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Finding the Missing Link between Landscape Structure and Population Dynamics: A Spatially Explicit Perspective.

We construct and explore a general modeling framework that allows for a systematic investigation of the impact of changes in landscape structure on population dynamics. The essential parts of the framework are a landscape generator with independent control over landscape composition and physiognomy, an individual-based spatially explicit population model that simulates population dynamics within heterogeneous landscapes, and scale-dependent landscape indices that depict the essential aspects of landscape that interact with dispersal and demographic processes. Landscape maps are represented by a grid of [Formula: see text] cells and consist of good-quality, poor-quality, or uninhabitable matrix habitat cells. The population model was shaped in accordance to the biology of European brown bears (Ursus arctos), and demographic parameters were adjusted to yield a source-sink configuration. Results obtained with the spatially explicit model do not confirm results of earlier nonspatial source-sink models where addition of sink habitat resulted in a decrease of total population size because of dilution of high-quality habitat. Our landscape indices, which describe scale-dependent correlation between and within habitat types, were able to explain variations in variables of population dynamics (mean number of females with sink home ranges, mean number of females with source home ranges, and mean dispersal distance) caused by different landscape structure. When landscape structure changed, changes in these variables generally followed the corresponding change of an appropriate landscape index in a linear way. Our general approach incorporates source-sink dynamics as well as metapopulation dynamics, and the population model can easily be modified for other species groups.

habitat connectivity↗

Spatially explicit models for the population dynamics of a species colonizing an island.

We construct reaction-diffusion models for the population dynamics of a species colonizing an island from a source population on a continent. We view the source population as inducing a density or flux of immigrants onto the island and interpret colonization as succeeding if the population on the island is predicted to persist even when immigration from the continent is stopped. To capture the observation that a sufficiently large population or density must be attained for colonization to succeed, we assume Allee (i.e., bistable) dynamics rather than logistic dynamics for the colonizing population. We consider the cases of colonization in both the absence and presence of a competitor. We use reaction-diffusion theory, especially comparison methods and sub- and supersolutions, to determine how parameters such as the distance from the continent to the island and the dispersal, birth and mortality rates, carrying capacity, and minimum viable population density of the colonizing species affect the outcome of the attempted colonization. In the case of colonization in the presence of a competitor we consider a number of scenarios involving different types and strengths of competition. Our analysis permits us to draw conclusions about the characteristics of a species that make it a good colonizer.

Animals↗

Synchronism in a metapopulation model.

We consider a spatially explicit meta-population model with interaction among the two nearest neighbors to relate, with a simple mathematical expression, chaos in the local, uncoupled, populations, the degree of interaction among patches, size of the meta-population, and the stability of the synchronized attractor. Since synchronism is strongly correlated with extinction, our results can provide useful information on factors leading to population extinction.

Animals↗

Spatially explicit model of transposon-based genetic drive mechanisms for displacing fluctuating populations of anopheline vector mosquitoes.

To evaluate the prospect of transposon-based genetic drive mechanisms for replacing African vectors of malaria with nonvector anopheline mosquitoes, we developed a spatially explicit simulation model that determined the likelihood that released transgenic mosquitoes may proceed to fixation or extinction under diverse conditions. We compared the effect on fixation of long breeding seasons with relatively subtle population fluctuations to short breeding seasons with severe bottlenecks. Assuming 100% transposition efficiency among heterozygotes with fitness varying between 50 and 100% of that of wild-type mosquitoes, we simulated releases of 1, 10, 50, 90, and 99% of transposon-bearers in relation to wild mosquitoes as well as 1 and 10% releases that were repeated annually. We also evaluated diverse patterns of release including linear, marginal, focused, and scattered distribution. Random dispersal provided the most rapid fixation of transposons within populations. More massive releases allowed longer persistence of transposon-bearers but did not promote fixation, especially when breeding seasons were long. Relative fitness of transposon-bearers, however, proved more powerful than pattern or number of releases in determining whether a construct will become fixed or extinct. Even when fitness approaches that of the wild-type, fixation of a construct may require 150 generations or more.

Animals↗

Source-sink dynamics between transgenic and non-transgenic habitats and their role in the evolution of resistance.

