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How organisms gather and utilize information about their landscapes is central to understanding land-use patterns and population distributions. When such information originates beyond an individual's immediate vicinity, movement decisions require integrating information out to some perceptual range. Such nonlocal information, whether obtained visually, acoustically, or via chemosensation, provides a field of stimuli that guides movement. Classically, however, models have assumed movement based on purely local information (e.g., chemotaxis, step-selection functions). Here we explore how foragers can exploit nonlocal information to improve their success in dynamic landscapes. Using a continuous time/continuous space model in which we vary both random (diffusive) movement and resource-following (advective) movement, we characterize the optimal perceptual ranges for foragers in dynamic landscapes. Nonlocal information can be highly beneficial, increasing the spatiotemporal concentration of foragers on their resources up to twofold compared with movement based on purely local information. However, nonlocal information is most useful when foragers possess both high advective movement (allowing them to react to transient resources) and low diffusive movement (preventing them from drifting away from resource peaks). Nonlocal information is particularly beneficial in landscapes with sharp (rather than gradual) patch edges and in landscapes with highly transient resources.

Climate change drives uneven phenology shifts across taxa, and this can result in changes to the phenological match between interacting species. Shifts in the relative phenology of partner species are well documented, but few studies have addressed the effects of such changes on population dynamics. To explore this, we develop a phenologically explicit model describing consumer-resource interactions. Focusing on scenarios for univoltine insects, we show how changes in resource phenology can be reinterpreted as transformations in the year-to-year recursion relationships defining consumer population dynamics. This perspective provides a straightforward path for interpreting the long-term population consequences of phenology change. Specifically, by relating the outcome of phenological shifts to species traits governing recursion relationships (e.g., consumer fecundity or competitive scenario), we demonstrate how changes in relative phenology can force systems into different dynamical regimes, with major implications for resource management, conservation, and other areas of applied dynamics.

Habitat loss and climate change jointly threaten a large fraction of earth's biodiversity. A key goal is to understand how these threats play out differentially across species. Focusing on insects that undergo an ontogenetic shift in habitat requirements, we use critical patch size models to examine how breeding strategy influences the abilities of different kinds of species to persist in small habitat patches. In general, we find that income breeders require larger habitat patches for population persistence than do capital breeders. However, increases in patch size requirements as a result of factors that limit oviposition (e.g., resource availability, weather conditions) are more severe for capital breeders than for income breeders. From a conservation perspective, our work suggests that a species' sensitivity to habitat loss, both today and in the future, can depend critically on evolved behavioral strategies. Explicit consideration of such behavioral strategies, including a careful accounting of their relationship with dispersal and survival, provides a map linking life-history spectra, spatial requirements, and management.

Understanding animal movement is a key challenge in ecology and conservation biology. Relocation data often represent a complex mixture of different movement behaviors, and reliably decomposing this mix into its component parts is an unresolved problem in movement ecology. Traditional approaches, such as composite random walk models, require that the timescales characterizing the movement are all similar to the usually arbitrary data-sampling rate. Movement behaviors such as long-distance searching and fine-scale foraging, however, are often intermixed but operate on vastly different spatial and temporal scales. An approach that integrates the full sweep of movement behaviors across scales is currently lacking. Here we show how the semivariance function (SVF) of a stochastic movement process can both identify multiple movement modes and solve the sampling rate problem. We express a broad range of continuous-space, continuous-time stochastic movement models in terms of their SVFs, connect them to relocation data via variogram regression, and compare them using standard model selection techniques. We illustrate our approach using Mongolian gazelle relocation data and show that gazelle movement is characterized by ballistic foraging movements on a 6-h timescale, fast diffusive searching with a 10-week timescale, and asymptotic diffusion over longer timescales.

The relative importance of plant facilitation and competition during primary succession depends on the development of ecosystem nutrient pools, yet the interaction of these processes remains poorly understood. To explore how these mechanisms interact to drive successional dynamics, we devised a stoichiometric ecosystem-level model that considers the role of nitrogen and phosphorus limitation in plant primary succession. We applied this model to the primary plant community on Mount St. Helens, Washington State, to check the validity of the proposed mechanisms. Our results show that the plant community is colimited by nitrogen and phosphorus, and they confirm previous suggestions that the presence of a nitrogen-fixing legume, Lupinus lepidus, can enhance community biomass. In addition, the observed nutrient supply rates may promote alternative successional trajectories that depend on the initial plant abundances, which may explain the observed heterogeneity in community development. The model further indicates the importance of mineralization rates and other ecosystem parameters to successional rates. We conclude that a model framework based on ecological stoichiometry allows integration of key biotic processes that interact nonlinearly with biogeochemical aspects of succession. Extension of this approach will improve the understanding of the process of primary succession and its application to ecosystem rehabilitation.

