Beyond simple adaptation: Incorporating other evolutionary processes and concepts into eco-evolutionary dynamics

Studies of eco-evolutionary dynamics have integrated evolution with ecological processes at multiple scales (populations, communities and ecosystems) and with multiple interspecific interactions (antagonistic, mutualistic and competitive). However, evolution has often been conceptualised as a simple process: short-term directional adaptation that increases population growth. Here we argue that diverse other evolutionary processes, well studied in population genetics and evolutionary ecology, should also be considered to explore the full spectrum of feedback between ecological and evolutionary processes. Relevant but underappreciated processes include (1) drift and mutation, (2) disruptive selection causing lineage diversification or speciation reversal and (3) evolution driven by relative fitness differences that may decrease population growth. Because eco-evolutionary dynamics have often been studied by population and community ecologists, it will be important to incorporate a variety of concepts in population genetics and evolutionary ecology to better understand and predict eco-evolutionary dynamics in nature.     

Recent advances in ecological stoichiometry: insights for population and community ecology

Conventional theories of population and community dynamics are based on a single currency such as number of individuals, biomass, carbon or energy. However, organisms are constructed of multiple elements and often require them (in particular carbon, phosphorus and nitrogen) in different ratios than provided by their resources; this mismatch may constrain the net transfer of energy and elements through trophic levels. Ecological stoichiometry, the study of the balance of elements in ecological processes, offers a framework for exploring ecological effects of such constraints. We review recent theoretical and empirical studies that have considered how stoichiometry may affect population and community dynamics. These studies show that stoichiometric constraints can affect several properties of populations (e.g. stability, oscillations, consumer extinction) and communities (e.g. coexistence of competitors, competitive interactions between different guilds). We highlight gaps in general knowledge and focus on areas of population and community ecology where incorporation of stoichiometric constraints may be particularly fruitful, such as studies of demographic bottlenecks, spatial processes, and multi-species interactions. Finally, we suggest promising directions for new research by recommending potential study systems (terrestrial insects, detritivory-based webs, soil communities) to improve our understanding of populations and communities. Our conclusion is that a better integration of stoichiometric principles and other theoretical approaches in ecology may allow for a richer understanding of both population and community structure and dynamics.     

The effect of dilution on eco‐evolutionary dynamics of experimental microbial communities

Changing environmental conditions can infer structural modifications of predator‐prey communities. New conditions often increase mortality which reduces population sizes. Following this, predation pressure may decrease until populations are dense again. Dilution may thus have substantial impact not only on ecological but also on evolutionary dynamics because it amends population densities. Experimental studies, in which microbial populations are maintained by a repeated dilution into fresh conditions after a certain period, are extensively used approaches allowing us to obtain mechanistic insights into fundamental processes. By design, dilution, which depends on transfer volume (modifying mortality) and transfer interval (determining the time of interaction), is an inherent feature of these experiments, but often receives little attention. We further explore previously published data from a live predator‐prey (bacteria and ciliates) system which investigated eco‐evolutionary principles and apply a mathematical model to predict how various transfer volumes and transfer intervals would affect such an experiment. We find not only the ecological dynamics to be modified by both factors but also the evolutionary rates to be affected. Our work predicts that the evolution of the anti‐predator defense in the bacteria, and the evolution of the predation efficiency in the ciliates, both slow down with lower transfer volume, but speed up with longer transfer intervals. Our results provide testable hypotheses for future studies of predator‐prey systems, and we hope this work will help improve our understanding of how ecological and evolutionary processes together shape composition of microbial communities.     

