Local intraspecific aggregation in phytoplankton model communities: spatial scales of occurrence and implications for coexistence.

The coexistence of multiple phytoplankton species despite their reliance on similar resources is often explained with mean-field models assuming mixed populations. In reality, observations of phytoplankton indicate spatial aggregation at all scales, including at the scale of a few individuals. Local spatial aggregation can hinder competitive exclusion since individuals then interact mostly with other individuals of their own species, rather than competitors from different species. To evaluate how microscale spatial aggregation might explain phytoplankton diversity maintenance, an individual-based, multispecies representation of cells in a hydrodynamic environment is required. We formulate a three-dimensional and multispecies individual-based model of phytoplankton population dynamics at the Kolmogorov scale. The model is studied through both simulations and the derivation of spatial moment equations, in connection with point process theory. The spatial moment equations show a good match between theory and simulations. We parameterized the model based on phytoplankters' ecological and physical characteristics, for both large and small phytoplankton. Defining a zone of potential interactions as the overlap between nutrient depletion volumes, we show that local species composition-within the range of possible interactions-depends on the size class of phytoplankton. In small phytoplankton, individuals remain in mostly monospecific clusters. Spatial structure therefore favours intra- over inter-specific interactions for small phytoplankton, contributing to coexistence. Large phytoplankton cell neighbourhoods appear more mixed. Although some small-scale self-organizing spatial structure remains and could influence coexistence mechanisms, other factors may need to be explored to explain diversity maintenance in large phytoplankton.

No sensitivity to functional forms in the Rosenzweig-MacArthur model with strong environmental stochasticity.

The classic Rosenzweig-MacArthur predator-prey model has been shown to exhibit, like other coupled nonlinear ordinary differential equations (ODEs) from ecology, worrying sensitivity to model structure. This sensitivity manifests as markedly different community dynamics arising from saturating functional responses with nearly identical shapes but different mathematical expressions. Using a stochastic differential equation (SDE) version of the Rosenzweig-MacArthur model with the three functional responses considered by Fussmann & Blasius (2005), I show that such sensitivity seems to be solely a property of ODEs or stochastic systems with weak noise. SDEs with strong environmental noise have by contrast very similar fluctuations patterns, irrespective of the mathematical formula used. Although eigenvalues of linearized predator-prey models have been used as an argument for structural sensitivity, they can also be an argument against structural sensitivity. While the sign of the eigenvalues' real part is sensitive to model structure, its magnitude and the presence of imaginary parts are not, which suggests noise-driven oscillations for a broad range of carrying capacities. I then discuss multiple other ways to evaluate structural sensitivity in a stochastic setting, for predator-prey or other ecological systems.

Reduced within-population quantitative genetic variation is associated with climate harshness in maritime pine.

How evolutionary forces interact to maintain genetic variation within populations has been a matter of extensive theoretical debates. While mutation and exogenous gene flow increase genetic variation, stabilizing selection and genetic drift are expected to deplete it. To date, levels of genetic variation observed in natural populations are hard to predict without accounting for other processes, such as balancing selection in heterogeneous environments. We aimed to empirically test three hypotheses: (i) admixed populations have higher quantitative genetic variation due to introgression from other gene pools, (ii) quantitative genetic variation is lower in populations from harsher environments (i.e., experiencing stronger selection), and (iii) quantitative genetic variation is higher in populations from heterogeneous environments. Using growth, phenological and functional trait data from three clonal common gardens and 33 populations (522 clones) of maritime pine (Pinus pinaster Aiton), we estimated the association between the population-specific total genetic variances (i.e., among-clone variances) for these traits and ten population-specific indices related to admixture levels (estimated based on 5165 SNPs), environmental temporal and spatial heterogeneity and climate harshness. Populations experiencing colder winters showed consistently lower genetic variation for early height growth (a fitness-related trait in forest trees) in the three common gardens. Within-population quantitative genetic variation was not associated with environmental heterogeneity or population admixture for any trait. Our results provide empirical support for the potential role of natural selection in reducing genetic variation for early height growth within populations, which indirectly gives insight into the adaptive potential of populations to changing environments.

