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Stock composition of the historical New York Bight Atlantic sturgeon (Acipenser oxyrinchus oxyrinchus) intercept fishery revealed through microsatellite analysis of archived spines

A targeted commercial fishery for Atlantic Sturgeon Acipenser oxyrinchus oxyrinchus once operated in the New York Bight, where it was assumed that most harvested Atlantic Sturgeon were natal to the Hudson River population. However, more recent evidence suggests that the fishery may have been targeting a mixed-stock aggregation, in which case harvested Atlantic Sturgeon could have been comprised of individuals from multiple populations throughout the species’ range. Although there is now a moratorium on Atlantic Sturgeon harvest in the New York Bight, modern molecular approaches provide an opportunity to use archived tissues to perform a retrospective mixed-stock analysis on the fishery. Genomic DNA extracted from archived fin spines from 80 Atlantic Sturgeon collected nearly 30 years ago suggests that the fishery primarily harvested individuals from the Hudson River population. However, based on individual-based assignment tests, our results indicate that the fishery also harvested individuals from at least eight other populations located throughout the species’ range. This study highlights how archival hard parts that were previously used for age and growth analyses can be employed for retrospective genetic analyses. Further, because the New York Bight harbors relatively high concentrations of Atlantic Sturgeon, the study shows how localized management decisions can influence Atlantic Sturgeon conservation at rangewide scales. When integrated with more recent knowledge of species ecology, these analyses can be used to evaluate the efficacy of previous management strategies and understand the effects of historical processes on contemporary demography.

New York↗

Sample design effects in landscape genetics

An important research gap in landscape genetics is the impact of different field sampling designs on the ability to detect the effects of landscape pattern on gene flow. We evaluated how five different sampling regimes (random, linear, systematic, cluster, and single study site) affected the probability of correctly identifying the generating landscape process of population structure. Sampling regimes were chosen to represent a suite of designs common in field studies. We used genetic data generated from a spatially-explicit, individual-based program and simulated gene flow in a continuous population across a landscape with gradual spatial changes in resistance to movement. Additionally, we evaluated the sampling regimes using realistic and obtainable number of loci (10 and 20), number of alleles per locus (5 and 10), number of individuals sampled (10-300), and generational time after the landscape was introduced (20 and 400). For a simulated continuously distributed species, we found that random, linear, and systematic sampling regimes performed well with high sample sizes (>200), levels of polymorphism (10 alleles per locus), and number of molecular markers (20). The cluster and single study site sampling regimes were not able to correctly identify the generating process under any conditions and thus, are not advisable strategies for scenarios similar to our simulations. Our research emphasizes the importance of sampling data at ecologically appropriate spatial and temporal scales and suggests careful consideration for sampling near landscape components that are likely to most influence the genetic structure of the species. In addition, simulating sampling designs a priori could help guide filed data collection efforts.

Conservation Genetics↗

A standard protocol for describing individual-based and agent-based models

Simulation models that describe autonomous individual organisms (individual based models, IBM) or agents (agent-based models, ABM) have become a widely used tool, not only in ecology, but also in many other disciplines dealing with complex systems made up of autonomous entities. However, there is no standard protocol for describing such simulation models, which can make them difficult to understand and to duplicate. This paper presents a proposed standard protocol, ODD, for describing IBMs and ABMs, developed and tested by 28 modellers who cover a wide range of fields within ecology. This protocol consists of three blocks (Overview, Design concepts, and Details), which are subdivided into seven elements: Purpose, State variables and scales, Process overview and scheduling, Design concepts, Initialization, Input, and Submodels. We explain which aspects of a model should be described in each element, and we present an example to illustrate the protocol in use. In addition, 19 examples are available in an Online Appendix. We consider ODD as a first step for establishing a more detailed common format of the description of IBMs and ABMs. Once initiated, the protocol will hopefully evolve as it becomes used by a sufficiently large proportion of modellers.

