Evolving metacommunities: toward an evolutionary perspective on metacommunities

The metacommunity framework predicts that local coexistence depends on the outcome of local species interactions and regional migration. In analogous fashion, spatial structure among populations can shape species interactions through evolutionary mechanisms. Yet, most metacommunity theories assume that populations do not evolve. Here, we evaluate how evolution shapes local species coexistence and exclusion within the multiscale and multispecies context embodied by the metacommunity framework. In general, coexistence in joint ecological-evolutionary models requires low to intermediate dispersal rates that can promote maintenance of both regional species and genetic diversity. These conditions support a set of key mechanisms that modify patterns of species coexistence including local adaptation, gene storage effects, genetic rescue effects, spatial genetic subsidies, and metacommunity evolution. Multispecies extensions indicate that correlated selection can further alter the outcome of interspecific interactions depending on the magnitude and direction of correlations and shape of fitness trade-offs. We suggest that an evolving metacommunity perspective has the potential to generate novel predictions about community structure and function by incorporating the genetic and species diversity that characterize natural communities. In adopting such a perspective, we seek to facilitate understanding about the interactions between evolutionary and metacommunity dynamics.     

Bite forces and evolutionary adaptations to feeding ecology in carnivores

The Carnivora spans the largest ecological and body size diversity of any mammalian order, making it an ideal basis for studies of evolutionary ecology and functional morphology. For animals with different feeding ecologies, it may be expected that bite force represents an important evolutionary adaptation, but studies have been constrained by a lack of bite force data. In this study we present predictions of bite forces for 151 species of extant carnivores, comprising representatives from all eight families and the entire size and ecological spectrum within the order. We show that, when normalized for body size, bite forces differ significantly between the various feeding categories. At opposing extremes and independent of genealogy, consumers of tough fibrous plant material and carnivores preying on large prey both have high bite forces for their size, while bite force adjusted for body mass is low among specialized insectivores. Omnivores and carnivores preying on small prey have more moderate bite forces for their size. These findings indicate that differences in bite force represent important adaptations to and indicators of differing feeding ecologies throughout carnivoran evolution. Our results suggest that the incorporation of bite force data may assist in the construction of more robust evolutionary and palaeontological analyses of feeding ecology.     

Local extinction of a foundation species in a hypoxic estuary: integrating individuals to ecosystem

We integrated across individual, population, community, and ecosystem levels to understand the impact of environmental stress by tracking the foundation species Mytilus edulis in the hypoxic estuary Narragansett Bay, Rhode Island, USA. Our initial surveys revealed that the mussels occurred in nine extensive (2-28 ha) dense (814-9943 individuals/m2) subtidal reefs that attracted a diverse suite of predators (sea stars, crabs, gastropods). Hypoxia occurred in the summer of 2001, and a mussel transplant experiment revealed overall reduced growth rates of individuals, and higher mortality rates among larger mussels. At the population level, large decreases in densities and cover of mussels were correlated with dissolved oxygen concentrations, leading to extinction at one site and reductions of over an order of magnitude at others. Within one year, seven of the eight remaining populations were edged to extinction, and the previously extinct population was recolonized. At the community level, a predator exclusion experiment indicated that predation was an unimportant source of mussel mortality during the hypoxic period, in part due to the emigration of sea stars, as predicted by the Consumer Stress Model. However, mussels were too intolerant to hypoxia to have a net benefit from the predation refuge. The seasonal (summer) occurrence of hypoxia allowed sea stars to return following a lag, as predicted by a stress return time model, and the resumption of predation contributed to the subsequent extinction of mussel populations. At the ecosystem level, the initial filtration rate of the mussel reefs was estimated at 134.6 x 10(6) m3/d, equivalent to filtering the volume of the bay 1.3 times during the 26-d average residence time. That function was reduced by >75% following hypoxia. The effect of hypoxia on each level of organization had consequences at others. For example, size-specific mortality and decreased growth of individuals, and reduced filtration capacity of reefs, indicated a loss of the ability of mussels to entrain planktonic productivity and potential to control future eutrophication and hypoxia. Our study quantified patterns of loss and identified pathways within an integrative framework of feedbacks, summarized in a conceptual model that is applicable to similar foundation species subjected to environmental stress.     