Mate finding, which is essential to both population growth and gene exchange, involves both spatial and temporal components. From a population dynamics perspective, spatial mate-finding problems are well studied, and decreased mate-finding efficiency at low population densities is a well-recognized mechanism for the Allee effect. Temporal aspects of mate finding have been rarely considered, but reproductive asynchrony may engender an Allee effect in which some females go mateless by virtue of temporal isolation. Here we develop and explore a model that unifies previously disparate theoretical considerations of spatial and temporal aspects of mate finding. Specifically, we develop a two-sex reaction-diffusion system to examine the interplay between reproductive asynchrony and the dispersal of individuals out of a patch. We also consider additional behavioral complications, including several alternative functional forms for mating efficiency and advective movements in which males actively seek out females. By calculating the fraction of females expected to go mateless as a joint function of reproductive asynchrony and patch size, we find that the population-level reproductive rates necessary to offset female matelessness may be quite high. These results suggest that Allee effects engendered by reproductive asynchrony will be greatly exacerbated in spatially isolated populations.

How biological processes such as reproduction and dispersal relate to the size of species' geographic ranges constitutes a major challenge in spatial ecology and biogeography. Here we develop a spatially explicit theoretical framework that links fundamental population-level ecological traits (e.g., rates of dispersal and population growth or decay) with landscape heterogeneity to derive estimates of species' geographic range sizes and, further, distributions of geographic range sizes across species. Although local (patch-scale) population dynamics in this model are completely deterministic, we consider a fragmented landscape of patches and gaps in which the spatial heterogeneity is itself stochastic. This stochastic spatial structure, which juxtaposes landscape-level patch and gap characteristics against population-level critical patch sizes and maximum gap-crossing abilities, determines how far a novel species can spread from its evolutionary origin. Given reasonable assumptions about landscape structure and about the distribution of critical patch sizes and critical gap lengths among species, we obtain distributions of geographic range sizes that are qualitatively similar to those routinely found in nature (e.g., many species with small geographic ranges). Collectively, our results suggest that both interspecific differences in population-level traits and the landscapes through which species spread help determine patterns of occupancy and geographic extent.

Central-place foragers, such as ants, beavers, and colonial seabirds, can act as biological conduits, subsidizing local communities with allochthonous resources. To explore the consequences of such biologically vectored resource redistribution, we draw on an example from cave ecology and develop a population-level model of central-place foraging based on the dispersal kernel framework. We explore how the size of the patch in which central-place foraging occurs and the spatial distribution of foragers within that patch feed back to influence the population dynamics of the central-place forager and the species richness of the associated recipient community. We demonstrate that the particular way in which a population of central-place foragers uses space has two important effects. First, space use determines the stability of the forager population and establishes patch size thresholds for persistence, stable equilibria, and limit cycles. Second, alternative foraging kernels lead to qualitatively different scaling relationships between the size of the foraging patch and species richness back at the central place. These analyses provide a new link among elements of ecology related to animal behavior, population dynamics, and species diversity while also providing a novel perspective on the utility of integrodifference equations for problems in spatial ecology.

Plant biomass and nutrient allocation explicitly links the evolved strategies of plant species to the material and energy cycles of ecosystems. Allocation of nitrogen (N) and phosphorus (P) is of particular interest because N and P play pivotal roles in many aspects of plant biology, and their availability frequently limits plant growth. Here we present a comparative scaling analysis of a global data compilation detailing the N and P contents of leaves, stems, roots, and reproductive structures of 1,287 species in 152 seed plant families. We find that P and N contents (as well as N : P) are generally highly correlated both within and across organs and that differences exist between woody and herbaceous taxa. Between plant organs, the quantitative form of the scaling relationship changes systematically, depending on whether the organs considered are primarily structural (i.e., stems, roots) or metabolically active (i.e., leaves, reproductive structures). While we find significant phylogenetic signals in the data, similar scaling relationships occur in independently evolving plant lineages, which implies that both the contingencies of evolutionary history and some degree of environmental convergence have led to a common set of rules that constrain the partitioning of nutrients among plant organs.