Ecological Modelling↗

A multi-model framework for simulating wildlife population response to land-use and climate change

Reliable assessments of how human activities will affect wildlife populations are essential for making scientifically defensible resource management decisions. A principle challenge of predicting effects of proposed management, development, or conservation actions is the need to incorporate multiple biotic and abiotic factors, including land-use and climate change, that interact to affect wildlife habitat and populations through time. Here we demonstrate how models of land-use, climate change, and other dynamic factors can be integrated into a coherent framework for predicting wildlife population trends. Our framework starts with land-use and climate change models developed for a region of interest. Vegetation changes through time under alternative future scenarios are predicted using an individual-based plant community model. These predictions are combined with spatially explicit animal habitat models to map changes in the distribution and quality of wildlife habitat expected under the various scenarios. Animal population responses to habitat changes and other factors are then projected using a flexible, individual-based animal population model. As an example application, we simulated animal population trends under three future land-use scenarios and four climate change scenarios in the Cascade Range of western Oregon. We chose two birds with contrasting habitat preferences for our simulations: winter wrens (Troglodytes troglodytes), which are most abundant in mature conifer forests, and song sparrows (Melospiza melodia), which prefer more open, shrubby habitats. We used climate and land-use predictions from previously published studies, as well as previously published predictions of vegetation responses using FORCLIM, an individual-based forest dynamics simulator. Vegetation predictions were integrated with other factors in PATCH, a spatially explicit, individual-based animal population simulator. Through incorporating effects of landscape history and limited dispersal, our framework predicted population changes that typically exceeded those expected based on changes in mean habitat suitability alone. Although land-use had greater impacts on habitat quality than did climate change in our simulations, we found that small changes in vital rates resulting from climate change or other stressors can have large consequences for population trajectories. The ability to integrate bottom-up demographic processes like these with top-down constraints imposed by climate and land-use in a dynamic modeling environment is a key advantage of our approach. The resulting framework should allow researchers to synthesize existing empirical evidence, and to explore complex interactions that are difficult or impossible to capture through piecemeal modeling approaches. ?? 2008 Elsevier B.V.

Ecological Modelling↗

Modeling the impacts of hunting on the population dynamics of red howler monkeys ( Alouatta seniculus )

Overexploitation of wildlife populations occurs across the humid tropics and is a significant threat to the long-term survival of large-bodied primates. To investigate the impacts of hunting on primates and ways to mitigate them, we developed a spatially explicit, individual-based model for a landscape that included hunted and un-hunted areas. We used the large-bodied neotropical red howler monkey ( Alouatta seniculus ) as our case study species because its life history characteristics make it vulnerable to hunting. We modeled the influence of different rates of harvest and proportions of landscape dedicated to un-hunted reserves on population persistence, population size, social dynamics, and hunting yields of red howler monkeys. In most scenarios, the un-hunted populations maintained a constant density regardless of hunting pressure elsewhere, and allowed the overall population to persist. Therefore, the overall population was quite resilient to extinction; only in scenarios without any un-hunted areas did the population go extinct. However, the total and hunted populations did experience large declines over 100 years under moderate and high hunting pressure. In addition, when reserve area decreased, population losses and losses per unit area increased disproportionately. Furthermore, hunting disrupted the social structure of troops. The number of male turnovers and infanticides increased in hunted populations, while birth rates decreased and exacerbated population losses due to hunting. Finally, our results indicated that when more than 55% of the landscape was harvested at high (30%) rates, hunting yields, as measured by kilograms of biomass, were less than those obtained from moderate harvest rates. Additionally, hunting yields, expressed as the number of individuals hunted/year/km 2 , increased in proximity to un-hunted areas, and suggested that dispersal from un-hunted areas may have contributed to hunting sustainability. These results indicate that un-hunted areas serve to enhance hunting yields, population size, and population persistence in hunted landscapes. Therefore, spatial regulation of hunting via a reserve system may be an effective management strategy for sustainable hunting, and we recommend it because it may also be more feasible to implement than harvest quotas or restrictions on season length.

