phoretic dispersal influence the population genetic structure of two different parasites of the same host species. We focus on two species of host‐specific feather lice (Phthiraptera: Ischnocera) that co‐occur on feral rock pigeons (Columba livia). Although these lice are ecologically very similar, “wing lice” (Columbicola columbae) disperse phoretically by “hitchhiking” on pigeon flies (Diptera: Hippoboscidae), while “body lice” (Campanulotes compar) do not. Differences in the phoretic dispersal of these species are thought to underlie observed differences in host specificity, as well as the degree of host–parasite cospeciation. These ecological and macroevolutionary patterns suggest that body lice should exhibit more genetic differentiation than wing lice. We tested this prediction among lice on individual birds and among lice on birds from three pigeon flocks. We found higher levels of genetic differentiation in body lice compared to wing lice at two spatial scales. Our results indicate that differences in phoretic dispersal can explain microevolutionary differences in population genetic structure and are consistent with macroevolutionary differences in the degree of host–parasite cospeciation.     

Confounding factors in the detection of species responses to habitat fragmentation

Habitat loss has pervasive and disruptive impacts on biodiversity in habitat remnants. The magnitude of the ecological impacts of habitat loss can be exacerbated by the spatial arrangement -- or fragmentation -- of remaining habitat. Fragmentation per se is a landscape-level phenomenon in which species that survive in habitat remnants are confronted with a modified environment of reduced area, increased isolation and novel ecological boundaries. The implications of this for individual organisms are many and varied, because species with differing life history strategies are differentially affected by habitat fragmentation. Here, we review the extensive literature on species responses to habitat fragmentation, and detail the numerous ways in which confounding factors have either masked the detection, or prevented the manifestation, of predicted fragmentation effects.Large numbers of empirical studies continue to document changes in species richness with decreasing habitat area, with positive, negative and no relationships regularly reported. The debate surrounding such widely contrasting results is beginning to be resolved by findings that the expected positive species-area relationship can be masked by matrix-derived spatial subsidies of resources to fragment-dwelling species and by the invasion of matrix-dwelling species into habitat edges. Significant advances have been made recently in our understanding of how species interactions are altered at habitat edges as a result of these changes. Interestingly, changes in biotic and abiotic parameters at edges also make ecological processes more variable than in habitat interiors. Individuals are more likely to encounter habitat edges in fragments with convoluted shapes, leading to increased turnover and variability in population size than in fragments that are compact in shape. Habitat isolation in both space and time disrupts species distribution patterns, with consequent effects on metapopulation dynamics and the genetic structure of fragment-dwelling populations. Again, the matrix habitat is a strong determinant of fragmentation effects within remnants because of its role in regulating dispersal and dispersal-related mortality, the provision of spatial subsidies and the potential mediation of edge-related microclimatic gradients.We show that confounding factors can mask many fragmentation effects. For instance, there are multiple ways in which species traits like trophic level, dispersal ability and degree of habitat specialisation influence species-level responses. The temporal scale of investigation may have a strong influence on the results of a study, with short-term crowding effects eventually giving way to long-term extinction debts. Moreover, many fragmentation effects like changes in genetic, morphological or behavioural traits of species require time to appear. By contrast, synergistic interactions of fragmentation with climate change, human-altered disturbance regimes, species interactions and other drivers of population decline may magnify the impacts of fragmentation. To conclude, we emphasise that anthropogenic fragmentation is a recent phenomenon in evolutionary time and suggest that the final, long-term impacts of habitat fragmentation may not yet have shown themselves.     

Dynamics of Adaptation in Spatially Heterogeneous Metapopulations

The selection pressure experienced by organisms often varies across the species range. It is hence crucial to characterise the link between environmental spatial heterogeneity and the adaptive dynamics of species or populations. We address this issue by studying the phenotypic evolution of a spatial metapopulation using an adaptive dynamics approach. The singular strategy is found to be the mean of the optimal phenotypes in each habitat with larger weights for habitats present in large and well connected patches. The presence of spatial clusters of habitats in the metapopulation is found to facilitate specialisation and to increase both the level of adaptation and the evolutionary speed of the population when dispersal is limited. By showing that spatial structures are crucial in determining the specialisation level and the evolutionary speed of a population, our results give insight into the influence of spatial heterogeneity on the niche breadth of species.     