Blood groups and natural selection by genetical environment

A survey is given of a number of investigations indicating the importance in natural selection of the genetical environment of populations and individuals.In the introduction it is observed that blood group frequency patterns are very stable, even in very small populations, and appear independent of environmental factors. They appear to be race-specific, maintained by a process of natural selection which is dependent of the racial genetical composition. Indications in favour of this hypothesis are obtained from several studies carried out in populations of mixed origin:The introduction on a small scale into the populations of New Guinea of foreign elements with some S genes may result in a population with relatively high S frequencies; the frequencies of certain of the blood group genes in the mixed negroid populations of Curaçao are not in agreement with the racial compositions of the mixtures as they have been calculated from the frequencies of other blood group genes, and the same appears to be the case in the populations of the Himalayas. The marked variation of MNSsHe frequencies in Africa may perhaps be explained by a powerful selective pressure exercized by the genetical backgrounds of the various populations, as is demonstrated by the absence of some expected genotypes among male Bush Negroes in Surinam.The effects of natural selection by genetical environment can also be demonstrated by family studies:In families with elliptocytosis Rhesus segregation shows some deviation from Mendelian laws, and the ratio of elliptocytosis-positive and-negative children appears to depend on the Rh genotype of the elliptocytosis-positive parent. From blood group studies in selected pedigrees and dizygotic twins it appears that twin pairs are more often doubly concordant for both MN and Rh than is to be expected.Some implications of the observed effects of natural selection in the study of human genetics and population dynamics are briefly discussed.     

Inference for nonlinear epidemiological models using genealogies and time series

Phylodynamics - the field aiming to quantitatively integrate the ecological and evolutionary dynamics of rapidly evolving populations like those of RNA viruses - increasingly relies upon coalescent approaches to infer past population dynamics from reconstructed genealogies. As sequence data have become more abundant, these approaches are beginning to be used on populations undergoing rapid and rather complex dynamics. In such cases, the simple demographic models that current phylodynamic methods employ can be limiting. First, these models are not ideal for yielding biological insight into the processes that drive the dynamics of the populations of interest. Second, these models differ in form from mechanistic and often stochastic population dynamic models that are currently widely used when fitting models to time series data. As such, their use does not allow for both genealogical data and time series data to be considered in tandem when conducting inference. Here, we present a flexible statistical framework for phylodynamic inference that goes beyond these current limitations. The framework we present employs a recently developed method known as particle MCMC to fit stochastic, nonlinear mechanistic models for complex population dynamics to gene genealogies and time series data in a Bayesian framework. We demonstrate our approach using a nonlinear Susceptible-Infected-Recovered (SIR) model for the transmission dynamics of an infectious disease and show through simulations that it provides accurate estimates of past disease dynamics and key epidemiological parameters from genealogies with or without accompanying time series data.     

Genetic population structure and neighbourhood population size estimates of the caddisfly Plectrocnemia conspersa

1. We used both genetic and ecological methods to evaluate the role of history and the scale of colonisation in structuring populations of the caddisfly Plectrocnemia conspersa. There was no genetic differentiation between sites up to 20 km apart, despite population sizes suggesting that genetic drift could create substantial variation at this scale. 2. Genetic differentiation between populations separated by more than 20 km was greater than expected given the contrasting short‐range trend, and implied a neighbourhood population size that is implausibly small. Therefore, the evolutionary processes that affect the short‐range trend do not explain differentiation over greater distances. 3. At small scales (<20 km), relatively short flights by winged adults spread over a number of generations could account for the spread of genes. For instance, dispersing individuals could found small (often temporary) populations, which may then grow and exchange genes with larger and more permanent local populations, amplifying the effects of the initial gene flow. 4. Over larger scales (20–500 km), substantial gaps between regions containing suitable habitat patches could reduce the number of colonisation events. Genetic patterns at this scale may date from the time they were last colonised. Previous ecological studies have rarely examined the dynamics of aquatic insect populations over these larger geographical scales, yet these processes may be central to their persistence and spread.     

Evolution of plant breeding systems

Breeding systems are important, and often neglected, aspects of the natural biology of organisms, affecting homozygosity and thus many aspects of their biology, including levels and patterns of genetic diversity and genome evolution. Among the different plant mating systems, it is useful to distinguish two types of systems: 'sex systems', hermaphroditic versus male/female and other situations; and the 'mating systems' of hermaphroditic populations, inbreeding, outcrossing or intermediate. Evolutionary changes in breeding systems occur between closely related species, and some changes occur more often than others. Understanding why such changes occur requires combined genetical and ecological approaches. I review the ideas of some of the most important theoretical models, showing how these are based on individual selection using genetic principles to ask whether alleles affecting plants' outcrossing rates or sex morphs will spread in populations. After discussing how the conclusions are affected by some of the many relevant ecological factors, I relate these theoretical ideas to empirical data from some of the many recent breeding system studies in plant populations.     