Feeding ecology and evolutionary survival of the living coelacanth Latimeria chalumnae

One concept of evolutionary ecology holds that a living fossil is the result of past evolutionary events, and is adapted to recent selective forces only if they are similar to the selective forces in the past. We describe the present environment of the living coelacanth Latimeria chalumnae Smith, 1939 at Grande Comore, western Indian Ocean and report depth-dependent cave distribution, temperature, salinity and oxygen values which are compared to the fish's distribution and its physiological demands. We studied the activity pattern, feeding behaviour, prey abundance and hunting success to evaluate possible links between environmental conditions, feeding ecology and evolutionary success of this ancient fish. Transmitter tracking experiments indicate nocturnal activity of the piscivorous predator which hunts between approximately 200 m below the surface to 500 m depth. Fish and prey density were measured between 200 and 400 m, both increase with depth. Feeding tracks and feeding strikes of the coelacanth at various depths were simulated with the help of video and laser techniques. Along a 9447 m video transect a total of 31 potential feeding strikes occurred. Assuming 100% hunting success, medium-sized individuals would obtain 122 g and large females 299 g of prey. Estimates of metabolic rates revealed for females 3.7 ml O₂ kg⁻¹ h⁻¹ and for males 4.5 ml O₂ kg⁻¹ h⁻¹. Today coelacanths are considered to be a specialist deep-water form and to inhabit, with their ancient morphology, a contemporary environment where they compete with advanced, modern fish. Received: 5 July 1999 / Accepted: 11 November 1999     

Testing hypotheses in evolutionary ecology with imperfect detection: capture-recapture structural equation modeling

Studying evolutionary mechanisms in natural populations often requires testing multifactorial scenarios of causality involving direct and indirect relationships among individual and environmental variables. It is also essential to account for the imperfect detection of individuals to provide unbiased demographic parameter estimates. To cope with these issues, we developed a new approach combining structural equation models with capture-recapture models (CR-SEM) that allows the investigation of competing hypotheses about individual and environmental variability observed in demographic parameters. We employ Markov chain Monte Carlo sampling in a Bayesian framework to (1) estimate model parameters, (2) implement a model selection procedure to evaluate competing hypotheses about causal mechanisms, and (3) assess the fit of models to data using posterior predictive checks. We illustrate the value of our approach using two case studies on wild bird populations. We first show that CR-SEM can be useful to quantify the action of selection on a set of phenotypic traits with an analysis of selection gradients on morphological traits in Common Blackbirds (Turdus merula). In a second case study on Blue Tits (Cyanistes caeruleus), we illustrate the use of CR-SEM to study evolutionary trade-offs in the wild, while accounting for varying environmental conditions.     

Information-theoretic approaches to statistical analysis in behavioural ecology: an introduction

Scientific thinking may require the consideration of multiple hypotheses, which often call for complex statistical models at the level of data analysis. The aim of this introduction is to provide a brief overview on how competing hypotheses are evaluated statistically in behavioural ecological studies and to offer potentially fruitful avenues for future methodological developments. Complex models have traditionally been treated by model selection approaches using threshold-based removal of terms, i.e. stepwise selection. A recently introduced method for model selection applies an information-theoretic (IT) approach, which simultaneously evaluates hypotheses by balancing between model complexity and goodness of fit. The IT method has been increasingly propagated in the field of ecology, while a literature survey shows that its spread in behavioural ecology has been much slower, and model simplification using stepwise selection is still more widespread than IT-based model selection. Why has the use of IT methods in behavioural ecology lagged behind other disciplines? This special issue examines the suitability of the IT method for analysing data with multiple predictors, which researchers encounter in our field. The volume brings together different viewpoints to aid behavioural ecologists in understanding the method, with the hope of enhancing the statistical integration of our discipline.     

