The evolutionary ecology of interactive synchronism: The illusion of the optimal phenotype

In this article, we discuss some ecological-evolutionary strategies that allow synchronization of organisms, resources, and conditions. Survival and reproduction require synchronization of life cycles of organisms with favourable environmental and ecological features and conditions. This interactive synchronization can occur directly, through pairwise or diffuse co-evolution, or indirectly, for example, as a result of actions of ecosystem engineers and facilitator species. Observations of specific interactions, especially those which have coevolved, may give the false impression that evolution results in optimal genotypes or phenotypes. However, some phenotypes may arise under evolutionary constraints, such as simultaneous evolution of multiple traits, lack of a chain of fit transitional forms leading to an optimal phenotype, or by limits inherent in the process of selection, set by the number of selective deaths and by interference between linked variants. Although there are no optimal phenotypes, optimization models applied to particular species may be useful for a better understanding of the nature of adaptations. The evolution of adaptive strategies results in variable life histories. These strategies can minimize adverse impacts on the fitness of extreme or severe environmental conditions on survival and reproduction, and may include reproductive strategies such as semelparity and iteroparity, or morphological, physiological, or behavioural traits such as diapause, seasonal polyphenism, migration, or bet-hedging. However, natural selection cannot indefinitely maintain intra-population variation, and lack of variation can ultimately extinguish populations

Extinction risk and eco-evolutionary dynamics in a variable environment with increasing frequency of extreme events

One of the most dramatic consequences of climate change will be the intensification and increased frequency of extreme events. I used numerical simulations to understand and predict the consequences of directional trend (i.e. mean state) and increased variability of a climate variable (e.g. temperature), increased probability of occurrence of point extreme events (e.g. floods), selection pressure and effect size of mutations on a quantitative trait determining individual fitness, as well as the their effects on the population and genetic dynamics of a population of moderate size. The interaction among climate trend, variability and probability of point extremes had a minor effect on risk of extinction, time to extinction and distribution of the trait after accounting for their independent effects. The survival chances of a population strongly and linearly decreased with increasing strength of selection, as well as with increasing climate trend and variability. Mutation amplitude had no effects on extinction risk, time to extinction or genetic adaptation to the new climate. Climate trend and strength of selection largely determined the shift of the mean phenotype in the population. The extinction or persistence of the populations in an 'extinction window' of 10 years was well predicted by a simple model including mean population size and mean genetic variance over a 10-year time frame preceding the 'extinction window', although genetic variance had a smaller role than population size in predicting contemporary risk of extinction. © 2014 The Author(s) Published by the Royal Society

Is Bacterial Persistence a Social Trait?

The ability of bacteria to evolve resistance to antibiotics has been much reported in recent years. It is less well-known that within populations of bacteria there are cells which are resistant due to a non-inherited phenotypic switch to a slow-growing state. Although such ‘persister’ cells are receiving increasing attention, the evolutionary forces involved have been relatively ignored. Persistence has a direct benefit to cells because it allows survival during catastrophes–a form of bet-hedging. However, persistence can also provide an indirect benefit to other individuals, because the reduced growth rate can reduce competition for limiting resources. This raises the possibility that persistence is a social trait, which can be influenced by kin selection. We develop a theoretical model to investigate the social consequences of persistence. We predict that selection for persistence is increased when: (a) cells are related (e.g. a single, clonal lineage); and (b) resources are scarce. Our model allows us to predict how the level of persistence should vary with time, across populations, in response to intervention strategies and the level of competition. More generally, our results clarify the links between persistence and other bet-hedging or social behaviours

The Stabilizing Effect Of Intraspecific Genetic Variation On Population Dynamics In Novel And Ancestral Habitats

Recent studies show that intraspecific genetic variation in asexual species may have large effects on community and ecosystem functions, increasing their stability, productivity, and species richness. However, major questions regarding its population-level impact remain empirically unanswered: (a) How does intraspecific genetic diversity affect the ecological characteristics of sexual species, in which recombination can alter the outcome of causal mechanisms such as selection and niche diversification? (b) Does genetic diversity increase population dynamic stability? (c) Is the impact of genetic diversity dependent on the selective environment? To answer these questions, I founded replicate flour beetle (Tribolium castaneum) populations with different degrees of ecologically relevant, heritable trait variation and monitored their dynamics for approximately eight generations. I show that population stability and persistence increased with greater genetic variation but that the stabilizing effect was independent of the selective habitat (different proportions of ancestral and novel resources). Alleles from a single founding strain underwent a selective sweep in the homogeneous ancestral habitat but not in a novel heterogeneous habitat. These results expand current understanding of the ecological impacts of genetic diversity by showing that genetically more diverse sexual populations persist longer and are more stable but that the selective environment determines the mechanistic basis of increased stability.Integrative Biolog

The cultural evolution of age-at-marriage norms

We present an agent-based model designed to study the cultural evolution of age-at-marriage norms. We review theoretical arguments and empirical evidence on the existence of norms proscribing marriage outside of an acceptable age interval. Using a definition of norms as constraints built in agents, we model the transmission of norms, and of mechanisms of intergenerational transmission of norms. Agents can marry each other only if they share part of the acceptable age interval. We perform several simulation experiments on the evolution across generations. In particular, we study the conditions under which norms persist in the long run, the impact of initial conditions, the role of random mutations, and the impact of social influence. Although the agent-based model we use is highly stylized, it gives important insights on the societal-level dynamics of life-course norms.

