Environmental degradation amplifies species' responses to temperature variation in a trophic interaction.

Land-use and climate change are two of the primary drivers of the current biodiversity crisis. However, we lack understanding of how single-species and multispecies associations are affected by interactions between multiple environmental stressors. We address this gap by examining how environmental degradation interacts with daily stochastic temperature variation to affect individual life history and population dynamics in a host-parasitoid trophic interaction, using the Indian meal moth, Plodia interpunctella, and its parasitoid wasp Venturia canescens. We carried out a single-generation individual life-history experiment and a multigeneration microcosm experiment during which individuals and microcosms were maintained at a mean temperature of 26°C that was either kept constant or varied stochastically, at four levels of host resource degradation, in the presence or absence of parasitoids. At the individual level, resource degradation increased juvenile development time and decreased adult body size in both species. Parasitoids were more sensitive to temperature variation than their hosts, with a shorter juvenile stage duration than in constant temperatures and a longer adult life span in moderately degraded environments. Resource degradation also altered the host's response to temperature variation, leading to a longer juvenile development time at high resource degradation. At the population level, moderate resource degradation amplified the effects of temperature variation on host and parasitoid populations compared with no or high resource degradation and parasitoid overall abundance was lower in fluctuating temperatures. Top-down regulation by the parasitoid and bottom-up regulation driven by resource degradation contributed to more than 50% of host and parasitoid population responses to temperature variation. Our results demonstrate that environmental degradation can strongly affect how species in a trophic interaction respond to short-term temperature fluctuations through direct and indirect trait-mediated effects. These effects are driven by species differences in sensitivity to environmental conditions and modulate top-down (parasitism) and bottom-up (resource) regulation. This study highlights the need to account for differences in the sensitivity of species' traits to environmental stressors to understand how interacting species will respond to simultaneous anthropogenic changes.

Boom-bust dynamics in biological invasions: towards an improved application of the concept.

Boom-bust dynamics - the rise of a population to outbreak levels, followed by a dramatic decline - have been associated with biological invasions and offered as a reason not to manage troublesome invaders. However, boom-bust dynamics rarely have been critically defined, analyzed, or interpreted. Here, we define boom-bust dynamics and provide specific suggestions for improving the application of the boom-bust concept. Boom-bust dynamics can arise from many causes, some closely associated with invasions, but others occurring across a wide range of ecological settings, especially when environmental conditions are changing rapidly. As a result, it is difficult to infer cause or predict future trajectories merely by observing the dynamic. We use tests with simulated data to show that a common metric for detecting and describing boom-bust dynamics, decline from an observed peak to a subsequent trough, tends to severely overestimate the frequency and severity of busts, and should be used cautiously if at all. We review and test other metrics that are better suited to describe boom-bust dynamics. Understanding the frequency and importance of boom-bust dynamics requires empirical studies of large, representative, long-term data sets that use clear definitions of boom-bust, appropriate analytical methods, and careful interpretations.

Navigating the complexity of ecological stability.

Human actions challenge nature in many ways. Ecological responses are ineluctably complex, demanding measures that describe them succinctly. Collectively, these measures encapsulate the overall 'stability' of the system. Many international bodies, including the Intergovernmental Science-Policy Platform on Biodiversity and Ecosystem Services, broadly aspire to maintain or enhance ecological stability. Such bodies frequently use terms pertaining to stability that lack clear definition. Consequently, we cannot measure them and so they disconnect from a large body of theoretical and empirical understanding. We assess the scientific and policy literature and show that this disconnect is one consequence of an inconsistent and one-dimensional approach that ecologists have taken to both disturbances and stability. This has led to confused communication of the nature of stability and the level of our insight into it. Disturbances and stability are multidimensional. Our understanding of them is not. We have a remarkably poor understanding of the impacts on stability of the characteristics that define many, perhaps all, of the most important elements of global change. We provide recommendations for theoreticians, empiricists and policymakers on how to better integrate the multidimensional nature of ecological stability into their research, policies and actions.

Testing for effects of climate change on competitive relationships and coexistence between two bird species.