Effects of Stage Structure on Coexistence: Mixed Benefits.

The properties of competition models where all individuals are identical are relatively well-understood; however, juveniles and adults can experience or generate competition differently. We study here less well-known structured competition models in discrete time that allow multiple life history parameters to depend on adult or juvenile population densities. A numerical study with Ricker density-dependence suggested that when competition coefficients acting on juvenile survival and fertility reflect opposite competitive hierarchies, stage structure could foster coexistence. We revisit and expand those results. First, through a Beverton-Holt two-species juvenile-adult model, we confirm that these findings do not depend on the specifics of density-dependence or life cycles, and obtain analytical expressions explaining how this coexistence emerging from stage structure can occur. Second, we show using a community-level sensitivity analysis that such emergent coexistence is robust to perturbations of parameter values. Finally, we ask whether these results extend from two to many species, using simulations. We show that they do not, as coexistence emerging from stage structure is only seen for very similar life-history parameters. Such emergent coexistence is therefore not likely to be a key mechanism of coexistence in very diverse ecosystems, although it may contribute to explaining coexistence of certain pairs of intensely competing species.

Combining Climatic and Genomic Data Improves Range-Wide Tree Height Growth Prediction in a Forest Tree.

AbstractPopulation response functions based on climatic and phenotypic data from common gardens have long been the gold standard for predicting quantitative trait variation in new environments. However, prediction accuracy might be enhanced by incorporating genomic information that captures the neutral and adaptive processes behind intrapopulation genetic variation. We used five clonal common gardens containing 34 provenances (523 genotypes) of maritime pine (Pinus pinaster Aiton) to determine whether models combining climatic and genomic data capture the underlying drivers of height growth variation and thus improve predictions at large geographical scales. The plastic component explained most of the height growth variation, probably resulting from population responses to multiple environmental factors. The genetic component stemmed mainly from climate adaptation and the distinct demographic and selective histories of the different maritime pine gene pools. Models combining climate of origin and gene pool of the provenances as well as height-associated positive-effect alleles (PEAs) captured most of the genetic component of height growth and better predicted new provenances compared with the climate-based population response functions. Regionally selected PEAs were better predictors than globally selected PEAs, showing high predictive ability in some environments even when included alone in the models. These results are therefore promising for the future use of genome-based prediction of quantitative traits.

Looking for compensation at multiple scales in a wetland bird community.

Compensatory dynamics, during which community composition shifts despite a near-constant total community size, are usually rare: Synchronous dynamics prevail in natural communities. This is a puzzle for ecologists, because of the key role of compensation in explaining the relation between biodiversity and ecosystem functioning. However, most studies so far have considered compensation in either plants or planktonic organisms, so that evidence for the generality of such synchrony is limited. Here, we extend analyses of community-level synchrony to wetland birds. We analyze a 35-year monthly survey of a community where we suspected that compensation might occur due to potential competition and changes in water levels, favoring birds with different habitat preferences. We perform both year-to-year analyses by season, using a compensation/synchrony index, and multiscale analyses using a wavelet-based measure, which allows for both scale- and time-dependence. We analyze synchrony both within and between guilds, with guilds defined either as tightknit phylogenetic groups or as larger functional groups. We find that abundance and biomass compensation are rare, likely due to the synchronizing influence of climate (and other drivers) on birds, even after considering several temporal scales of covariation (during either cold or warm seasons, above or below the annual scale). Negative covariation in abundance at the guild or community level did only appear at the scale of a few months or several years. We also found that synchrony varies with taxonomic and functional scale: The rare cases where compensation appeared consistently in year-to-year analyses were between rather than within functional groups. Our results suggest that abundance compensation may have more potential to emerge between broad functional groups rather than between species, and at relatively long temporal scales (multiple years for vertebrates), above that of the dominant synchronizing driver.

Seed banks can help to maintain the diversity of interacting phytoplankton species.