Modeling the ecology and evolution of biodiversity: Biogeographical cradles, museums, and graves

Individual processes shaping geographical patterns of biodiversity are increasingly understood, but their complex interactions on broad spatial and temporal scales remain beyond the reach of analytical models and traditional experiments. To meet this challenge, we built a spatially explicit, mechanistic simulation model implementing adaptation, range shifts, fragmentation, speciation, dispersal, competition, and extinction, driven by modeled climates of the past 800,000 years in South America. Experimental topographic smoothing confirmed the impact of climate heterogeneity on diversification. The simulations identified regions and episodes of speciation (cradles), persistence (museums), and extinction (graves). Although the simulations had no target pattern and were not parameterized with empirical data, emerging richness maps closely resembled contemporary maps for major taxa, confirming powerful roles for evolution and diversification driven by topography and climate

Plant hydraulic traits reveal islands as refugia from worsening drought.

Relatively mesic environments within arid regions may be important conservation targets as 'climate change refugia' for species persistence in the face of worsening drought conditions. Semi-arid southern California and the relatively mesic environments of California's Channel Islands provide a model system for examining drought responses of plants in potential climate change refugia. Most methods for detecting refugia are focused on 'exposure' of organisms to certain abiotic conditions, which fail to assess how local adaptation or acclimation of plant traits (i.e. 'sensitivity') contribute to or offset the benefits of reduced exposure. Here, we use a comparative plant hydraulics approach to characterize the vulnerability of plants to drought, providing a framework for identifying the locations and trait patterns that underlie functioning climate change refugia. Seasonal water relations, xylem hydraulic traits and remotely sensed vegetation indices of matched island and mainland field sites were used to compare the response of native plants from contrasting island and mainland sites to hotter droughts in the early 21st century. Island plants experienced more favorable water relations and resilience to recent drought. However, island plants displayed low plasticity/adaptation of hydraulic traits to local conditions, which indicates that relatively conserved traits of island plants underlie greater hydraulic safety and localized buffering from regional drought conditions. Our results provide an explanation for how California's Channel Islands function as a regional climate refugia during past and current climate change and demonstrate a physiology-based approach for detecting potential climate change refugia in other systems

Ecological boundaries and constraints on viable eco-evolutionary pathways

Evolutionary dynamics are subject to constraints ranging from limitations on what is physically possible to limitations on the pathways that evolution can take. One set of evolutionary constraints, known as ‘demographic constraints’, constrain what can occur evolutionarily due to the demographic or dynamical consequences of evolution leading to conditions that make populations susceptible to extinction. These demographic constraints can limit the strength of selection or the rates of environmental change populations can experience while remaining extant and the trait values a population can express. Here we further hypothesize that the population demographic and dynamic consequences of evolution also can constrain the eco-evolutionary pathways that populations can traverse by defining ecological boundaries represented by areas of likely extinction. We illustrate this process using a model of predator evolution. Our results show that the populations that persist over time tend to be those whose eco-evolutionary dynamics have avoided ecological boundaries representing areas of likely extinction due to stochastic deviations from a deterministic eco-evolutionary expectation. We term this subset of persisting pathways viable eco-evolutionary pathways. The potential existence of ecological boundaries constraining evolutionary pathways has important implications for predicting evolutionary dynamics, interpreting past evolution, and understanding the role of stochasticity and ecological constraints on eco-evolutionary dynamics

Ecological boundaries and constraints on viable eco-evolutionary pathways

Evolutionary dynamics are subject to constraints ranging from limitations on what is physically possible to limitations on the pathways that evolution can take. One set of evolutionary constraints, known as ‘demographic constraints’, constrain what can occur evolutionarily due to the demographic or dynamical consequences of evolution leading to conditions that make populations susceptible to extinction. These demographic constraints can limit the strength of selection or the rates of environmental change populations can experience while remaining extant and the trait values a population can express. Here we further hypothesize that the population demographic and dynamic consequences of evolution also can constrain the eco-evolutionary pathways that populations can traverse by defining ecological boundaries represented by areas of likely extinction. We illustrate this process using a model of predator evolution. Our results show that the populations that persist over time tend to be those whose eco-evolutionary dynamics have avoided ecological boundaries representing areas of likely extinction due to stochastic deviations from a deterministic eco-evolutionary expectation. We term this subset of persisting pathways viable eco-evolutionary pathways. The potential existence of ecological boundaries constraining evolutionary pathways has important implications for predicting evolutionary dynamics, interpreting past evolution, and understanding the role of stochasticity and ecological constraints on eco-evolutionary dynamics