Climate change is expected to have profound ecological effects, yet shifts in competitive abilities among species are rarely studied in this context. Blue tits (Cyanistes caeruleus) and great tits (Parus major) compete for food and roosting sites, yet coexist across much of their range. Climate change might thus change the competitive relationships and coexistence between these two species. Analysing four of the highest-quality, long-term datasets available on these species across Europe, we extend the textbook example of coexistence between competing species to include the dynamic effects of long-term climate variation. Using threshold time-series statistical modelling, we demonstrate that long-term climate variation affects species demography through different influences on density-dependent and density-independent processes. The competitive interaction between blue tits and great tits has shifted in one of the studied sites, creating conditions that alter the relative equilibrium densities between the two species, potentially disrupting long-term coexistence. Our analyses show that long-term climate change can, but does not always, generate local differences in the equilibrium conditions of spatially structured species assemblages. We demonstrate how long-term data can be used to better understand whether (and how), for instance, climate change might change the relationships between coexisting species. However, the studied populations are rather robust against competitive exclusion.

Ecological and evolutionary implications of food subsidies from humans.

Human activities are the main current driver of global change. From hunter-gatherers through to Neolithic societies-and particularly in contemporary industrialised countries-humans have (voluntarily or involuntarily) provided other animals with food, often with a high spatio-temporal predictability. Nowadays, as much as 30-40% of all food produced in Earth is wasted. We argue here that predictable anthropogenic food subsidies (PAFS) provided historically by humans to animals has shaped many communities and ecosystems as we see them nowadays. PAFS improve individual fitness triggering population increases of opportunistic species, which may affect communities, food webs and ecosystems by altering processes such as competition, predator-prey interactions and nutrient transfer between biotopes and ecosystems. We also show that PAFS decrease temporal population variability, increase resilience of opportunistic species and reduce community diversity. Recent environmental policies, such as the regulation of dumps or the ban of fishing discards, constitute natural experiments that should improve our understanding of the role of food supply in a range of ecological and evolutionary processes at the ecosystem level. Comparison of subsidised and non-subsidised ecosystems can help predict changes in diversity and the related ecosystem services that have suffered the impact of other global change agents.

Colonization, covariance and colour: environmental and ecological drivers of diversity-stability relationships.

Understanding the mechanisms that underlie the relationship between community diversity and biomass stability is a fundamental topic in ecology. Theory has emphasized differences in species-specific responses to environmental fluctuations as an important stabiliser of total biomass fluctuations. However, previous analyses have often been based on simplifying assumptions, such as uniform species abundance distributions, uniform environmental variance across species, and uniform environmental responses across species pairs. We compare diversity-stability relationships in model communities, based on multi-species Ricker dynamics, that follow different colonization rules during community assembly (fixed or flexible resource use) forced by temporally uncorrelated (white) or correlated (red) environmental fluctuations. The colonization rules generate characteristic niche-dependent (hierarchical, HR) environmental covariance structures, which we compare with uncorrelated (independent, IR) species' environmental responses. Environmental reddening increases biomass stability and qualitatively alters diversity-stability patterns in HR communities, under both colonization rules. Diversity-stability patterns in IR communities are qualitatively altered by colonization rules but not by environmental colour. Our results demonstrate that diversity-stability patterns are contingent upon species' colonization strategies (resource use), emergent or independent responses to environmental fluctuations, and the colour of environmental fluctuations. We describe why our results arise through differences in species traits associated with niche position. These issues are often overlooked when considering the statistical components commonly used to describe diversity-stability patterns (e.g., Overyielding, Portfolio and Covariance effects). Mechanistic understanding of different diversity-stability relationships requires consideration of the biological processes that drive different population and community level behaviours.

Species dynamics alter community diversity-biomass stability relationships.

The relationship between community diversity and biomass variability remains a crucial ecological topic, with positive, negative and neutral diversity-stability relationships reported from empirical studies. Theory highlights the relative importance of Species-Species or Species-Environment interactions in driving diversity-stability patterns. Much previous work is based on an assumption of identical (stable) species-level dynamics. We studied ecosystem models incorporating stable, cyclic and more complex species-level dynamics, with either linear or non-linear density dependence, within a locally stable community framework. Species composition varies with increasing diversity, interacting with the correlation of species' environmental responses to drive either positive or negative diversity-stability patterns, which theory based on communities with only stable species-level dynamics fails to predict. Including different dynamics points to new mechanisms that drive the full range of diversity-biomass stability relationships in empirical systems where a wider range of dynamical behaviours are important.