Niche and fitness differences relate the maintenance of diversity to ecosystem function

The frequently observed positive correlation between species diversity and community biomass is thought to depend on both the degree of resource partitioning and on competitive dominance between consumers, two properties that are also central to theories of species coexistence. To make an explicit link between theory on the causes and consequences of biodiversity, we define in a precise way two kinds of differences among species: niche differences, which promote coexistence, and relative fitness differences, which promote competitive exclusion. In a classic model of exploitative competition, promoting coexistence by increasing niche differences typically, although not universally, increases the "relative yield total", a measure of diversity's effect on the biomass of competitors. In addition, however, we show that promoting coexistence by decreasing relative fitness differences also increases the relative yield total. Thus, two fundamentally different mechanisms of species coexistence both strengthen the influence of diversity on biomass yield. The model and our analysis also yield insight on the interpretation of experimental diversity manipulations. Specifically, the frequently reported "complementarity effect" appears to give a largely skewed estimate of resource partitioning. Likewise, the "selection effect" does not seem to isolate biomass changes attributable to species composition rather than species richness, as is commonly presumed. We conclude that past inferences about the cause of observed diversity-function relationships may be unreliable, and that new empirical estimates of niche and relative fitness differences are necessary to uncover the ecological mechanisms responsible for diversity-function relationships.     

Resistance and re-organization of an ecosystem in response to biological invasion: Some hypotheses

Using a dynamic model of Lake Chozas developed by Marchi et al. (2011), we tested three hypotheses about recovery of the indigenous community and water quality after radical changes caused by introduction of an invasive allochthonous crayfish, Procambarus clarkii:

1.

Can the lake resist the pressure of an invasive species, like P. clarkii, by adaptation?

2.

Can the ecosystem recover when all the crayfish are removed and low phosphorus concentrations persist in inflow water?

3.

Does the simulated recovery of submerged vegetation occur at a total phosphorus concentration below 100 mg TP m⁻³, as estimated by Scheffer et al. (1993), Scheffer (1997), Jeppesen et al. (1998) and Zhang et al. (2003)?

We obtained the following answers:

1.

Lake Chozas can at least partly resist by adaptation. A combination of possible parameter changes could lead to a significant increase in eco-exergy.

2.

Removal of the phosphorus represented by crayfish (by harvesting) implies complete recovery of the lake and its eco-exergy, albeit not necessarily with the same organisms having the same properties.

3.

The expected hysteresis created by introduction and harvesting of crayfish is observed under the following conditions: phytoplankton dominance at total phosphorus ≥ about 200-250 mg TP m⁻³ and submerged vegetation returns at total phosphorus < 100 mg TP m⁻³.

     

Biodiversity in the ecosystem of the Portuguese continental shelf: distributional ecology and the role of benthic amphipods

The biodiversity and distributional ecology of amphipod crustaceans were analysed from benthic surveys in 1985 and 1986 along the coast of Portugal, from the Tagus canyon to the coast of Minho, in order to determine their role in the ecosystem of the Portuguese continental self. In samples obtained from different types of substrate between 18 and 545 m, we identified 113 species belonging to 51 genera of 24 families. Seventeen of these species were new records for the Portuguese coast. The relative frequencies and number of individuals of all species were analysed. The genus Ampelisca clearly dominated the group of the most frequent and abundant species, which also exhibited a large range of vertical distribution. Most of the 113 species were found only down to 150 m depth, corresponding roughly to the lower limit of the continental shelf. Fifteen species exhibited clear bathyal traits, being consistently found deeper than 100 m, especially in the Tagus canyon and the upper zone of the continental slope. The most frequent and abundant species were found predominatly or in large numbers in medium to fine sand bottoms. This type of substrate also contained the highest number of species. The granulometric structure of the habitat could constitute the most important factor controlling both biodiversity and development of large amphipod populations, since oxygen level, water exchange, number of available shelters, and probably the quantity and quality of nutritional resources all vary as a function of substrate texture. Because of their dominance in both frequency and abundance, several Ampelisca species may constitute an important food source for many secondary consumers of the shelf sediments, especially for demersal fishes.     

Ecodynamics: Towards an evolutionary thermodynamics of ecosystems

This paper collects, with few minor formal changes, two of the latest scientific contributions ( [Tiezzi, 2006a] and [Tiezzi et al., 2010]) written by Prof. Enzo Tiezzi, where he introduced new concepts and tools to formalize and understand the role of thermodynamics in ecosystems theory.Particular attention is devoted to goal functions, to the relation of matter, energy, space and time and to the interdisciplinary approach connecting thermodynamics and biology. Entropy is discussed as a fundamental goal function in the far from equilibrium framework. The relationship between entropy, as a non-state function, and the state-function energy is stressed and discussed, at the light of the role of information. The theory of probability is also discussed in the light of new theoretical findings related to the role of events, also in terms of entropy and evolutionary thermodynamics.Confined Ontic Open Systems (COOS) represent the latest model proposed by Prof. Tiezzi based on his Ecodynamic theory, evolutionary thermodynamics, Ulanowicz's ontic. The model has a wide range of applications, including ecosystems, ecological economics, urban organization, the supra-molecular structure of water and global biosphere's models. The model is explained in terms of evolutionary thermodynamics and Jørgensen's ecosystems theory.     