Here we study the spatial dynamics of a coinvading consumer-resource pair. We present a theoretical treatment with extensive empirical data from a long-studied field system in which native herbivorous insects attack a population of lupine plants recolonizing a primary successional landscape created by the 1980 volcanic eruption of Mount St. Helens. Using detailed data on the life history and interaction strengths of the lupine and one of its herbivores, we develop a system of integrodifference equations to study plant-herbivore invasion dynamics. Our analyses yield several new insights into the spatial dynamics of coinvasions. In particular, we demonstrate that aspects of plant population growth and the intensity of herbivory under low-density conditions can determine whether the plant population spreads across a landscape or is prevented from doing so by the herbivore. In addition, we characterize the existence of threshold levels of spatial extent and/or temporal advantage for the plant that together define critical values of "invasion momentum," beyond which herbivores are unable to reverse a plant invasion. We conclude by discussing the implications of our findings for successional dynamics and the use of biological control agents to limit the spread of pest species.

Identifying linkages between life-history traits and small population processes is essential to effective multispecies conservation. Reproductive asynchrony, which occurs when individuals are reproductively active for only a portion of the population-level breeding period, may provide one such link. Traditionally, reproductive asynchrony has been considered from evolutionary perspectives as an advantageous bet-hedging strategy in temporally unpredictable environments. Here, we explore the dynamic consequences of reproductive asynchrony as a density-dependent life-history trait. To examine how asynchrony affects population growth rate and extinction risk, we used a general model of reproductive timing to quantify the temporal overlap of opposite-sex individuals and to simulate population dynamics over a range of initial densities and empirical estimates of reproductive asynchrony. We also considered how protandry, a sexually selected life-history strategy that often accompanies asynchrony, modulates the population-level effects of reproductive asynchrony. We found that asynchrony decreases the number of males a female overlaps with, decreases the average probability of mating per male/female pair that does overlap, and leaves some females completely isolated in time. This loss of reproductive potential, which is exacerbated by protandry, reduces population growth rate at low density and can lead to extinction via an Allee effect. Thus reproductive asynchrony and protandry, both of which can be evolutionarily advantageous at higher population densities, may prove detrimental when population density declines.

Disparities in nutrient content (nitrogen and phosphorus) between herbivores and their plant resources have lately proven to have major consequences for herbivore success, consumer-driven nutrient cycling, and the fate of primary production in ecosystems. Here we extend these findings by examining patterns of nutrient content between animals at higher trophic levels, specifically between insect herbivores and predators. Using a recently compiled database on insect nutrient content, we found that predators exhibit on average 15% greater nitrogen content than herbivores. This difference persists after accounting for variation from phylogeny and allometry. Among herbivorous insects, we also found evidence that recently derived lineages (e.g., herbivorous Diptera and Lepidoptera) have, on a relative basis, 15%-25% less body nitrogen than more ancient herbivore lineages (e.g., herbivorous Orthoptera and Hemiptera). We elaborate several testable hypotheses for the origin of differences in nitrogen content between trophic levels and among phylogenetic lineages. For example, interspecific variation in insect nitrogen content may be directly traceable to differences in dietary nitrogen (including dilution by gut contents), selected for directly in response to the differential scarcity of dietary nitrogen, or an indirect consequence of adaptation to different feeding habits. From some functional perspectives, the magnitude rather than the source of the interspecific differences in nitrogen content may be most critical. We conclude by discussing the implications of the observed patterns for both the trophic complexity of food webs and the evolutionary radiation of herbivorous insects.

Understanding the impact of habitat edges provides a key to deciphering how community dynamics change as functions of habitat structure and spatial scale. Motivated by studies of predation on bird nests in forest fragments and other cases of "cross-boundary subsidies," we present results from a partial differential equation model in which a patch-resident prey species suffers incidental mortality from a generalist predator species residing in the surrounding matrix habitat. We demonstrate that predator intrusions have the potential to induce critical patch size effects for the prey species, even when the prey's dynamics would otherwise preclude such effects. We also demonstrate that the existence of critical patch size effects depends on the functional response of the predator, with Lotka-Volterra and Type II functional responses generating the effect (but not Type III). We conclude by discussing how predator-induced critical patch size effects can influence opportunities for regionwide persistence of the prey by altering the fraction and spatial distribution of meaningful patches within a metapopulation.

Many species exhibit widespread spatial synchrony in population fluctuations. This pattern is of great ecological interest and can be a source of concern when the species is rare or endangered. Both dispersal and spatial correlations in the environment have been implicated as possible causes of this pattern, but these two factors have rarely been studied in combination. We develop a spatially structured population model, simple enough to obtain analytic solutions for the population correlation, that incorporates both dispersal and environmental correlation. We ask whether these two synchronizing factors contribute additively to the total spatial population covariance. We find that there is always an interaction between these two factors and that this interaction is small only when one or both of the environmental correlation and the dispersal rate are small. The interaction is opposite in sign to the environmental correlation; so, in the normal case of positive environmental correlation across sites, the population synchrony will be lower than predicted by simply adding the effects of dispersal and environmental correlation. We also find that population synchrony declines as the strength of population regulation increases. These results indicate that dispersal and environmental correlation need to be considered in combination as explanations for observed patterns of population synchrony.