Seed formation is part of the reproductive cycle, leading to the accumulation of resistance stages that can withstand harsh environmental conditions for long periods of time. At the community level, multiple species with such long-lasting life stages can be more likely to coexist. While the implications of this process for biodiversity have been studied in terrestrial plants, seed banks are usually neglected in phytoplankton multispecies dynamic models, in spite of widespread empirical evidence for such seed banks. In this study, we build a metacommunity model of interacting phytoplankton species, including a resting stage supplying the seed bank. The model is parameterized with empirically-driven growth rate functions and field-based interaction estimates, which include both facilitative and competitive interactions. Exchanges between compartments (coastal pelagic cells, coastal resting cells on the seabed, and open ocean pelagic cells) are controlled by hydrodynamical parameters to which the sensitivity of the model is assessed. We consider two models, i.e., with and without a saturating effect of the interactions on the growth rates. Our results are consistent between models, and show that a seed bank allows to maintain all species in the community over 30 years. Indeed, a fraction of the species are vulnerable to extinction at specific times within the year, but this process is buffered by their survival in their resting stage. We thus highlight the potential role of the seed bank in the recurrent re-invasion of the coastal community, and of coastal environments in re-seeding oceanic regions. Moreover, the seed bank enables populations to tolerate stronger interactions within the community as well as more severe changes to the environment, such as those predicted in a climate change context. Our study therefore shows how resting stages may help phytoplanktonic diversity maintenance.

Survival rates of adult and juvenile gyrfalcons in Iceland: estimates and drivers.

Knowledge of survival rates and their potential covariation with environmental drivers, for both adults and juveniles, is paramount to forecast the population dynamics of long-lived animals. Long-lived bird and mammal populations are indeed very sensitive to change in survival rates, especially that of adults. Here we report the first survival estimates for the Icelandic gyrfalcon (Falco rusticolus) obtained by capture-mark-recapture methods. We use a mark-recapture-recovery model combining live and dead encounters into a unified analysis, in a Bayesian framework. Annual survival was estimated at 0.83 for adults and 0.40 for juveniles. Positive effects of main prey density on juvenile survival (5% increase in survival from min to max density) were possible though not likely. Weather effects on juvenile survival were even less likely. The variability in observed lifespan suggests that adult birds could suffer from human-induced alteration of survival rates.

Fitting stochastic predator-prey models using both population density and kill rate data.

Most mechanistic predator-prey modelling has involved either parameterization from process rate data or inverse modelling. Here, we take a median road: we aim at identifying the potential benefits of combining datasets, when both population growth and predation processes are viewed as stochastic. We fit a discrete-time, stochastic predator-prey model of the Leslie type to simulated time series of densities and kill rate data. Our model has both environmental stochasticity in the growth rates and interaction stochasticity, i.e., a stochastic functional response. We examine what the kill rate data brings to the quality of the estimates, and whether estimation is possible (for various time series lengths) solely with time series of population counts or biomass data. Both Bayesian and frequentist estimation are performed, providing multiple ways to check model identifiability. The Fisher Information Matrix suggests that models with and without kill rate data are all identifiable, although correlations remain between parameters that belong to the same functional form. However, our results show that if the attractor is a fixed point in the absence of stochasticity, identifying parameters in practice requires kill rate data as a complement to the time series of population densities, due to the relatively flat likelihood. Only noisy limit cycle attractors can be identified directly from population count data (as in inverse modelling), although even in this case, adding kill rate data - including in small amounts - can make the estimates much more precise. Overall, we show that under process stochasticity in interaction rates, interaction data might be essential to obtain identifiable dynamical models for multiple species. These results may extend to other biotic interactions than predation, for which similar models combining interaction rates and population counts could be developed.

Within Reach? Habitat Availability as a Function of Individual Mobility and Spatial Structuring.