Turning defence into offence? Intrusion of cladoceran brood chambers by a green alga leads to reproductive failure.

Microalgae are the foundation of aquatic food webs. Their ability to defend against grazers is paramount to their survival, and modulates their ecological functions. Here, we report a novel anti-grazer strategy in the common green alga Chlorella vulgaris against two grazers, Daphnia magna and Simocephalus sp. The algal cells entered the brood chamber of both grazers, presumably using the brood current generated by the grazer's abdominal appendages. Once inside, the alga densely colonized the eggs, significantly reducing reproductive success. The effect was apparent under continuous light or higher light intensity. The algal cells remained viable following removal from the brood chamber, continuing to grow when inoculated in fresh medium. No brood chamber colonization was found when the grazers were fed the reference diet Raphidocelis subcapitata under the same experimental conditions, despite the fact that both algal species were readily ingested by the grazers and were small enough to enter their brood chambers. These observations suggest that C. vulgaris can directly inflict harm on the grazers' reproductive structure. There is no known prior example of brood chamber colonization by a microalgal prey; our results point to a new type of grazer-algae interaction in the plankton that fundamentally differs from other antagonistic ecological interactions.

When can we distinguish between neutral and non-neutral processes in community dynamics under ecological drift?

Determining whether the composition of ecological communities (species presence and abundance), can be predicted from species demographic traits, rather than being a result of neutral drift, is a key ecological question. Here we compare the similarity of community composition, from different community assembly models run under identical environmental conditions, where interspecific competition is assumed to be either neutral or niche-based. In both cases, species colonize a focal patch from a network of neighbouring patches in a metacommunity. We highlight the circumstances (rate and spatial scale of dispersal, and the relative importance of ecological drift) where commonly used community similarity metrics or species rank-abundance relationships are likely to give similar results, regardless of the underlying processes (neutral or non-neural) driving species' dynamics. As drift becomes more important in driving species abundances, deterministic niche structure has a smaller influence. Our ability to discriminate between different underlying processes driving community organization depends on the relative importance of different drift processes that operate on different spatial scales.

Increasing community size and connectance can increase stability in competitive communities.

The relationship between community complexity and stability has been the subject of an enduring debate in ecology over the last 50 years. Results from early model communities showed that increased complexity is associated with decreased local stability. I demonstrate that increasing both the number of species in a community and the connectance between these species results in an increased probability of local stability in discrete-time competitive communities, when some species would show unstable dynamics in the absence of competition. This is shown analytically for a simple case and across a wider range of community sizes using simulations, where individual species have dynamics that can range from stable point equilibria to periodic or more complex. Increasing the number of competitive links in the community reduces per-capita growth rates through an increase in competitive feedback, stabilising oscillating dynamics. This result was robust to the introduction of a trade-off between competitive ability and intrinsic growth rate and changes in species interaction strengths. This throws new light on the discrepancy between the theoretical view that increased complexity reduces stability and the empirical view that more complex systems are more likely to be stable, giving one explanation for the relative lack of complex dynamics found in natural systems. I examine how these results relate to diversity-biomass stability relationships and show that an analytical solution derived in the region of stable equilibrium dynamics captures many features of the change in biomass fluctuations with community size in communities including species with oscillating dynamics.

Community stability under different correlation structures of species' environmental responses.

The outcome of species interactions in a variable environment is expected to depend on how similarly different species react to variation in environmental conditions. We study community stability (evenness and species diversity) in competitive communities that are either closed or subjected to random migration, under different regimes of environmental forcing. Community members respond to environmental variation: (i) independently (IR), (ii) in a positively correlated way (CR), or (iii) hierarchically, according to niche differences (HR). Increasing the amplitude of environmental variation and environmental reddening both reduce species evenness in closed communities through a reduction in species richness and increased skew in species abundances, under all three environmental response scenarios, although autocorrelation only has a minor effect with HR. Open communities show important qualitative differences, according to changes in the correlation structure of species' environmental responses. There is an intermediate minimum in evenness for HR communities with increasing environmental amplitude, explained by the interaction of changes in species richness and changes in the variance of within-species environmental responses across the community. Changes in autocorrelation also lead to qualitative differences between IR, CR and HR communities. Our results highlight the importance of considering mechanistically derived, hierarchical environmental correlations between species when addressing the influence of environmental variation on ecological communities, not only uniform environmental correlation across all species within a community.