Costs of aggregation: shadow competition in a sit-and-wait predator

The role of top-down (e.g. parasitism) and bottom-up (e.g. resource competition) processes is of fundamental importance for the stability and persistence of insect herbivore populations. Although emphasis has often focused on single regulatory agents, the processes underpinning tri-trophic interactions may actually be more pluralistic. Recently, further complexities involved in the regulation of tri-trophic systems have been highlighted. In particular, life history characteristics may have a concomitant role when coupled with the regulatory effects of resource competition and/or parasitism. Here we present an age-structured model to investigate the effects of larval development period, parasitism and resource competition on the stability and persistence of herbivore–parasitoid interactions. We show that the influence of weak density dependent parasitism is sufficient to stabilise the interaction when the period of host susceptibility to parasitism is short. For longer periods of host susceptibility, parasitism needs to be highly non-linear to overcome the destabilising effects of the time delays. In systems where host development is protracted through the season we predict that resource competition is likely to be the dominant process for herbivore regulation. We use this age-structured approach to explore the population dynamics of two field studies from temperate ecosystems. Predictions from these case studies show 1) that both the strength and type of competition and parasitism are important for the stability and persistence of the particular system, and 2) that the length of the developmental period of the vulnerable host is critical to understand the influence of different regulatory processes. Host demography is of overriding importance in determining whether herbivores show outbreaks, and which particular ecological processes and mechanisms are responsible for generating such overcompensatory dynamics.     

Reproducibility of Vibrionaceae population structure in coastal bacterioplankton

How reproducibly microbial populations assemble in the wild remains poorly understood. Here, we assess evidence for ecological specialization and predictability of fine-scale population structure and habitat association in coastal ocean Vibrionaceae across years. We compare Vibrionaceae lifestyles in the bacterioplankton (combinations of free-living, particle, or zooplankton associations) measured using the same sampling scheme in 2006 and 2009 to assess whether the same groups show the same environmental association year after year. This reveals complex dynamics with populations falling primarily into two categories: (i) nearly equally represented in each of the two samplings and (ii) highly skewed, often to an extent that they appear exclusive to one or the other sampling times. Importantly, populations recovered at the same abundance in both samplings occupied highly similar habitats suggesting predictable and robust environmental association while skewed abundances of some populations may be triggered by shifts in ecological conditions. The latter is supported by difference in the composition of large eukaryotic plankton between years, with samples in 2006 being dominated by copepods, and those in 2009 by diatoms. Overall, the comparison supports highly predictable population-habitat linkage but highlights the fact that complex, and often unmeasured, environmental dynamics in habitat occurrence may have strong effects on population dynamics.     

The interplay between determinism and stochasticity in childhood diseases

An important issue in the history of ecology has been the study of the relative importance of deterministic forces and processes noise in shaping the dynamics of ecological populations. We address this question by exploring the temporal dynamics of two childhood infections, measles and whooping cough, in England and Wales. We demonstrate that epidemics of whooping cough are strongly influenced by stochasticity; fully deterministic approaches cannot achieve even a qualitative fit to the observed data. In contrast, measles dynamics are extremely well explained by a deterministic model. These differences are shown to be caused by their contrasting responses to dynamical noise due to different infectious periods.     

Time-series forecasting offers novel quantitative measure to assess loud sound event in an urban park with restored prairie

Soundscape ecology and ecoacoustics study the spatiotemporal dynamics of a soundscape and how the dynamics reflect and influence ecological processes in the environment. Soundscape analysis methods employ acoustic recording units (ARUs) that collect acoustic data in study areas over time. Analyzing these data includes computation of several acoustic diversity indices developed to quantify species abundance, richness, or habitat condition through digital audio processing and algorithm adaptations for within-group populations. However, the success of specific indices is often dependent on habitat type and biota richness present and analyzing these data can be challenging due to temporal pseudo-replication. Time-series analytical methods address the inherent problems of temporal autocorrelation for soundscape analyses challenges. Animal population dynamics fluctuate in a variety of ways due to changes in habitat or natural patterns of a landscape and chronic noise exposure, with soundscape phenology patterns evident in terrestrial and aquatic environments. Historical phenological soundscape patterns have been used to predict expected soundscape patterns in long-term studies but limited work has explored how forecasting can quantify changes in short-term studies. We evaluate how forecasting from an acoustic index can be used to quantify change in an acoustic community response to a loud, acute noise. We found that the acoustic community of a Midwestern restored prairie had decreased acoustic community activity after a loud sound event (LSE), a Civil War Reenactment, mainly driven by observed changes in the bird community and quantified using two methods: an automated acoustic index and species richness. Time-series forecasting maybe considered an underutilized tool in analyzing acoustic data whose experimental design can be flawed with temporal autocorrelation. Forecasting using auto ARIMA with acoustic indices could benefit decision makers who consider ecological questions at different time scales.     