Organisms need access to particular habitats for their survival and reproduction. However, even if all necessary habitats are available within the broader environment, they may not all be easily reachable from the position of a single individual. Many species distribution models consider populations in environmental (or niche) space, hence overlooking this fundamental aspect of geographical accessibility. Here, we develop a formal way of thinking about habitat availability in environmental spaces by describing how limitations in accessibility can cause animals to experience a more limited or simply different mixture of habitats than those more broadly available. We develop an analytical framework for characterizing constrained habitat availability based on the statistical properties of movement and environmental autocorrelation. Using simulation experiments, we show that our general statistical representation of constrained availability is a good approximation of habitat availability for particular realizations of landscape-organism interactions. We present two applications of our approach, one to the statistical analysis of habitat preference (using step-selection functions to analyze harbor seal telemetry data) and a second that derives theoretical insights about population viability from knowledge of the underlying environment. Analytical expressions for habitat availability, such as those we develop here, can yield gains in analytical speed, biological realism, and conceptual generality by allowing us to formulate models that are habitat sensitive without needing to be spatially explicit.

From winter to summer and back: Lessons from the parameterization of a seasonal food web model for the Bialowieza forest.

Dynamic food web models describe how species abundances change over time as a function of trophic and life-history parameters. They are essential to predicting the response of ecosystems to perturbations. However, they are notoriously difficult to parameterize, so that most models rely heavily either on allometric scaling of parameters or inverse estimation of biomass flows. The allometric approach makes species of comparable body mass have near-identical parameters which can generate extinctions within a trophic level. The biomass flow approach is more precise, but is restricted to steady-states, which is not appropriate for time-varying environments. Adequately parameterizing large food webs of temperate and arctic environments requires dealing both with many species of similar sizes and a strongly seasonal environment. Inspired by the rich empirical knowledge on the vertebrate food web of the Bialowieza forest, we parameterize a bipartite food web model comprising 21 predators and 124 prey species. Our model is a non-autonomous coupled ordinary differential equations system that allows for seasonality in life-history and predation parameters. Birth and death rates, seasonal descriptions of diet for each species, food requirements and biomass information are combined into a seasonal parameterization of a dynamic food web model. Food web seasonality is implemented with time-varying intrinsic growth rate and interaction parameters, while predation is modelled with both type I and type II functional responses. All our model variants allow for >80% persistence in spite of massive apparent competition, and a quantitative match to observed (seasonal) biomasses. We also identify trade-offs between maximizing persistence, reproducing observed biomasses, and ensuring model robustness to sampling errors. Although multi-annual cycles are expected with type II functional responses, they are here prevented by a strong predator self-regulation. We discuss these results and possible improvements on the model. We provide a general workflow to parameterize dynamic food web models in seasonal environments, based on a real case study. This may help to better predict how biodiverse food webs respond to changing environments.

The effect of seasonal strength and abruptness on predator-prey dynamics.

Coupled dynamical systems in ecology are known to respond to the seasonal forcing of their parameters with multiple dynamical behaviours, ranging from seasonal cycles to chaos. Seasonal forcing is predominantly modelled as a sine wave. However, the transition between seasons is often more sudden as illustrated by the effect of snow cover on predation success. A handful of studies have mentioned the robustness of their results to the shape of the forcing signal but did not report any detailed analyses. Therefore, whether and how the shape of seasonal forcing could affect the dynamics of coupled dynamical systems remains unclear, while abrupt seasonal transitions are widespread in ecological systems. To provide some answers, we conduct a numerical analysis of the dynamical response of predator-prey communities to the shape of the forcing signal by exploring the joint effect of two features of seasonal forcing: the magnitude of the signal, which is classically the only one studied, and the shape of the signal, abrupt or sinusoidal. We consider both linear and saturating functional responses, and focus on seasonal forcing of the predator's discovery rate, which fluctuates with changing environmental conditions and prey's ability to escape predation. Our numerical results highlight that a more abrupt seasonal forcing mostly alters the magnitude of population fluctuations and triggers period-doubling bifurcations, as well as the emergence of chaos, at lower forcing strength than for sine waves. Controlling the variance of the forcing signal mitigates this trend but does not fully suppress it, which suggests that the variance is not the only feature of the shape of seasonal forcing that acts on community dynamics. Although theoretical studies may predict correctly the sequence of bifurcations using sine waves as a representation of seasonality, there is a rationale for applied studies to implement as realistic seasonal forcing as possible to make precise predictions of community dynamics.