The predictability of ecological stability in a noisy world.

Random environmental variation, or stochasticity, is a key determinant of ecological dynamics. While we have some appreciation of how environmental stochasticity can moderate the variability and persistence of communities, we know little about its implications for the nature and predictability of ecological responses to large perturbations. Here, we show that shifts in the temporal autocorrelation (colour) of environmental noise provoke trade-offs in ecological stability across a wide range of different food-web structures by stabilizing dynamics in some dimensions, while simultaneously destabilizing them in others. Specifically, increasingly positive autocorrelation (reddening) of environmental noise increases resilience by hastening the recovery of food webs following a large perturbation, but reduces their resistance to perturbation and increases their temporal variability (reduces biomass stability). In contrast, all stability dimensions become less predictable, showing increased variability around the mean response, as environmental noise reddens. Moreover, we found environmental reddening to be a considerably more important determinant of stability than intrinsic food-web characteristics. These findings reveal the fundamental and dominant role played by environmental stochasticity in determining the dynamics and stability of ecosystems, and extend our understanding of how the multiple dimensions of stability relate to each other beyond simple white noise environments.

Community extinction patterns in coloured environments.

Understanding community responses to environmental variation is a fundamental aspect of ecological research, with direct ecological, conservation and economic implications. Here, we examined the role of the magnitude, correlation and autocorrelation structures of environmental variation on species' extinction risk (ER), and the probability of actual extinction events in model competitive communities. Both ER and probability increased with increasing positive autocorrelation when species responded independently to the environment, yet both decreased with a strong correlation between species-specific responses. These results are framed in terms of the synchrony between--and magnitude of variation within--species population sizes and are explained in terms of differences in noise amplification under different conditions. The simulation results are robust to changes in the strength of interspecific density dependence, and whether noise affects density-independent or density-dependent population processes. Similar patterns arose under different ranges of noise severity when these different model assumptions were examined. We compared our results with those from an analytically derived solution, which failed to capture many features of the simulation results.

Population synchrony in small-world networks.

Network topography ranges from regular graphs (linkage between nearest neighbours only) via small-world graphs (some random connections between nodes) to completely random graphs. Small-world linkage is seen as a revolutionary architecture for a wide range of social, physical and biological networks, and has been shown to increase synchrony between oscillating subunits. We study small-world topographies in a novel context: dispersal linkage between spatially structured populations across a range of population models. Regular dispersal between population patches interacting with density-dependent renewal provides one ecological explanation for the large-scale synchrony seen in the temporal fluctuations of many species, for example, lynx populations in North America, voles in Fennoscandia and grouse in the UK. Introducing a small-world dispersal kernel leads to a clear reduction in synchrony with both increasing dispersal rate and small-world dispersal probability across a variety of biological scenarios. Synchrony is also reduced when populations are affected by globally correlated noise. We discuss ecological implications of small-world dispersal in the frame of spatial synchrony in population fluctuations.

Multiple dimensions of biodiversity drive human interest in tide pool communities.

Activities involving observation of wild organisms (e.g. wildlife watching, tidepooling) can provide recreational and learning opportunities, with biologically diverse animal assemblages expected to be more stimulating to humans. In turn, more diverse communities may enhance human interest and facilitate provisioning of cultural services. However, no experimental tests of this biodiversity-interest hypothesis exist to date. We therefore investigated the effects of different dimensions of animal biodiversity (species richness, phyletic richness and functional diversity) on self-reported interest using tide pools as a model system. We performed two experiments by manipulating: (1) the richness of lower (species) and higher taxonomic levels (phyla) in an image based, online survey, and (2) the richness of the higher taxonomic level (phyla) in live public exhibits. In both experiments, we further quantified functional diversity, which varied freely, and within the online experiment we also included the hue diversity and colourfulness arising from the combination of organisms and the background scenes. Interest was increased by phyletic richness (both studies), animal species richness (online study) and functional diversity (online study). A structural equation model revealed that functional diversity and colourfulness (of the whole scene) also partially mediated the effects of phyletic richness on interest in the online study. In both studies, the presence of three of four phyla additively increased interest, supporting the importance of multiple, diverse phyla rather than a single particularly interesting phylum. These results provide novel experimental evidence that multiple dimensions of biodiversity enhance human interest and suggest that conservation initiatives that maintain or restore biodiversity will help stimulate interest in ecosystems, facilitating educational and recreational benefits.