Movement ecology: size-specific behavioral response of an invasive snail to food availability

Immigration, emigration, migration, and redistribution describe processes that involve movement of individuals. These movements are an essential part of contemporary ecological models, and understanding how movement is affected by biotic and abiotic factors is important for effectively modeling ecological processes that depend on movement. We asked how phenotypic heterogeneity (body size) and environmental heterogeneity (food resource level) affect the movement behavior of an aquatic snail (Tarebia granifera), and whether including these phenotypic and environmental effects improves advection-diffusion models of movement. We postulated various elaborations of the basic advection diffusion model as a priori working hypotheses. To test our hypotheses we measured individual snail movements in experimental streams at high- and low-food resource treatments. Using these experimental movement data, we examined the dependency of model selection on resource level and body size using Akaike's Information Criterion (AIC). At low resources, large individuals moved faster than small individuals, producing a platykurtic movement distribution; including size dependency in the model improved model performance. In stark contrast, at high resources, individuals moved upstream together as a wave, and body size differences largely disappeared. The model selection exercise indicated that population heterogeneity is best described by the advection component of movement for this species, because the top-ranked model included size dependency in advection, but not diffusion. Also, all probable models included resource dependency. Thus population and environmental heterogeneities both influence individual movement behaviors and the population-level distribution kernels, and their interaction may drive variation in movement behaviors in terms of both advection rates and diffusion rates. A behaviorally informed modeling framework will integrate the sentient response of individuals in terms of movement and enhance our ability to accurately model ecological processes that depend on animal movement.     

Forest to reclaimed mine land use change leads to altered ecosystem structure and function.

The United States' use of coal results in many environmental alterations. In the Appalachian coal belt region, one widespread alteration is conversion of forest to reclaimed mineland. The goal of this study was to quantify the changes to ecosystem structure and function associated with a conversion from forest to reclaimed mine grassland by comparing a small watershed containing a 15-year-old reclaimed mine with a forested, reference watershed in western Maryland. Major differences were apparent between the two watersheds in terms of biogeochemistry. Total C, N, and P pools were all substantially lower at the mined site, mainly due to the removal of woody biomass but also, in the case of P, to reductions in soil pools. Mineral soil C, N, and P pools were 96%, 79%, and 69% of native soils, respectively. Although annual runoff from the watersheds was similar, the mined watershed exhibited taller, narrower storm peaks as a result of a higher soil bulk density and decreased infiltration rates. Stream export of N was much lower in the mined watershed due to lower net nitrification rates and nitrate concentrations in soil. However, stream export of sediment and P and summer stream temperature were much higher. Stream leaf decomposition was reduced and macroinvertebrate community structure was altered as a result of these changes to the stream environment. This land use change leads to substantial, long-term changes in ecosystem capital and function.     

Microbial stress-response physiology and its implications for ecosystem function

Microorganisms have a variety of evolutionary adaptations and physiological acclimation mechanisms that allow them to survive and remain active in the face of environmental stress. Physiological responses to stress have costs at the organismal level that can result in altered ecosystem-level C, energy, and nutrient flows. These large-scale impacts result from direct effects on active microbes' physiology and by controlling the composition of the active microbial community. We first consider some general aspects of how microbes experience environmental stresses and how they respond to them. We then discuss the impacts of two important ecosystem-level stressors, drought and freezing, on microbial physiology and community composition. Even when microbial community response to stress is limited, the physiological costs imposed on soil microbes are large enough that they may cause large shifts in the allocation and fate of C and N. For example, for microbes to synthesize the osmolytes they need to survive a single drought episode they may consume up to 5% of total annual net primary production in grassland ecosystems, while acclimating to freezing conditions switches Arctic tundra soils from immobilizing N during the growing season to mineralizing it during the winter. We suggest that more effectively integrating microbial ecology into ecosystem ecology will require a more complete integration of microbial physiological ecology, population biology, and process ecology.     