The interaction of population dynamics and movement among two habitat types (toxic transgenic fields and nontoxic refuge fields) on the evolution of insecticide resistance was examined in two different simulation models. The two models were developed to test the hypothesis that increasing habitat grain from fine-grained to coarse-grained, and the resultant increase in nonrandom mating, would increase the rate of local adaptation, here the evolution of resistance. The first model, a complex, stochastic spatially explicit model, altered habitat grain by varying adult dispersal rates between habitat patches. In contrast to the expectation that increasing patch isolation and increasing the coarseness of the habitats would increase the rate of resistance evolution, intermediate levels of dispersal actually delayed resistance by as much as fivefold over the range of dispersal levels observed. Source-sink dynamics related to ovipositional patterns and the related population dynamics appear to explain the results. A simple deterministic model was developed to abstract out the separate impacts of mating and ovipositional behaviors. This model showed qualitatively the same results, although under similar assumptions it predicted much longer delays in resistance evolution. In this model, nonrandom mating alone always increased the rate at which insects adapted to transgenic crops, but nonrandom mating in combination with nonrandom oviposition could significantly delay resistance evolution. Differences between the two models may be due to the population regulation incorporated in the spatially explicit model. The models clearly suggest that resistance management programs using untreated refuges should not over-emphasize random mating at the cost of making the habitat too fine-grained.

Animals↗

Spatial distributions of genotypes under isolation by distance.

The spatial distributions of single-locus diploid genotypes, produced within populations of sexually reproducing individuals under isolation by distance, are measured and characterized in detail by quantifying the join-counts for simulated model populations. The models more realistically reflect spatially explicit distributions of genotypes in populations, because unlike the classical theory, they include the stochasticity inherent in the process of matings between genotypes. This stochasticity causes the formation of large areas or patches containing mostly one homozygous genotype, which is not predicted by the classical theory. A number of previously uncharacterized features of the spatial structures produced under isolation by distance are revealed. Spatial autocorrelation measures based on counts of pairs of homozygotes are highly consistent quantifications of the concentrations of homozygotes in patchy genotypic distributions for a given level of dispersal. Most strikingly, the degree of intermixing of homozygotes with heterozygotes over small spatial scales is much higher than previously thought, unless dispersal is extremely limited (e.g., Wright's neighborhood size less than approximately 5.0). Among implications for studies of spatial structure of real populations, one is that the results provide a basis for using join-counts as estimators of gene dispersal based solely on genetic data.

Diploidy↗

Host spatial heterogeneity and extinction of an SIS epidemic.

Spatially explicit epidemic models explore population-level consequences of interactions between neighboring infectious and susceptible individuals. Most such models equate local and global host density, so that each individual interacts with the same number of neighbors. But many natural populations exhibit heterogeneity in local abundance. Therefore, we let host dispersion vary from uniform to clumped in a spatial epidemic with recovery. We analytically approximated the epidemic with a branching process to show how the probability of pathogen extinction could depend on the degree of host clumping. We then simulated the detailed model across a range of parameter combinations. Both approaches to the problem indicate that host spatial aggregation strongly increases the chance of pathogen extinction.

Animals↗

Pollen movement in declining populations of California Valley oak, Quercus lobata: where have all the fathers gone?

The fragmented populations and reduced population densities that result from human disturbance are issues of growing importance in evolutionary and conservation biology. A key issue is whether remnant individuals become reproductively isolated. California Valley oak (Quercus lobata) is a widely distributed, endemic species in California, increasingly jeopardized by anthropogenic changes in biota and land use. We studied pollen movement in a savannah population of Valley oak at Sedgwick Reserve, Santa Barbara County, to estimate effective number of pollen donors (Nep) and average distance of effective pollen movement (delta). Using twogener, our recently developed hybrid model of paternity and genetic structure treatments that analyses maternal and progeny multilocus genotypes, we found that current Nep = 3.68 individuals. Based on an average adult density of d= 1.19 stems/ha, we assumed a bivariate normal distribution to model current average pollen dispersal distance (delta) and estimated delta= 64.8 m. We then deployed our parameter estimates in spatially explicit models of the Sedgwick population to evaluate the extent to which Nep may have changed, as a consequence of progressive stand thinning between 1944 and 1999. Assuming that pollen dispersal distance has not changed, we estimate Nep was 4.57 individuals in 1944, when stand density was 1.48. Both estimates indicate fewer effective fathers than one might expect for wind-pollinated species and fewer than observed elsewhere. The results presented here provide a basis for further refinements on modelling pollen movement. If the trends continue, then ongoing demographic attrition could further reduce neighbourhood size in Valley oak resulting in increased risk of reproductive failure and genetic isolation.

