Niche construction through phenological plasticity: life history dynamics and ecological consequences

The ability of an organism to alter the environment that it experiences has been termed 'niche construction'. Plants have several ways whereby they can determine the environment to which they are exposed at different life stages. This paper discusses three of these: plasticity in dispersal, flowering timing and germination timing. It reviews pathways through which niche construction alters evolutionary and ecological trajectories by altering the selective environment to which organisms are exposed, the phenotypic expression of plastic characters, and the expression of genetic variation. It provides examples whereby niche construction creates positive or negative feedbacks between phenotypes and environments, which in turn cause novel evolutionary constraints and novel life-history expression.     

Joint evolution of sex ratio and dispersal: conditions for higher dispersal rates from good habitats

We analyze models of evolution of sex ratio conditional on habitat quality and with sex specific dispersal. Previous analysis concluded that the main constraint on sex ratio is habitat choice and leads to overproduction of the most dispersing sex in low quality habitat. Here, we analyze three models with finite local populations and show that constraints on sex ratio can balance constraints on habitat choice. In the first model, dispersal rates are fixed. In the second, sex specific dispersal can evolve independently of the habitat quality. These models suggests that sex ratio evolution can lead to higher global dispersal rates (mean of male and female dispersal rates) from high quality habitats. In the last model dispersal evolves conditionally with both sex and habitat. Our models suggests that conditions for overproduction of the most dispersing sex in high quality habitat are frequent. The predictions of the models with evolving dispersal contrast with patterns generally described in nature. We discuss possible reasons of this difference.     

Forecasting ecological and evolutionary strategies to global change: an example from habitat selection by lemmings

Ecologists and evolutionary biologists must develop theories that can predict the consequences of global warming and other impacts on Earth's biota. Theories of adaptive habitat selection are particularly promising because they link distribution and density with fitness. The evolutionarily stable strategy that emerges from adaptive habitat choice is given by the system's habitat isodar, the graph of densities in pairs of habitats such that the expectation of fitness is the same in each. We illustrate how isodars can be converted into adaptive landscapes of habitat selection that display the density‐ and frequency‐dependent fitness of competing strategies of habitat use. The adaptive landscape varies with the abundance of habitats and can thus be used to predict future adaptive distributions of individuals under competing scenarios of habitat change. Application of the theory to three species of Arctic rodents living on Herschel Island in the Beaufort Sea predicts changes in selection gradients as xeric upland increases in frequency with global warming. Selection gradients will become more shallow for brown lemming (Lemmus trimucronatus) and tundra vole (Microtus oeconomus) strategies that preferentially exploit mesic habitat. Climate change will cause selection gradients for the alternative strategy of using mostly xeric habitat to become much steeper. Meanwhile, the adaptive landscape for collared lemmings (Dicrostonyx groenlandicus), which specialize on xeric Dryas‐covered upland, will become increasingly convex. Changes in the adaptive landscapes thus predict expanding niches for Lemmus and Microtus, and a narrower niche for Dicrostonyx. The ability to draw adaptive landscapes from current patterns of distribution represents one of the few methods available to forecast the consequences of climate change on the future distribution and evolution of affected species.     

Modelling single nucleotide effects in phosphoglucose isomerase on dispersal in the Glanville fritillary butterfly: coupling of ecological and evolutionary dynamics

Dispersal comprises a complex life-history syndrome that influences the demographic dynamics of especially those species that live in fragmented landscapes, the structure of which may in turn be expected to impose selection on dispersal. We have constructed an individual-based evolutionary sexual model of dispersal for species occurring as metapopulations in habitat patch networks. The model assumes correlated random walk dispersal with edge-mediated behaviour (habitat selection) and spatially correlated stochastic local dynamics. The model is parametrized with extensive data for the Glanville fritillary butterfly. Based on empirical results for a single nucleotide polymorphism (SNP) in the phosphoglucose isomerase (Pgi) gene, we assume that dispersal rate in the landscape matrix, fecundity and survival are affected by a locus with two alleles, A and C, individuals with the C allele being more mobile. The model was successfully tested with two independent empirical datasets on spatial variation in Pgi allele frequency. First, at the level of local populations, the frequency of the C allele is the highest in newly established isolated populations and the lowest in old isolated populations. Second, at the level of sub-networks with dissimilar numbers and connectivities of patches, the frequency of C increases with decreasing network size and hence with decreasing average metapopulation size. The frequency of C is the highest in landscapes where local extinction risk is high and where there are abundant opportunities to establish new populations. Our results indicate that the strength of the coupling of the ecological and evolutionary dynamics depends on the spatial scale and is asymmetric, demographic dynamics having a greater immediate impact on genetic dynamics than vice versa.     