Functional responses of cougars (Puma concolor) in a multiple prey-species system.

The study of predator-prey interactions is commonly analyzed using functional responses to gain an understanding of predation patterns and the impact they have on prey populations. Despite this, little is known about predator-prey systems with multiple prey species in sites near the equator. Here we studied the functional response of cougars (Puma concolor) in relation to their main prey, armadillo (Dasypus novemcinctus), coati (Nasua narica) and white-tailed deer (Odocoileus virginianus). Between 2004 and 2010, cougar scats were collected along 5 transects to estimate the consumption of different prey species. A relative abundance index (RAI) was calculated for each prey species and cougar using 18 camera traps. We compared Holling type I, II and III functional response models to determine patterns in prey consumption based on the relative abundance and biomass of each prey species consumed. The 3 main prey species comprised 55% (armadillo), 17% (coati) and 8% (white-tailed deer) of the diet. Type I and II functional responses described consumption of the 2 most common prey species armadillos and coati similarly well, while a type I response best characterized consumption of white-tailed deer. A negative correlation between the proportions of armadillo versus coati and white-tailed deer biomass in cougar scats suggests switching to consume alternative prey, confirming high foraging plasticity of this carnivore. This work represents one of the few studies to compare functional responses across multiple prey species, combined with evidence for prey-switching at low densities of preferred prey.

The impacts of different management strategies and environmental forcing in ecological communities.

Understanding the effects of population management on the community a target species belongs to is of key importance for successful management. It is known that the removal or extinction of a single species in a community may lead to extinctions of other community members. In our study, we assess the impacts of population management on competitive communities, studying the response of both locally stable and unstable communities of varying size (between four and 10 species) to three different management strategies; harvesting of a target species, harvesting with non-targeted catch, and stocking of the target species. We also studied the consequences of selecting target species with different relative abundances, as well as the effects of varying environmental conditions. We show here how the effects of management in competitive communities extend far beyond the target population. A crucial role is played by the underlying stability properties of the community under management. In general, locally unstable communities are more vulnerable to perturbation through management. Furthermore, the community response is shown to be sensitive to the relative density of the target species. Of considerable interest is the result that even a small (2.5%) increase in the population size of the target species through stocking may lead to extinction of other community members. These results emphasize the importance of considering and understanding multi-species interactions in population management.

Confounding environmental colour and distribution shape leads to underestimation of population extinction risk.

The colour of environmental variability influences the size of population fluctuations when filtered through density dependent dynamics, driving extinction risk through dynamical resonance. Slow fluctuations (low frequencies) dominate in red environments, rapid fluctuations (high frequencies) in blue environments and white environments are purely random (no frequencies dominate). Two methods are commonly employed to generate the coloured spatial and/or temporal stochastic (environmental) series used in combination with population (dynamical feedback) models: autoregressive [AR(1)] and sinusoidal (1/f) models. We show that changing environmental colour from white to red with 1/f models, and from white to red or blue with AR(1) models, generates coloured environmental series that are not normally distributed at finite time-scales, potentially confounding comparison with normally distributed white noise models. Increasing variability of sample Skewness and Kurtosis and decreasing mean Kurtosis of these series alter the frequency distribution shape of the realised values of the coloured stochastic processes. These changes in distribution shape alter patterns in the probability of single and series of extreme conditions. We show that the reduced extinction risk for undercompensating (slow growing) populations in red environments previously predicted with traditional 1/f methods is an artefact of changes in the distribution shapes of the environmental series. This is demonstrated by comparison with coloured series controlled to be normally distributed using spectral mimicry. Changes in the distribution shape that arise using traditional methods lead to underestimation of extinction risk in normally distributed, red 1/f environments. AR(1) methods also underestimate extinction risks in traditionally generated red environments. This work synthesises previous results and provides further insight into the processes driving extinction risk in model populations. We must let the characteristics of known natural environmental covariates (e.g., colour and distribution shape) guide us in our choice of how to best model the impact of coloured environmental variation on population dynamics.