Variability of blood group frequencies in Romania

This paper presents the results of a study of two population samples from Dobruja and Muntenia (Romania) as well as data from the literature (AB0, MN, Rhesus (D/d) and HPA). The data have been pooled according to the different regions of Romania. Interestingly, the Carpathian Mountains do not form a genetical barrier: there are clear genetical similarities between the provinces east and west as well as between the provinces south and north from the mountains. The differences and similarities between Romania as a whole and other populations in this region reflect the historical processes of the last centuries.     

The restriction fragment length polymorphism of the DNA loci of human chromosome 7]

The results of comparative RFLP analysis in some DNA loci of chromosome 7 in the populations of different Ukrainian regions are presented. Significant differences in RFLP-genotype distributions among regional populations are found. The role of different genetical processes which take place in the populations of different regions of the Ukraine is under discussion.     

Evolutionary game theory as a framework for studying biological invasions

Although biological invasions pose serious threats to biodiversity, they also provide the opportunity to better understand interactions between the ecological and evolutionary processes structuring populations and communities. However, ecoevolutionary frameworks for studying species invasions are lacking. We propose using game theory and the concept of an evolutionarily stable strategy (ESS) as a conceptual framework for integrating the ecological and evolutionary dynamics of invasions. We suggest that the pathways by which a recipient community may have no ESS provide mechanistic hypotheses for how such communities may be vulnerable to invasion and how invaders can exploit these vulnerabilities. We distinguish among these pathways by formalizing the evolutionary contexts of the invader relative to the recipient community. We model both the ecological and the adaptive dynamics of the interacting species. We show how the ESS concept provides new mechanistic hypotheses for when invasions result in long- or short-term increases in biodiversity, species replacement, and subsequent evolutionary changes.     

Linking extinction-colonization dynamics to genetic structure in a salamander metapopulation

Theory predicts that founder effects have a primary role in determining metapopulation genetic structure. However, ecological factors that affect extinction-colonization dynamics may also create spatial variation in the strength of genetic drift and migration. We tested the hypothesis that ecological factors underlying extinction-colonization dynamics influenced the genetic structure of a tiger salamander (Ambystoma tigrinum) metapopulation. We used empirical data on metapopulation dynamics to make a priori predictions about the effects of population age and ecological factors on genetic diversity and divergence among 41 populations. Metapopulation dynamics of A. tigrinum depended on wetland area, connectivity and presence of predatory fish. We found that newly colonized populations were more genetically differentiated than established populations, suggesting that founder effects influenced genetic structure. However, ecological drivers of metapopulation dynamics were more important than age in predicting genetic structure. Consistent with demographic predictions from metapopulation theory, genetic diversity and divergence depended on wetland area and connectivity. Divergence was greatest in small, isolated wetlands where genetic diversity was low. Our results show that ecological factors underlying metapopulation dynamics can be key determinants of spatial genetic structure, and that habitat area and isolation may mediate the contributions of drift and migration to divergence and evolution in local populations.     

Eco‐evolutionary partitioning metrics: assessing the importance of ecological and evolutionary contributions to population and community change

Interest in eco‐evolutionary dynamics is rapidly increasing thanks to ground‐breaking research indicating that evolution can occur rapidly and can alter the outcome of ecological processes. A key challenge in this sub‐discipline is establishing how important the contribution of evolutionary and ecological processes and their interactions are to observed shifts in population and community characteristics. Although a variety of metrics to separate and quantify the effects of evolutionary and ecological contributions to observed trait changes have been used, they often allocate fractions of observed changes to ecology and evolution in different ways. We used a mathematical and numerical comparison of two commonly used frameworks – the Price equation and reaction norms – to reveal that the Price equation cannot partition genetic from non‐genetic trait change within lineages, whereas the reaction norm approach cannot partition among‐ from within‐lineage trait change. We developed a new metric that combines the strengths of both Price‐based and reaction norm metrics, extended all metrics to analyse community change and also incorporated extinction and colonisation of species in these metrics. Depending on whether our new metric is applied to populations or communities, it can correctly separate intraspecific, interspecific, evolutionary, non‐evolutionary and interacting eco‐evolutionary contributions to trait change.     