California↗

Habitat structure and population persistence in an experimental community.

Understanding spatial population dynamics is fundamental for many questions in ecology and conservation. Many theoretical mechanisms have been proposed whereby spatial structure can promote population persistence, in particular for exploiter-victim systems (host-parasite/pathogen, predator-prey) whose interactions are inherently oscillatory and therefore prone to extinction of local populations. Experiments have confirmed that spatial structure can extend persistence, but it has rarely been possible to identify the specific mechanisms involved. Here we use a model-based approach to identify the effects of spatial population processes in experimental systems of bean plants (Phaseolus lunatus), herbivorous mites (Tetranychus urticae) and predatory mites (Phytoseiulus persimilis). On isolated plants, and in a spatially undivided experimental system of 90 plants, prey and predator populations collapsed; however, introducing habitat structure allowed long-term persistence. Using mechanistic models, we determine that spatial population structure did not contribute to persistence, and spatially explicit models are not needed. Rather, habitat structure reduced the success of predators at locating prey outbreaks, allowing between-plant asynchrony of local population cycles due to random colonization events.

Animals↗

Space and the persistence of male-killing endosymbionts in insect populations.

Male-killing bacteria are bacteria that are transmitted vertically through the females of their insect hosts. They can distort the sex ratio of their hosts by killing infected male offspring. In nature, male-killing endosymbionts (male killers) often have a 100% efficient vertical transmission, and multiple male-killing bacteria infecting a single population are observed. We use different model formalisms to study these observations. In mean-field models a male killer with perfect transmission drives the host population to extinction, and coexistence between multiple male killers within one population is impossible; however, in spatially explicit models, both phenomena are readily observed. We show how the spatial pattern formation underlies these results. In the case of high transmission efficiencies, waves with a high density of male killers alternate with waves of mainly wild-type hosts. The male killers cause local extinction, but this creates an opportunity for uninfected hosts to re-invade these areas. Spatial pattern formation also creates an opportunity for two male killers to coexist within one population: different strains create spatial regions that are qualitatively different; these areas then serve as different niches, making coexistence possible.

Animals↗

Selection for intermediate mortality and reproduction rates in a spatially structured population.

How local interactions influence both population and evolutionary dynamics is currently a key topic in theoretical ecology. We use a 'well-mixed' analytical model and spatially explicit individual-based models to investigate a system where a population is subject to rare disturbance events. The disturbance can only propagate through regions of the population where the density of individuals is sufficiently high and individuals affected by the disturbance die shortly after. We find that populations where individuals are sessile often exhibit very different dynamic behaviour when compared to populations where individuals are mobile and spatially well mixed. When mutations are allowed which affect either offspring birth rates or mortality rates, the well-mixed populations always evolve to a state where a single disturbance event leads to extinction. Populations often persist substantially longer if individuals are sessile and they disperse their offspring locally. We also find that for sessile populations selection may favour short-lived individuals with limited offspring production. Population dynamics are found to be strongly influenced by the host characters that are evolving and the rate at which host variation is introduced into the system.

Animals↗

Risk based management of contaminated sediments: consideration of spatial and temporal patterns in exposure modeling.