The empirical modeling of an ecosystem

The authors have endeavored to create a verified a-posteriori model of a planktonic ecosystem. Verification of an empirically derived set of first-order, quadratic differential equations proved elusive due to the sensitivity of the model system to changes in initial conditions. Efforts to verify a similarly derived set of linear differential equations were more encouraging, yielding reasonable behavior for half of the ten ecosystem compartments modeled. The well-behaved species models gave indications as to the rate-controlling process in the ecosystem.     

Fooled by local robustness: an applied ecology perspective

In this short discussion, we point out that it is apparently as easy to be fooled by robustness as it is to be fooled by randomness. Our objective is to bring to the attention of applied ecologists that radius-of-stability robustness models are models of local robustness. As such, these models are utterly unsuitable for the treatment/management of a severe uncertainty characterized by a vast uncertainty space and a likelihood-free quantification of the uncertainty. This observation is particularly pertinent to applications of info-gap decision theory in ecology, conservation biology, and environmental management, where the objective is to identify decisions that are robust against a severe uncertainty of this type.     

Application of an evolutionary algorithm to the inverse parameter estimation of an individual-based model

Inverse parameter estimation of individual-based models (IBMs) is a research area which is still in its infancy, in a context where conventional statistical methods are not well suited to confront this type of models with data. In this paper, we propose an original evolutionary algorithm which is designed for the calibration of complex IBMs, i.e. characterized by high stochasticity, parameter uncertainty and numerous non-linear interactions between parameters and model output. Our algorithm corresponds to a variant of the population-based incremental learning (PBIL) genetic algorithm, with a specific “optimal individual” operator. The method is presented in detail and applied to the individual-based model OSMOSE. The performance of the algorithm is evaluated and estimated parameters are compared with an independent manual calibration. The results show that automated and convergent methods for inverse parameter estimation are a significant improvement to existing ad hoc methods for the calibration of IBMs.     

Using the Price Equation to partition the effects of biodiversity loss on ecosystem function

Species loss can impact ecosystem functioning, but no general framework for analyzing these impacts exists. Here I derive a general partitioning of the effects of species loss on any ecosystem function comprising the summed contributions of individual species (e.g., primary productivity). The approach partitions the difference in ecosystem function between two sites (a "pre-loss" site, and a "post-loss" site comprising a strict subset of the species at the pre-loss site) into additive components attributable to different effects. The approach does not assume a particular experimental design or require monoculture data, making it more general than previous approaches. Using the Price Equation from evolutionary biology, I show that three distinct effects cause ecosystem function to vary between sites: the "species richness effect" (SRE; random loss of species richness), the "species composition effect" (SCE; nonrandom loss of high- or low-functioning species), and the "context dependence effect" (CDE; post-loss changes in the functioning of the remaining species). The SRE reduces ecosystem function without altering mean function per species. The SCE is analogous to natural selection in evolution. Nonrandom loss of, for example, high-functioning species will reduce mean function per species, and thus total function, just as selection against large individuals in an evolving population reduces mean body size in the next generation. The CDE is analogous to imperfect transmission in evolution. For instance, any factor (e.g., an environmental change) causing offspring to attain smaller body sizes than their parents (imperfect transmission) will reduce the mean body size in the next generation. Analogously, any factor causing the species remaining at the post-loss site to make smaller functional contributions than at the pre-loss site will reduce mean function per species, and thus total function. I use published data to illustrate how this new partition generalizes previous approaches, facilitates comparative analyses, and generates new empirical insights. In particular, the SCE often is less important than other effects.     

Toward an ecosystem approach to ICM: assessing ecological provinces at sea by remote sensing

The ecosystem approach requires that all elements of an ecosystem, and their mutual interactions, be taken into consideration in any management effort. The selection of suitable geographical units, where this approach can be taken, requires the assessment of ecological provinces, characterized by a coherent set of environmental traits. The marine side of coastal zones, where the interaction between atmosphere, land and sea is not bounded by evident geographic markers, represents a critical factor in this assessment. A coastal province can be defined by physical setting, but also by its bio-geo-chemical features, ideally on the basis of synoptic remote sensing data, collected at space/time scales not accessible by other means. Classifications based on indicators such as temperature, wind speed and chlorophyll-like pigments, demonstrate the identification of potential ecological provinces in the Mediterranean Sea. The results suggest remote sensing as the ideal tool to set up the basis for an ecosystem approach to the management of each province.