Predator-prey feedback in a gyrfalcon-ptarmigan system?

Specialist predators with oscillating dynamics are often strongly affected by the population dynamics of their prey, yet they are not always the cause of prey cycling. Only those that exert strong (delayed) regulation of their prey can be. Inferring predator-prey coupling from time series therefore requires contrasting models with top-down versus bottom-up predator-prey dynamics. We study here the joint dynamics of population densities of the Icelandic gyrfalcon Falco rusticolus, and its prey, the rock ptarmigan Lagopus muta. The dynamics of both species are likely not only linked to each other but also to stochastic weather variables acting as confounding factors. We infer the degree of coupling between populations, as well as forcing by abiotic variables, using multivariate autoregressive models MAR(p), with p = 1 and 2 time lags. MAR(2) models, allowing for species to cycle independently from each other, further suggest alternative scenarios where a cyclic prey influences its predator but not the other way around (i.e., bottom-up scenarios). The classical MAR(1) model predicts that the time series exhibit predator-prey feedback (i.e., reciprocal dynamic influence between prey and predator), and that weather effects are weak and only affecting the gyrfalcon population. Bottom-up MAR(2) models produced a better fit but less realistic cross-correlation patterns. Simulations of MAR(1) and MAR(2) models further demonstrate that the top-down MAR(1) models are more likely to be misidentified as bottom-up dynamics than vice versa. We therefore conclude that predator-prey feedback in the gyrfalcon-ptarmigan system is likely the main cause of observed oscillations, though bottom-up dynamics cannot yet be excluded with certainty. Overall, we showed how to make more out of ecological time series by using simulations to gauge the quality of model identification, and paved the way for more mechanistic modeling of this system by narrowing the set of important biotic and abiotic drivers.

Moving forward in circles: challenges and opportunities in modelling population cycles.

Population cycling is a widespread phenomenon, observed across a multitude of taxa in both laboratory and natural conditions. Historically, the theory associated with population cycles was tightly linked to pairwise consumer-resource interactions and studied via deterministic models, but current empirical and theoretical research reveals a much richer basis for ecological cycles. Stochasticity and seasonality can modulate or create cyclic behaviour in non-intuitive ways, the high-dimensionality in ecological systems can profoundly influence cycling, and so can demographic structure and eco-evolutionary dynamics. An inclusive theory for population cycles, ranging from ecosystem-level to demographic modelling, grounded in observational or experimental data, is therefore necessary to better understand observed cyclical patterns. In turn, by gaining better insight into the drivers of population cycles, we can begin to understand the causes of cycle gain and loss, how biodiversity interacts with population cycling, and how to effectively manage wildly fluctuating populations, all of which are growing domains of ecological research.

Continental-scale travelling waves in forest geometrids in Europe: an evaluation of the evidence.

A recent paper claims the existence of one of the most large-scale travelling waves ever recorded for any animal population. Here we address why conceptual and methodological pitfalls may have served to exaggerate or even impose the spatial patterns reported. Photo credit: Jane U. Jepsen.

Indirect effects of primary prey population dynamics on alternative prey.

We develop a theory of generalist predation showing how alternative prey species are affected by changes in both mean abundance and variability (coefficient of variation) of their predator's primary prey. The theory is motivated by the indirect effects of cyclic rodent populations on ground-breeding birds, and developed through progressive analytic simplifications of an empirically-based model. It applies nonetheless to many other systems where primary prey have fast life-histories and can become superabundant, thus facilitating impact on alternative prey species and generating highly asymmetric interactions. Our results suggest that predator effects on alternative prey should generally decrease with mean primary prey abundance, and increase with primary prey variability (low to high CV)-unless predators have strong aggregative responses, in which case these results can be reversed. Approximations of models including predator dynamics (general numerical response with possible delays) confirm these results but further suggest that negative temporal correlation between predator and primary prey is harmful to alternative prey. Finally, we find that measurements of predator numerical responses are crucial to predict-even qualitatively-the response of ecosystems to changes in the dynamics of outbreaking prey species.

