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Abstract Many animals show avoidance behavior in response to disease. For instance, in some species of frogs, individuals that survive infection of the fungal disease chytridiomycosis may learn to avoid areas where the pathogen is present. As chytridiomycosis has caused substantial declines in many amphibian populations worldwide, it is a highly relevant example for studying these behavioral dynamics. Here we develop compartmental ODE models to study the epidemiological consequences of avoidance behavior of animals in response to waterborne infectious diseases. Individuals with avoidance behavior are less likely to become infected, but avoidance may also entail increased risk of mortality. We compare the outbreak dynamics with avoidance behavior that is innate (present from birth) or learned (gained after surviving infection). We also consider how management to induce learned avoidance might affect the resulting dynamics. Using methods from dynamical systems theory, we calculate the basic reproduction number$$R_0$$for each model, analyze equilibrium stability of the systems, and perform a detailed bifurcation analysis. We show that disease persistence when$$R_0< 1$$is possible with learned avoidance, but not with innate avoidance. Our results imply that management to induce behavioral avoidance can actually cause such a scenario, but it is also less likely to occur for high-mortality diseases (e.g., chytridiomycosis). Furthermore, the learned avoidance model demonstrates a variety of codimension-1 and -2 bifurcations not found in the innate avoidance model. Simulations with parameters based on chytridiomycosis are used to demonstrate these features and compare the outcomes with innate, learned, and no avoidance behavior. 

Abstract Integral projection models (IPMs) are widely used for studying continuously size‐structured populations. IPMs require a growth sub‐model that describes the probability of future size conditional on current size and any covariates. Most IPM studies assume that this distribution is Gaussian, despite calls for non‐Gaussian models that accommodate skewness and excess kurtosis. We provide a general workflow for accommodating non‐Gaussian growth patterns while retaining important covariates and random effects. Our approach emphasizes visual diagnostics from pilot Gaussian models and quantile‐based metrics of skewness and kurtosis that guide selection of a non‐Gaussian alternative, if necessary. Across six case studies, skewness and excess kurtosis were common features of growth data, and non‐Gaussian models consistently generated simulated data that were more consistent with real data than pilot Gaussian models. However, effects of “improved” growth modeling on IPM results were moderate to weak and differed in direction or magnitude between different outputs from the same model. Using tools not available when IPMs were first developed, it is now possible to fit non‐Gaussian models to growth data without sacrificing ecological complexity. Doing so, as guided by careful interrogation of the data, will result in models that better represent the populations for which they are intended. 

Species distribution models (SDMs) are frequently data-limited. In aquatic habitats, emerging environmental DNA (eDNA) sampling methods can be quicker and more cost-efficient than traditional count and capture surveys, but their utility for fitting SDMs is complicated by dilution, transport, and loss processes that modulate DNA concentrations and mix eDNA from different locations. Past models for estimating organism densities from measured species-specific eDNA concentrations have accounted for how these processes affect expected concentrations. We built off this previous work to construct a linear hierarchical model that also accounts for how they give rise to spatially correlated concentration errors. We applied our model to 60 simulated stream networks and three types of species niches in order to answer two questions: 1) what is the D-optimal sampling design, i.e. where should eDNA samples be positioned to most precisely estimate species–environment relationships? and 2) How does parameter estimation accuracy depend on the stream network’s topological and hydrologic properties? We found that correcting for eDNA dynamics was necessary to obtain consistent parameter estimates, and that relative to a heuristic benchmark design, optimizing sampling locations improved design efficiency by an average of 41.5%. Samples in the D-optimal design tended to be positioned near downstream ends of stream reaches high in the watershed, where eDNA concentration was high and mostly from homogeneous source areas, and they collectively spanned the full ranges of covariates. When measurement error was large, it was often optimal to collect replicate samples from high-information reaches. eDNA-based estimates of species–environment regression parameters were most precise in stream networks that had many reaches, large geographic size, slow flows, and/or high eDNA loss rates. Our study demonstrates the importance and viability of accounting for eDNA dilution, transport, and loss in order to optimize sampling designs and improve the accuracy of eDNA-based species distribution models. 

In many species, a few individuals produce most of the next generation. How much of this reproductive skew is driven by variation among individuals in fixed traits, how much by external factors, and how much by random chance? And what does it take to have truly exceptional lifetime reproductive output (LRO)? In the past, we and others have partitioned the variance of LRO as a proxy for reproductive skew. Here we explain how to partition LRO skewness itself into contributions from fixed trait variation, four forms of “demographic luck” (birth state, fecundity luck, survival trajectory luck, and growth trajectory luck), and two kinds of “environmental luck” (birth environment and environment trajectory). Each of these is further partitioned into contributions at different ages.We also determine what we can infer about individuals with exceptional LRO. We find that reproductive skew is largely driven by random variation in lifespan, and exceptional LRO generally results from exceptional lifespan. Other kinds of luck frequently bring skewness down rather than increasing it. In populations where fecundity varies greatly with environmental conditions, getting a good year at the right time can be an alternate route to exceptional LRO, so that LRO is less predictive of lifespan. 