This paper addresses interactions among foraging behavior, habitat preferences, site characteristics, and spatial distribution of contaminants in developing PCB exposure estimates for winter flounder at a hypothetical open water dredged material disposal site in the coastal waters of New York and New Jersey (NY-NJ). The implications of these interactions for human health risk estimates for local recreational anglers who fish for and eat flounder are described. The models implemented in this study include a spatial submodel to account for spatial and temporal characteristics of fish exposures and a probabilistic adaptation of the Gobas bioaccumulation model that accounts for temporal variation in concentrations of hydrophobic contaminants in sediment and water. We estimated the geographic distribution of a winter flounder subpopulation offshore of NY-NJ based on species biology and its vulnerability to local recreational fishing, the foraging area of individual fish, and their migration patterns. We incorporated these parameters and an estimate of differential attraction to a management site into a spatially explicit model to assess the range of exposures within the population. The output of this modeling effort, flounder PCB tissue concentrations, provided exposure point concentrations for an estimate of human health risk through ingestion of locally caught flounder. The risks obtained for the spatially nonexplicit case are as much as 1 order of magnitude higher than those obtained with explicit consideration of spatial and temporal characteristics of winter flounder foraging and seasonal migration. This practice of "defaulting" to extremely conservative estimates for exposure parameters in the face of uncertainty ill serves the decision-making process for management of contaminated sediments in general and specifically for disposal of dredged materials. Consideration of realistic spatial and temporal scales in food chain models can help support sediment management decisions by providing a quantitative expression of the confidence in risk estimates.

Animals↗

A mechanistic model for partial preferences.

Classic prey optimal foraging model assumes that individual predators are globally omniscient; that is, they have exact knowledge of prey population densities in the environment. This study examines a spatially explicit individual-based model of a one-predator two-prey system where individual predators are assumed to be omniscient only locally, i.e., to know prey population densities only in the range of their perception. Due to local variations in prey numbers, the probability of acceptance of less profitable prey shifts from the zero-one rule to a gradually decreasing function, for which an explicit formula is derived, giving way to partial preferences. A corresponding predator functional response to more profitable prey is shown to have a sigmoid-like form.

Animals↗

Comparing approximations to spatio-temporal models for epidemics with local spread.

Analytical methods for predicting and exploring the dynamics of stochastic, spatially interacting populations have proven to have useful application in epidemiology and ecology. An important development has been the increasing interest in spatially explicit models, which require more advanced analytical techniques than the usual mean-field or mass-action approaches. The general principle is the derivation of differential equations describing the evolution of the expected population size and other statistics. As a result of spatial interactions no closed set of equations is obtained. Nevertheless, approximate solutions are possible using closure relations for truncation. Here we review and report recent progress on closure approximations applicable to lattice models with nearest-neighbour interactions, including cluster approximations and elaborations on the pair (or pairwise) approximation. This study is made in the context of an SIS model for plant-disease epidemics introduced in Filipe and Gibson (1998, Studying and approximating spatio-temporal models for epidemic spread and control, Phil. Trans. R. Soc. Lond. B 353, 2153-2162) of which the contact process [Harris, T. E. (1974), Contact interactions on a lattice, Ann. Prob. 2, 969] is a special case. The various methods of approximation are derived and explained and their predictions are compared and tested against simulation. The merits and limitations of the various approximations are discussed. A hybrid pairwise approximation is shown to provide the best predictions of transient and long-term, stationary behaviour over the whole parameter range of the model.

Cluster Analysis↗

Time and space issues in ecotoxicology: population models, landscape pattern analysis, and long-range environmental chemistry.

Advances in ecotoxicology addressing problems of time and spatial scales are presented and interpreted in the frame of concepts on population/community dynamics and landscape pattern analysis. Example deterministic/probabilistic modeling experiments are used to illustrate key concepts. Space and time scales analyzed are single and multigenerations of local populations, metapopulations, community, and ecosystem/landscape. Most population models used in recent ecotoxicology studies are deterministic and do not include a formal treatment of spatial processes, like migration or local random extinction. Some metapopulation models have been applied with success. Upscaling of ecotoxicological results at the community level is less developed, probably because of the inherent complexity of indirect and direct coactions among organisms. Community and ecosystem toxicity end points that could find a broad use in regulatory applications have not yet been identified. Some practical issues like the estimation of the potential for the natural attenuation of toxicity and the transport of contaminants along food chains must be addressed at these scales/levels of biological complexity. The estimation of ecotoxicological effects has been increasingly evolving to integrate modeling and monitoring contaminant transport and fate, landscape pattern analysis, and spatially explicit population dynamics (including direct and indirect communal interactions).