How landscapes affect the evolution of dispersal behaviour in reef fishes: results from an individual‐based model

The vast majority of tropical reef fishes have a sedentary adult phase and pelagic larval phase that is potentially highly dispersive. Dispersal may be favoured by a wide range of factors including the arrangement of suitable habitat in space. In this paper the dispersal strategy of individuals is followed and allowed to evolve in a simplified model of three different landscapes: an enclosed sea, an open archipelago and a barrier reef. The three landscapes have very different characteristics, but all have similar spatial clumping of reef habitat. In all landscapes, as minimum time to settlement increases, evolved movement strategy also increases and longer settlement windows favour dispersal. In the archipelago movement is not maximized until the minimum pelagic duration is longer than in the other landscapes. The model predicts that, given the same pelagic duration, species from enclosed seas should have more dispersive behaviours than those from open archipelagos, because of the density of habitat and the aggregation of habitat in space affect the likelihood of larvae finding suitable habitat for settlement.     

Individual variation and population dynamics: lessons from a simple system

The mapping of environment, through variation in individuals' life histories, to dynamics can be complex and often poorly known. Consequently, it is not clear how important it is dynamically. To explore this, I incorporated lessons from an empirical system, a soil mite, into an individual-based model. Individuals compete for resource and allocate this according to eight 'genetic' rules that specify investment in growth or reserves (which influences survival or fecundity), size at maturation and reproductive allocation. Density dependence, therefore, emerges from competition for food, limiting individual's growth and fecundity. We use this model to examine the role that genetic and phenotypically plastic variation plays in dynamics, by fixing phenotypes, by allowing phenotypes to vary plastically and by creating genetic variation between individuals. Variation, and how it arises, influences short- and long-run dynamics in a way comparable in magnitude with halving food supply. In particular, by switching variation on and off, it is possible to identify a range of processes necessary to capture the dynamics of the 'full model'. Exercises like this can help identify key processes and parameters, but a concerted effort is needed across many different systems to search for shared understanding of both process and modelling.     

Experimentally induced host‐shift changes life‐history strategy in a seed beetle

Expansion of the host range in phytophagous insects depends on their ability to form an association with a novel plant through changes in host‐related traits. Phenotypic plasticity has important effects on initial survival of individuals faced with a new plant, as well as on the courses of evolutionary change during long‐term adaptation to novel conditions. Using experimental populations of the seed beetle that evolved on ancestral (common bean) or novel (chickpea) host and applying reciprocal transplant at both larval and adult stage on the alternative host plant, we studied the relationship between the initial (plastic) phases of host‐shift and the subsequent stages of evolutionary divergence in life‐history strategies between populations exposed to the host‐shift process. After 48 generations, populations became well adapted to chickpea by evolving the life‐history strategy with prolonged larval development, increased body mass, earlier reproduction, shorter lifespan and decreased plasticity of all traits compared with ancestral conditions. In chickpea‐adapted beetles, negative fitness consequences of low plasticity of pre‐adult development (revealed as severe decrease in egg‐to‐adult viability on beans) exhibited mismatch with positive effects of low plasticity (i.e. low host sensitivity) in oviposition and fecundity. In contrast, beetles adapted to the ancestral host showed high plasticity of developmental process, which enabled high larval survival on chickpea, whereas elevated plasticity in adult behaviour (i.e. high host sensitivity) resulted in delayed reproduction and decreased fecundity on chickpea. The analysis of population growth parameters revealed significant fluctuation during successive phases of the host‐shift process in A. obtectus.     

Population responses to anthropogenic disturbance: lessons from three‐spined sticklebacks Gasterosteus aculeatus in eutrophic habitats

Human‐induced environmental changes differ from most natural changes in which they happen at a faster rate and require quicker responses from populations. The first response of populations is usually phenotypically plastic alterations of morphology, physiology and behaviour. This plasticity can be favourable and move the population closer to an adaptive peak in the altered environment and, hence, maintain a viable population, or be maladaptive and move the population further from the peak and increase the risk of extinction. The radiation of the three‐spined stickleback Gasterosteus aculeatus from the ocean to different freshwater habitats has provided much information on adaptation to new environmental conditions. Currently, human‐induced eutrophication is changing the breeding areas of these fish, which creates a model system for investigation of responses to rapid environmental disturbance. Results show that a primary reaction is plastic alterations of behaviour, with some adjustments being adaptive while others are not. At the same time, the strength of sexual selection on several traits is relaxed, which could increase the relative importance of survival selection. Whether this will restore population viability depends on the amount of standing genetic variation in the right direction. Human disturbances can be dramatic and resolution of the limit of flexibility and the possibility of genetic adaptation should be important targets of future research.     