A life‐history perspective on the demographic drivers of structured population dynamics in changing environments

Current understanding of life‐history evolution and how demographic parameters contribute to population dynamics across species is largely based on assumptions of either constant environments or stationary environmental variation. Meanwhile, species are faced with non‐stationary environmental conditions (changing mean, variance, or both) created by climate and landscape change. To close the gap between contemporary reality and demographic theory, we develop a set of transient life table response experiments (LTREs) for decomposing realised population growth rates into contributions from specific vital rates and components of population structure. Using transient LTREs in a theoretical framework, we reveal that established concepts in population biology will require revision because of reliance on approaches that do not address the influence of unstable population structure on population growth and mean fitness. Going forward, transient LTREs will enhance understanding of demography and improve the explanatory power of models used to understand ecological and evolutionary dynamics.     

Dispersal: a central and independent trait in life history

The study of tradeoffs among major life history components (age at maturity, lifespan and reproduction) allowed the development of a quantitative framework to understand how environmental variation shapes patterns of biodiversity among and within species. Because every environment is inherently spatially structured, and in most cases temporally variable, individuals need to move within and among habitats to maximize fitness. Dispersal is often assumed to be tightly integrated into life histories through genetic correlations with other vital traits. This assumption is particularly strong within the context of a fast‐slow continuum of life‐history variation. Such a framework is to date used to explain many aspects of population and community dynamics. Evidence for a consistent and context‐independent integration of dispersal in life histories is, however, weak. We therefore advocate the explicit integration of dispersal into life history theory as a principal axis of variation influencing fitness, that is free to evolve, independently of other life history traits. We synthesize theoretical and empirical evidence on the central role of dispersal and its evolutionary dynamics on the spatial distribution of ecological strategies and its impact on population spread, invasions and coexistence. By applying an optimality framework we show that the inclusion of dispersal as an independent dimension of life histories might substantially change our view on evolutionary trajectories in spatially structured environments. Because changes in the spatial configuration of habitats affect the costs of movement and dispersal, adaptations to reduce these costs will increase phenotypic divergence among and within populations. We outline how this phenotypic heterogeneity is anticipated to further impact population and community dynamics.     

Restoration Biology: A Population Biology Perspective

A major goal of population biologists involved in restoration work is to restore populations to a level that will allow them to persist over the long term within a dynamic landscape and include the ability to undergo adaptive evolutionary change. We discuss five research areas of particular importance to restoration biology that offer potentially unique opportunities to couple basic research with the practical needs of restorationists. The five research areas are: (1) the influence of numbers of individuals and genetic variation in the initial population on population colonization, establishment, growth, and evolutionary potential; (2) the role of local adaptation and life history traits in the success of restored populations; (3) the influence of the spatial arrangement of landscape elements on metapopulation dynamics and population processes such as migration; (4) the effects of genetic drift, gene flow, and selection on population persistence within an often accelerated, successional time frame; and (5) the influence of interspecific interactions on population dynamics and community development. We also provide a sample of practical problems faced by practitioners, each of which encompasses one or more of the research areas discussed, and that may be solved by addressing fundamental research questions.     

Evolutionary dynamics in structured populations

Evolutionary dynamics shape the living world around us. At the centre of every evolutionary process is a population of reproducing individuals. The structure of that population affects evolutionary dynamics. The individuals can be molecules, cells, viruses, multicellular organisms or humans. Whenever the fitness of individuals depends on the relative abundance of phenotypes in the population, we are in the realm of evolutionary game theory. Evolutionary game theory is a general approach that can describe the competition of species in an ecosystem, the interaction between hosts and parasites, between viruses and cells, and also the spread of ideas and behaviours in the human population. In this perspective, we review the recent advances in evolutionary game dynamics with a particular emphasis on stochastic approaches in finite sized and structured populations. We give simple, fundamental laws that determine how natural selection chooses between competing strategies. We study the well-mixed population, evolutionary graph theory, games in phenotype space and evolutionary set theory. We apply these results to the evolution of cooperation. The mechanism that leads to the evolution of cooperation in these settings could be called ‘spatial selection’: cooperators prevail against defectors by clustering in physical or other spaces.