The implications of rapid eco‐evolutionary processes for biological control ‐ a review

Novel environmental conditions experienced by introduced species can drive rapid evolution of diverse traits. In turn, rapid evolution, both adaptive and non‐adaptive, can influence population size, growth rate, and other important ecological characteristics of populations. In addition, spatial evolutionary processes that arise from a combination of assortative mating between highly dispersive individuals at the expanding edge of populations and altered reproductive rates of those individuals can accelerate expansion speed. Growing experimental evidence shows that the effects of rapid evolution on ecological dynamics can be quite large, and thus it can affect establishment, persistence, and the distribution of populations. We review the experimental and theoretical literature on such eco‐evolutionary feedbacks and evaluate the implications of these processes for biological control. Experiments show that evolving populations can establish at higher rates and grow larger than non‐evolving populations. However, non‐adaptive processes, such as genetic drift and inbreeding depression can also lead to reduced fitness and declines in population size. Spatial evolutionary processes can increase spread rates and change the fitness of individuals at the expansion front. These examples demonstrate the power of eco‐evolutionary dynamics and indicate that evolution is likely more important in biocontrol programs than previously realized. We discuss how this knowledge can be used to enhance efficacy of biological control.     

Numerical bifurcation analysis of ecosystems in a spatially homogeneous environment

The dynamics of single populations up to ecosystems, are often described by one or a set of non-linear ordinary differential equations. In this paper we review the use of bifurcation theory to analyse these non-linear dynamical systems. Bifurcation analysis gives regimes in the parameter space with quantitatively different asymptotic dynamic behaviour of the system. In small-scale systems the underlying models for the populations and their interaction are simple Lotka-Volterra models or more elaborated models with more biological detail. The latter ones are more difficult to analyse, especially when the number of populations is large. Therefore for large-scale systems the Lotka-Volterra equations are still popular despite the limited realism. Various approaches are discussed in which the different time-scale of ecological and evolutionary biological processes are considered together.     

The impact of rapid evolution on population dynamics in the wild: experimental test of eco-evolutionary dynamics

Rapid evolution challenges the assumption that evolution is too slow to impact short-term ecological dynamics. This insight motivates the study of 'Eco-Evolutionary Dynamics' or how evolution and ecological processes reciprocally interact on short time scales. We tested how rapid evolution impacts concurrent population dynamics using an aphid (Myzus persicae) and an undomesticated host (Hirschfeldia incana) in replicated wild populations. We manipulated evolvability by creating non-evolving (single clone) and potentially evolving (two-clone) aphid populations that contained genetic variation in intrinsic growth rate. We observed significant evolution in two-clone populations whether or not they were exposed to predators and competitors. Evolving populations grew up to 42% faster and attained up to 67% higher density, compared with non-evolving control populations but only in treatments exposed to competitors and predators. Increased density also correlates with relative fitness of competing clones suggesting a full eco-evolutionary dynamic cycle defined as reciprocal interactions between evolution and density.     

Eco‐evolutionary dynamics in restored communities and ecosystems

Recent ecological studies have revealed that rapid evolution within populations can have significant impacts on the ecological dynamics of communities and ecosystems. These eco‐evolutionary dynamics (EED) are likely to have substantial and quantifiable effects in restored habitats over timescales that are relevant for the conservation and restoration of small populations and threatened communities. Restored habitats may serve as “hotspots” for EED due to mismatches between transplanted genotypes and the restored environment, and novel interactions among lineages that do not share a coevolutionary history, both of which can generate strong selection for rapid evolutionary change that has immediate demographic consequences. Rapid evolution that influences population dynamics and community processes is likely to have particularly large effects during the establishment phase of restoration efforts. Finally, restoration activities and their associated long‐term monitoring programs provide outstanding opportunities for using eco‐evolutionary experimental approaches. Results from such studies will address questions about the effects of rapid evolutionary change on the ecological dynamics of populations and interacting species, while simultaneously providing critical, but currently overlooked, information for conservation practices.