Morphological flexibility across an environmental gradient in the epiphytic orchid,Tolumnia variegata: complicating patterns of fitness

Deceit‐pollinated orchid species show substantial variation in floral traits, which may be maintained by genetic drift or various forms of selection, or may reflect phenotypic plasticity. We explored how much plasticity occurs in both vegetative and floral traits of Tolumnia variegata (Oncidiinae, Orchidaceae) across two different light environments in Puerto Rico using data from a reciprocal transplant experiment. We also examined how fruit set, a measure of reproductive success and a surrogate for fitness, is associated with this morphological variation, and whether it changes over time. Tolumnia variegata responded to environmental variables in multiple ways. Vegetative characters were more plastic than those associated with sexual reproduction. Transplant effects accounted for significant variation in flower length, lip length, number of inflorescences, peduncle length, leaf length and the total number of ramets, but responses were not always consistent among years. Phenotypic selection on morphological characters was dependent on plant location. The trends detected were complex, and often inconsistent across years, probably as a result of wetter and drier years than average. Overall fruit set was quite variable among plants, averaging 15%, with no significant differences among sun and shade plants. Although reproductive success was similar among sites, habitat heterogeneity and annual variation had an effect on morphological expression, which sometimes modified the trajectories of phenotypic selection. © 2010 The Linnean Society of London, Botanical Journal of the Linnean Society, 2010, 163 , 431–446.     

A way forward with eco evo devo: an extended theory of resource polymorphism with postglacial fishes as model systems

A major goal of evolutionary science is to understand how biological diversity is generated and altered. Despite considerable advances, we still have limited insight into how phenotypic variation arises and is sorted by natural selection. Here we argue that an integrated view, which merges ecology, evolution and developmental biology (eco evo devo) on an equal footing, is needed to understand the multifaceted role of the environment in simultaneously determining the development of the phenotype and the nature of the selective environment, and how organisms in turn affect the environment through eco evo and eco devo feedbacks. To illustrate the usefulness of an integrated eco evo devo perspective, we connect it with the theory of resource polymorphism (i.e. the phenotypic and genetic diversification that occurs in response to variation in available resources). In so doing, we highlight fishes from recently glaciated freshwater systems as exceptionally well‐suited model systems for testing predictions of an eco evo devo framework in studies of diversification. Studies on these fishes show that intraspecific diversity can evolve rapidly, and that this process is jointly facilitated by (i) the availability of diverse environments promoting divergent natural selection; (ii) dynamic developmental processes sensitive to environmental and genetic signals; and (iii) eco evo and eco devo feedbacks influencing the selective and developmental environments of the phenotype. We highlight empirical examples and present a conceptual model for the generation of resource polymorphism – emphasizing eco evo devo, and identify current gaps in knowledge.     

Female choice in Drosophila: evidence from Hawaii and implications for evolutionary biology

Details of female choice of mate in Drosophila silvestris of Hawaii strikingly parallels epigamic behavioral systems in many other animals and may be common in other species of Drosophilidae. Females respond selectively to male circling, wing displays, songs and tactile stimulation with foreleg cilia, a quantitative character that is highly variable in some populations. I hypothesize that the female can exert choice based on these cues from individual males that differ genetically by quantitative trait loci. Laboratory tests show that one third of courting males are repeatedly rejected in favor of a minority of alpha males. This result imposes non-random mating at the local population level. Past multiple-choice lab tests, widely used to measure isolation between pairs of populations or species of Drosophila may be flawed, since random mating has been assumed in the interpretation of results. Pre-mating sexual selection is clearly a powerful intrapopulation force in population biology. This view creates difficulties for discerning any proposed simultaneous interpopulation selective events in the presence of strong female choice. The long-held theory assuming that there is significant selection for pre-mating isolation between groups is questionable.     

Resurrecting complexity: the interplay of plasticity and rapid evolution in the multiple trait response to strong changes in predation pressure in the water flea Daphnia magna

A resurrection ecology reconstruction of 14 morphological, life history and behavioural traits revealed that a natural Daphnia magna population rapidly tracked changes in fish predation by integrating phenotypic plasticity and widespread evolutionary changes both in mean trait values and in trait plasticity. Increased fish predation mainly generated rapid adaptive evolution of plasticity (especially in the presence of maladaptive ancestral plasticity) resulting in an important change in the magnitude and direction of the multivariate reaction norm. Subsequent relaxation of the fish predation pressure resulted in reversed phenotypic plasticity and mainly caused evolution of the trait means towards the ancestral pre‐fish means. Relaxation from fish predation did, however, not result in a complete reversal to the ancestral fishless multivariate phenotype. Our study emphasises that the study population rapidly tracked environmental changes through a mosaic of plasticity, evolution of trait means and evolution of plasticity to generate integrated phenotypic changes in multiple traits.     