Agriculture↗

The use of spatial modeling in an aquatic food web to estimate exposure and risk.

This paper quantitatively evaluates interactions among foraging behavior, habitat preferences, site characteristics and the spatial distribution of contaminants in estimating PCB exposure concentrations for winter flounder at a hypothetical open water dredged material disposal site in the coastal waters of New York and New Jersey (NY-NJ). The models implemented in this study include a spatial submodel to account for spatial and temporal characteristics of fish exposure and a probabilistic adaptation of the Gobas bioaccumulation model to account for temporal variation in concentrations of polychlorinated biphenyls (PCBs) in sediment and water. We estimated the geographic distribution of an offshore winter flounder subpopulation based on species biology, including such variables as foraging area, habitat size, disposal site size and migration characteristics. We incorporated these variables together with an estimate of differential attraction to a management site within a spatially explicit model to assess the range of expected PCB exposures to a winter flounder population. The output of this modeling effort, flounder PCB tissue concentrations, provides exposure point concentrations for estimates of human health risk through ingestion of locally caught flounder. The risks obtained for the spatially non-explicit case are as much as one order of magnitude higher than those obtained after incorporating spatial and temporal characteristics of winter flounder foraging and seasonal migration. Incorporating spatial and temporal variables in food chain models can help support sediment management decisions by providing a quantitative expression of the confidence in risk estimates.

Animals↗

Habitat fragmentation resulting in overgrazing by herbivores.

Habitat fragmentation sometimes results in outbreaks of herbivorous insect and causes an enormous loss of primary production. It is hypothesized that the driving force behind such herbivore outbreaks is disruption of natural enemy attack that releases herbivores from top-down control. To test this hypothesis I studied how trophic community structure changes along a gradient of habitat fragmentation level using spatially implicit and explicit models of a tri-trophic (plant, herbivore and natural enemy) food chain. While in spatially implicit model number of trophic levels gradually decreases with increasing fragmentation, in spatially explicit model a relatively low level of habitat fragmentation leads to overgrazing by herbivore to result in extinction of the plant population followed by a total system collapse. This provides a theoretical support to the hypothesis that habitat fragmentation can lead to overgrazing by herbivores and suggests a central role of spatial structure in the influence of habitat fragmentation on trophic communities. Further, the spatially explicit model shows (i) that the total system collapse by the overgrazing can occur only if herbivore colonization rate is high; (ii) that with increasing natural enemy colonization rate, the fragmentation level that leads to the system collapse becomes higher, and the frequency of the collapse is lowered.

Animals↗

Spatial processes in the evolution of resistance in Helicoverpa zea (Lepidoptera: Noctuidae) to Bt transgenic corn and cotton in a mixed agroecosystem: a biology-rich stochastic simulation model.

A simulation model is developed to examine the role of spatial processes in the evolution of resistance in Helicoverpa zea populations to Bt corn and Bt cotton. The model is developed from the stochastic spatially explicit Heliothis virescens model described by Peck et al. (1999), to accommodate a spatial mix of two host crops (corn and cotton), and to reflect the agronomic practices, as well as the spatial and temporal population dynamics of H. zea, in eastern North Carolina. The model suggests that selection for resistance is more intense in Bt cotton fields than in Bt corn fields. It further suggests that local gene frequencies are highly dependent on local deployment levels of Bt crops despite the high mobility of the adult insects. Region-wide average gene frequencies depend on the region-wide level of Bt deployment, so incomplete technology adoption slows the rate of resistance evolution. However, on a local scale, H. zea populations in clusters of fields in which Bt use is high undergo far more rapid evolution than populations in neighboring clusters of fields in which Bt use is low. The model suggests that farm-level refuge requirements are important for managing the risk of resistance. The model can be used as an aid in designing plans for monitoring for resistance by suggesting the appropriate distribution of monitoring locations, which should focus on areas of highest Bt crop deployment. The findings need to be placed in the context of the input parameters, many of which are uncertain or highly variable in nature, and therefore, a thorough sensitivity analysis is warranted.

Agriculture↗