Abstract: One strand of modern coexistence theory (MCT) partitions invader growth rates (IGR) to quantify how different mechanisms contribute to species coexistence, highlighting fluctuation‐dependent mechanisms. A general conclusion from the classical analytic MCT theory is that coexistence mechanisms relying on temporal variation (such as the temporal storage effect) are generally less effective at promoting coexistence than mechanisms relying on spatial or spatiotemporal variation (primarily growth‐density covariance). However, the analytic theory assumes continuous population density, and IGRs are calculated for infinitesimally rare invaders that have infinite time to find their preferred habitat and regrow, without ever experiencing intraspecific competition. Here we ask if the disparity between spatial and temporal mechanisms persists when individuals are, instead, discrete and occupy finite amounts of space. We present a simulation‐based approach to quantifying IGRs in this situation, building on our previous approach for spatially non‐varying habitats. As expected, we found that spatial mechanisms are weakened; unexpectedly, the contribution to IGR from growth‐density covariance could even become negative, opposing coexistence. We also found shifts in which demographic parameters had the largest effect on the strength of spatial coexistence mechanisms. Our substantive conclusions are statements about one model, across parameter ranges that we subjectively considered realistic. Using the methods developed here, effects of individual discreteness should be explored theoretically across a broader range of conditions, and in models parameterized from empirical data on real communities. 

Species distribution models (SDMs) are frequently data‐limited. In aquatic habitats, emerging environmental DNA (eDNA) sampling methods can be quicker and more cost‐efficient than traditional count and capture surveys, but their utility for fitting SDMs is complicated by dilution, transport, and loss processes that modulate DNA concentrations and mix eDNA from different locations. Past models for estimating organism densities from measured species‐specific eDNA concentrations have accounted for how these processes affect expected concentrations. We built off this previous work to construct a linear hierarchical model that also accounts for how they give rise to spatially correlated concentration errors. We applied our model to 60 simulated stream networks and three types of species niches in order to answer two questions: 1) what is the D‐optimal sampling design, i.e. where should eDNA samples be positioned to most precisely estimate species–environment relationships? and 2) How does parameter estimation accuracy depend on the stream network's topological and hydrologic properties? We found that correcting for eDNA dynamics was necessary to obtain consistent parameter estimates, and that relative to a heuristic benchmark design, optimizing sampling locations improved design efficiency by an average of 41.5%. Samples in the D‐optimal design tended to be positioned near downstream ends of stream reaches high in the watershed, where eDNA concentration was high and mostly from homogeneous source areas, and they collectively spanned the full ranges of covariates. When measurement error was large, it was often optimal to collect replicate samples from high‐information reaches. eDNA‐based estimates of species–environment regression parameters were most precise in stream networks that had many reaches, large geographic size, slow flows, and/or high eDNA loss rates. Our study demonstrates the importance and viability of accounting for eDNA dilution, transport, and loss in order to optimize sampling designs and improve the accuracy of eDNA‐based species distribution models. 

Abstract Spatial environmental heterogeneity maintains species diversity in ecological communities, but it is difficult to model and its effects on individuals are often ignored in empirical studies of species coexistence. This is problematic because many studies estimate plant competition at the neighbourhood level, and modelling individual demographic responses without neighbourhood‐level environmental data could lead to biased results. We aimed to understand how and when ignoring spatial heterogeneity produces biased estimates of the strength of intraspecific and interspecific competition and, therefore, incorrect predictions about species coexistence.First, we used a spatially explicit individual‐based model (IBM), with neighbourhood competition depending on neighbour size and distance, to simulate two plant populations that coexist via differing responses to spatial heterogeneity. Next, we mimicked the action of ecologists collecting data in observational studies or short‐term experimental studies and sampled from the simulated populations. Then, we fitted demographic models for each species including parameters for the intensity of intra‐ and interspecific competitionwithoutinformation about spatial heterogeneity. Finally, we simulated invasions of species at low density using the IBM and estimated parameters to compute invasion growth rates (IGR).Demographic modelling with data from simulated observational studies resulted in underestimates of intraspecific competition and overestimates of interspecific competition. When we simulated invasions with the IBM and those estimated competition coefficients, the IGRs were negative or around zero and incorrectly predicted competitive exclusion. In simulated experimental studies, the estimates of competition were more accurate, but the model still incorrectly predicted competitive exclusion.Synthesis: Our study demonstrates how biases can arise in neighbourhood competition studies and highlights the importance of explicitly incorporating spatial heterogeneity into empirical coexistence studies. Failure to account for environmental heterogeneity at the level of individual plants may lead to mistaken conclusions about coexistence outcomes. 

Abstract: Chance pervades life. In turn, life histories are described by probabilities (e.g. survival and breeding) and averages across individuals (e.g. mean growth rate and age at maturity). In this study, we explored patterns of luck in lifetime outcomes by analysing structured population models for a wide array of plant and animal species. We calculated four response variables: variance and skewness in both lifespan and lifetime reproductive output (LRO), and partitioned them into contributions from different forms of luck. We examined relationships among response variables and a variety of life history traits. We found that variance in lifespan and variance in LRO were positively correlated across taxa, but that variance and skewness were negatively correlated for both lifespan and LRO. The most important life history trait was longevity, which shaped variance and skew in LRO through its effects on variance in lifespan. We found that luck in survival, growth, and fecundity all contributed to variance in LRO, but skew in LRO was overwhelmingly due to survival luck. Rapidly growing populations have larger variances in LRO and lifespan than shrinking populations. Our results indicate that luck‐induced genetic drift may be most severe in recovering populations of species with long mature lifespan and high iteroparity. 

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