Functional mechanisms of an inducible defence in tadpoles: morphology and behaviour influence mortality risk from predation

In many amphibian larvae a suite of morphological and behavioural characters varies together in an induced defence against predators, but it remains unclear which features are functionally related to defence. We independently manipulated behaviour and morphology in tadpoles of Hyla versicolor and assessed their consequences for swimming performance and predator escape. Data on burst swimming showed that tadpoles which accelerated rapidly were elongate, with shallow bodies and tails. Predator escape was measured by exposing tadpoles to predators (larval Anax dragonflies or larval Ambystoma salamanders) and recording time until death. Tadpoles were first reared for 30 days in ponds containing either caged Anax or no predators; individuals responded to predators by developing large brightly coloured tails and short bodies. We placed tadpoles of both morphological phenotypes into plastic tubs, and manipulated their behaviour using food and chemical cues from predators. Mortality risk experienced by the predator‐induced phenotype was about half that of the no‐predator phenotype, and risk increased with time spent swimming. An interaction between morphology and behaviour arose because increasing activity caused higher risk for tadpoles with deep tail fins but not shallow tail fins.     

Parallel invasions produce heterogenous patterns of life history adaptation: rapid divergence in an invasive insect

Biotic invasions provide a natural experiment in evolution: when invasive species colonize new ranges, they may evolve new clines in traits in response to environmental gradients. Yet it is not clear how rapidly such patterns can evolve and whether they are consistent between regions. We compare four populations of the invasive cabbage white butterfly (Pieris rapae) from North America and Japan, independently colonized by P. rapae 150 years ago and 300 years ago, respectively. On each continent, we employed a northern and southern population to compare the effects of latitude on body mass, development rate and immune function. For each population, we used a split‐sibling family design in which siblings were reared at either warm (26.7 °C) or cool (20 °C) temperatures to determine reaction norms for each trait. Latitudinal patterns in development time were similar between the two continents. In contrast, there were strong geographical differences in reaction norms for body size, but no consistent effects of latitude; there were no detectable effects of latitude or continent on immune function. These results imply that some life history traits respond consistently to selection along climatic gradients, whereas other traits may respond to local environmental factors, or not at all.     

(Not) far from home: No sex bias in dispersal,but limited genetic patch size,in an endangered species,the Spotted Turtle (Clemmys guttata)

Sex-biased dispersal is common in many animals, with male-biased dispersal often found in studies of mammals and reptiles, including interpretations of spatial genetic structure, ostensibly as a result of male–male competition and a lack of male parental care. Few studies of sex-biased dispersal have been conducted in turtles, but a handful of studies, in saltwater turtles and in terrestrial turtles, have detected male-biased dispersal as expected. We tested for sex-biased dispersal in the endangered freshwater turtle, the spotted turtle (Clemmys guttata) by investigating fine-scale genetic spatial structure of males and females. We found significant spatial genetic structure in both sexes, but the patterns mimicked each other. Both males and females typically had higher than expected relatedness at distances <25 km, and in many distance classes greater than 25 km, less than expected relatedness. Similar patterns were apparent whether we used only loci in Hardy–Weinberg equilibrium (n = 7) or also included loci with potential null alleles (n = 5). We conclude that, contrary to expectations, sex-biased dispersal is not occurring in this species, possibly related to the reverse sexual dimorphism in this species, with females having brighter colors. We did, however, detect significant spatial genetic structure in males and females, separate and combined, showing philopatry within a genetic patch size of <25 km in C. guttata, which is concerning for an endangered species whose populations are often separated by distances greater than the genetic patch size.     

Company matters: The presence of other genotypes alters traits and intraspecific selection in an Arctic diatom under climate change

Arctic phytoplankton and their response to future conditions shape one of the most rapidly changing ecosystems on the planet. We tested how much the phenotypic responses of strains from the same Arctic diatom population diverge and whether the physiology and intraspecific composition of multistrain populations differs from expectations based on single strain traits. To this end, we conducted incubation experiments with the diatom Thalassiosira hyalina under present‐day and future temperature and pCO₂ treatments. Six fresh isolates from the same Svalbard population were incubated as mono‐ and multistrain cultures. For the first time, we were able to closely follow intraspecific selection within an artificial population using microsatellites and allele‐specific quantitative PCR. Our results showed not only that there is substantial variation in how strains of the same species cope with the tested environments but also that changes in genotype composition, production rates, and cellular quotas in the multistrain cultures are not predictable from monoculture performance. Nevertheless, the physiological responses as well as strain composition of the artificial populations were highly reproducible within each environment. Interestingly, we only detected significant strain sorting in those populations exposed to the future treatment. This study illustrates that the genetic composition of populations can change on very short timescales through selection from the intraspecific standing stock, indicating the potential for rapid population level adaptation to climate change. We further show that individuals adjust their phenotype not only in response to their physicochemical but also to their biological surroundings. Such intraspecific interactions need to be understood in order to realistically predict ecosystem responses to global change.