The intensity of horizontal and vertical search in a diving forager: the harbour seal.

BACKGROUND: Free ranging foraging animals can vary their searching intensity in response to the profitability of the environment by modifying their movements. Marine diving animals forage in a three dimensional space and searching intensity can be varied in both the horizontal and vertical planes. Therefore understanding the relationship between the allocation of searching effort in these two spaces can provide a better understanding of searching strategies and a more robust identification of foraging behaviour from the multitude of foraging indices (FIs) available. We investigated the movement of a widespread marine coastal predator, the harbour seal (Phoca vitulina), and compared two sets of foraging indices reflecting searching intensity respectively in the horizontal plane (displacement speed, extensive vs. intensive movement types, residence time) and in the vertical dimension (time at the bottom of a dive). We then tested how several factors (dive depth, direction of the trip with respect to haul-out site, different predatory tactics, the presence of factors confounding the detection of foraging, and temporal resolution of the data) affected their relationships. RESULTS: Overall the indices only showed a very weak positive correlation across the two spaces. However controlling for various factors strengthened the relationships. Resting at sea, a behaviour intrinsically static in the horizontal plane, was found to be strongly negatively related to the time spent at the bottom of the dives, indirectly weakening the relationship between horizontal and vertical foraging indices. Predatory tactic (benthic vs. pelagic) was found to directly affect the relationship. In benthic (as opposed to pelagic) foraging a stronger positive relationship was found between vertical and horizontal indices. CONCLUSIONS: Our results indicated that movement responses, leading to an intensification of search, are similar in the two spaces (positive relationship), but additional factors need to be taken into account for this relationship to emerge. Foraging indices measuring residence in the horizontal plane tend to be inflated by resting events at sea, while vertical indices tend to distinguish mainly between periods of activity and inactivity, or of benthic and pelagic foraging. The simultaneous consideration of horizontal and vertical movements, as well as topographic information, allows additional behavioural states to be inferred, providing greater insight into the interpretation of foraging activity.

Overcompensation and phase effects in a cyclic common vole population: between first and second-order cycles.

Population cycles in voles are often thought to be generated by one-year delayed density dependence on the annual population growth rate. In common voles, however, it has been suggested by Turchin (2003) that some populations exhibit first-order cycles, resulting from strong overcompensation (i.e. carrying capacity overshoots in peak years, with only an effect of the current year abundance on annual growth rates). We focus on a common vole (Microtus arvalis) population from western France that exhibits 3-year cycles. Several overcompensating nonlinear models for populations dynamics are fitted to the data, notably those of Hassell, and Maynard-Smith and Slatkin. Overcompensating direct density dependence (DD) provides a satisfactory description of winter crashes, and one-year delayed density dependence is not responsible for the crashes, thus these are not classical second-order cycles. A phase-driven modulation of direct density dependence maintains a low-phase, explaining why the cycles last three years instead of two. Our analyses suggest that some of this phase dependence can be expressed as one-year delayed DD, but phase dependence provides a better description. Hence, modelling suggests that cycles in this population are first-order cycles with a low phase after peaks, rather than fully second-order cycles. However, based on the popular log-linear second-order autoregressive model, we would conclude only that negative delayed density dependence exists. The additive structure of this model cannot show when delayed DD occurs (here, during lows rather than peaks). Our analyses thus call into question the automated use of second-order log-linear models, and suggests that more attention should be given to non-(log)linear models when studying cyclic populations. From a biological viewpoint, the fast crashes through overcompensation that we found suggest they might be caused by parasites or food rather than predators, though predators might have a role in maintaining the low phase and spatial synchrony.