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Integrated population models (IPMs) combine multiple ecological data types such as capture-mark-recapture histories, reproduction surveys, and population counts into a single statistical framework. In such models, each data type is generated by a probabilistic submodel, and an assumption of independence between the different data types is usually made. The fact that the same biological individuals can contribute to multiple data types has been perceived as affecting their independence, and several studies have even investigated IPM robustness in this scenario. However, what matters from a statistical perspective is probabilistic independence: the joint probability of observing all data is equal to the product of the likelihoods of the various datasets. Contrary to a widespread perception, probabilistic non-independence does not automatically result from collecting data on the same physical individuals. Conversely, while there can be good reasons for non-independence of IPM submodels arising from sharing of individuals between data types, these relations do not seem to be included in IPMs whose robustness is being investigated. Furthermore, conditional rather than true independence is sometimes assumed. In this conceptual paper, I survey the various independence concepts used in IPMs, try to make sense of them by getting back to first principles in toy models, and show that it is possible to obtain probabilistic independence (or near-independence) despite two or three data types collected on the same set of biological individuals. I then revisit recommendations pertaining to component data collection and IPM robustness checks, and provide some suggestions to bridge the current gap between individual-level IPMs and their population-level approximations using composite likelihoods.

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., Comment: 51 pages; 16 figures

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 Fussman & 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 linearised 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 ways to evaluate structural sensitivity in a stochastic setting, for predator-prey or other ecological systems.

In a recent opinion article, Muff et al. recapitulate well-known objections to the Neyman-Pearson Null-Hypothesis Significance Testing (NHST) framework and call for reforming our practices in statistical reporting. We agree with them on several important points: the significance threshold P<0.05 is only a convention, chosen as a compromise between type I and II error rates; transforming the p-value into a dichotomous statement leads to a loss of information; and p-values should be interpreted together with other statistical indicators, in particular effect sizes and their uncertainty. In our view, a lot of progress in reporting results can already be achieved by keeping these three points in mind. We were surprised and worried, however, by Muff et al.'s suggestion to interpret the p-value as a "gradual notion of evidence". Muff et al. recommend, for example, that a P-value > 0.1 should be reported as "little or no evidence" and a P-value of 0.001 as "strong evidence" in favor of the alternative hypothesis H1., Comment: This is a letter in response to Muff, S., Nilsen, E. B., O'Hara, R. B., & Nater, C. R. (2022). Rewriting results sections in the language of evidence. Trends in Ecology & Evolution

The Traill island model of Gilg et al. (2003) is a landmark attempt at mechanistic modelling of the cyclic population dynamics of rodents, focusing on a high Arctic community. It models the dynamics of one prey, the collared lemming, and four predators : the stoat, the Arctic fox, the long-tailed skua and the snowy owl. In the present short note, we first summarize how the model works in light of theory on seasonally forced predator-prey systems, with a focus on the temporal dynamics of predation rates. We show notably how the impact of generalist predation, which is able here to initiate population declines, differs slightly from that of generalist predation in other mechanistic models of rodent-mustelid interactions such as Turchin & Hanski (1997). We then provide a low-dimensional approximation with a single generalist predator compartment that mimics the essential features of the Traill island model: cycle periodicity, amplitude, shape, as well as generalist-induced declines. This simpler model should be broadly applicable to model other lemming populations that predominantly grow under the snow during the winter period. Matlab computer codes for Gilg et al. (2003), its two-dimensional approximation, as well as alternative lemming population dynamics models are provided.

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.

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., Comment: 46 pages ; 13 figures

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 but the transition between seasons is often more sudden. Some studies mentioned the robustness of their results to the shape of the forcing signal, without detailed analyses. Therefore, whether and how the shape of seasonal forcing affects 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.

Identifying directed interactions between species from time series of their population densities has many uses in ecology. This key statistical task is equivalent to causal time series inference, which connects to the Granger causality (GC) concept: $x$ causes $y$ if $x$ improves the prediction of $y$ in a dynamic model. However, the entangled nature of nonlinear ecological systems has led to question the appropriateness of Granger causality, especially in its classical linear Multivariate AutoRegressive (MAR) model form. Convergent-cross mapping (CCM), a nonparametric method developed for deterministic dynamical systems, has been suggested as an alternative. Here, we show that linear GC and CCM are able to uncover interactions with surprisingly similar performance, for predator-prey cycles, 2-species deterministic (chaotic) or stochastic competition, as well as 10- and 20-species interaction networks. There is no correspondence between the degree of nonlinearity of the dynamics and which method performs best. Our results therefore imply that Granger causality, even in its linear MAR($p$) formulation, is a valid method for inferring interactions in nonlinear ecological networks; using GC or CCM (or both) can instead be decided based on the aims and specifics of the analysis.

Explaining coexistence in species-rich communities of primary producers remains a challenge for ecologists because of their likely competition for shared resources. Following Hutchinson's seminal suggestion, many theoreticians have tried to create diversity through a fluctuating environment, which impairs or slows down competitive exclusion. However, fluctuating-environment models often only produce a dozen of coexisting species at best. Here, we investigate how to create richer communities in fluctuating environments, using an empirically parameterized model. Building on the forced Lotka-Volterra model of Scranton and Vasseur (Theor Ecol 9(3):353-363, 2016), inspired by phytoplankton communities, we have investigated the effect of two coexistence mechanisms, namely the storage effect and higher intra- than interspecific competition strengths (i.e., strong self-regulation). We tuned the intra/inter competition ratio based on empirical analyses, in which self-regulation dominates interspecific interactions. Although a strong self-regulation maintained more species (50%) than the storage effect (25%), we show that none of the two coexistence mechanisms considered could ensure the coexistence of all species alone. Realistic seasonal environments only aggravated that picture, as they decreased persistence relative to a random environment. However, strong self-regulation and the storage effect combined superadditively so that all species could persist with both mechanisms at work. Our results suggest that combining different coexistence mechanisms into community models might be more fruitful than trying to find which mechanism best explains diversity. We additionally highlight that while biomass-trait distributions provide some clues regarding coexistence mechanisms, they cannot indicate unequivocally which mechanisms are at play., Comment: 27 pages, 9 figures, Theor Ecol (2019)

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.

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 locally superabundant, which facilitates impact on alternative prey species. In contrast to classic apparent competition theory based on symmetric 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. We find in general that predator numerical responses are crucial to predict the response of ecosystems to changes in key prey species exhibiting outbreaks, and extend the apparent competition/mutualism theory to asymmetric interactions.