Lupines (Lupinus lepidus var. lobbii), the earliest plant colonists of primary successional habitats at Mount St. Helens, were expected to strongly affect successional trajectories through facilitative effects. However, their effects remain localized because initially high rates of reinvasive spread were short lived, despite widespread habitat availability. We experimentally tested whether insect herbivores, by reducing plant growth and fecundity at the edge of the expanding lupine population, could curtail the rate of reinvasion and whether those herbivores had comparable impacts in the older, more successionally advanced core region. We found that removing insect herbivores increased both the areal growth of individual lupine plants and the production of new plants in the edge region, thereby accelerating the lupine's intrinsic rate of increase at the front of the lupine reinvasion. We found no such impacts of herbivory in the core region, where low plant quality or a complex of recently arrived natural enemies may hold herbivores in check. In the context of invasion theory, herbivore-mediated decreases in lupine population growth rate in the edge region translate into decreased rates of lupine spread, which we quantify here using diffusion models. In the Mount St. Helens system, decreased rate of lupine reinvasion will result in reductions in rates of soil formation, nitrogen input, and entrapment of seeds and detritus that are likely to postpone or alter trajectories of primary succession. If the type of spatial subtleties in herbivore effects we found here are common, with herbivory focused on the edge of an expanding plant population and suppressed or ineffective in the larger, denser central region (where the plants might be more readily noticed and studied), then insect herbivores may have stronger impacts on the dynamics of primary succession and plant invasions than previously recognized.

We examine the degree to which fitting simple dynamic models to time series of population counts can predict extinction probabilities. This is both an active branch of ecological theory and an important practical topic for resource managers. We introduce an approach that is complementary to recently developed techniques for estimating extinction risks (e.g., diffusion approximations) and, like them, requires only count data rather than the detailed ecological information available for traditional population viability analyses. Assuming process error, we use four different models of population growth to generate snapshots of population dynamics via time series of the lengths commonly available to ecologists. We then ask to what extent we can identify which of several broad classes of population dynamics is evident in the time series snapshot. Along the way, we introduce the idea of "variation thresholds," which are the maximum amount of process error that a population may withstand and still have a specified probability of surviving for a given length of time. We then show how these thresholds may be useful to both ecologists and resource managers, particularly when dealing with large numbers of poorly understood species, a common problem faced by those designing biodiversity reserves.

Traditionally, ecologists interested in habitat edges have focused on edge-related gradients in patterns of species richness or abiotic variables. Here, however, we take a different perspective, attempting to synthesize recent empirical results concerning the effects of habitat edges on population dynamics with contemporary theoretical developments to outline the ways in which species interactions, and the dynamics of the communities in which they are embedded, can be changed by habitat edges. We find a striking convergence between empirical notions of a patch's core area and analytical results from partial differential equation models. A review of both empirical and theoretical studies suggests four general classes of mechanisms through which habitat edges can have similar impacts on dissimilar types of species interactions. Specifically, we focus on edges' roles as dispersal barriers or filters, edges' influences on mortality, edges' involvement in spatial subsidies (in which dispersers' intrapatch impacts are maintained by their activities in other habitats), and edges' roles as generators of novel interactions. For each class of edge-mediated effects, we provide examples of how one can use spatial modeling to address the relevant questions on these topics, which together form a key link between community dynamics and landscape structure.

Omnivory-defined broadly as feeding on more than one trophic level-occupies a prominent position in discussions of food web architecture and dynamics, due in large part to an enduring conflict regarding omnivory's role in community dynamics. According to classical results from mathematical food web theory, omnivory destabilizes ecological communities, whereas more recent conceptual syntheses suggest that omnivory should be a strongly stabilizing factor in food webs. Working with an arthropod assemblage at Mount Saint Helens, I experimentally addressed this controversy using a two-way factorial design that crossed a manipulation of the degree of omnivory with another "disturbance" manipulation that targeted a specific component of the assemblage. In this statistical design, significant interaction effects (i.e., how the community impacts of the disturbance varied with the degree of omnivory) identified key stabilizing or destabilizing influences of omnivory. Overall, my experimental results indicated that increasing the degree of omnivory stabilized community dynamics, in keeping with recent conceptual syntheses.