Guárico↗

The ODD protocol: A review and first update

The ‘ODD’ (Overview, Design concepts, and Details) protocol was published in 2006 to standardize the published descriptions of individual-based and agent-based models (ABMs). The primary objectives of ODD are to make model descriptions more understandable and complete, thereby making ABMs less subject to criticism for being irreproducible. We have systematically evaluated existing uses of the ODD protocol and identified, as expected, parts of ODD needing improvement and clarification. Accordingly, we revise the definition of ODD to clarify aspects of the original version and thereby facilitate future standardization of ABM descriptions. We discuss frequently raised critiques in ODD but also two emerging, and unanticipated, benefits: ODD improves the rigorous formulation of models and helps make the theoretical foundations of large models more visible. Although the protocol was designed for ABMs, it can help with documenting any large, complex model, alleviating some general objections against such models.

Ecological Modelling↗

Using dynamic population simulations to extend resource selection analyses and prioritize habitats for conservation

Prioritizing habitats for conservation is a challenging task, particularly for species with fluctuating populations and seasonally dynamic habitat needs. Although the use of resource selection models to identify and prioritize habitat for conservation is increasingly common, their ability to characterize important long-term habitats for dynamic populations are variable. To examine how habitats might be prioritized differently if resource selection was directly and dynamically linked with population fluctuations and movement limitations among seasonal habitats, we constructed a spatially explicit individual-based model for a dramatically fluctuating population requiring temporally varying resources. Using greater sage-grouse ( Centrocercus urophasianus) in Wyoming as a case study, we used resource selection function maps to guide seasonal movement and habitat selection, but emergent population dynamics and simulated movement limitations modified long-term habitat occupancy. We compared priority habitats in RSF maps to long-term simulated habitat use. We examined the circumstances under which the explicit consideration of movement limitations, in combination with population fluctuations and trends, are likely to alter predictions of important habitats. In doing so, we assessed the future occupancy of protected areas under alternative population and habitat conditions. Habitat prioritizations based on resource selection models alone predicted high use in isolated parcels of habitat and in areas with low connectivity among seasonal habitats. In contrast, results based on more biologically-informed simulations emphasized central and connected areas near high-density populations, sometimes predicted to be low selection value. Dynamic models of habitat use can provide additional biological realism that can extend, and in some cases, contradict habitat use predictions generated from short-term or static resource selection analyses. The explicit inclusion of population dynamics and movement propensities via spatial simulation modeling frameworks may provide an informative means of predicting long-term habitat use, particularly for fluctuating populations with complex seasonal habitat needs. Importantly, our results indicate the possible need to consider habitat selection models as a starting point rather than the common end point for refining and prioritizing habitats for protection for cyclic and highly variable populations.

Wyoming↗

The confluences of ideas leading to, and the flow of ideas emerging from, individual-based modeling of riverine fishes

In this review article, we trace the history of events leading to the development of individual-based models (IBMs) to represent aquatic organisms in rivers and streams. As a metaphor, we present this history as a series of confluences between individual scientists (tributaries) sharing ideas. We describe contributions of these models to science and management. One iconic feature of river IBMs is the linkage between flow and the physical habitat experienced by individual animals, and the first model that focused on this linkage is briefly described. We continue by reviewing the contributions of riverine IBMs to eight broad areas of scientific inquiry. The first four areas include research to understand 1) the effects of flow regimes on fish populations, 2) species interactions (e.g., size-mediated competition and predation), 3) fish movement and habitat selection, and 4) contaminant and water quality impacts on populations. Next, we review research using IBMs 5) to guide conservation biology of imperiled taxa through population viability analysis, including research 6) to understand river fragmentation by dams and reconnection, 7) to understand genetic outcomes for riverine metapopulations, and 8) to anticipate the future effects of temperature and climate change. This rich body of literature has contributed to both theoretical insights (e.g., about animal behavior and life history) and applied insights (e.g., population-level effects of flow regimes, temperature, and the effects of hydropower and other industries that share rivers with aquatic biota). We finish by exploring promising branches that lie ahead in the braided, downstream channel that represents future river modeling research.

Ecological Modelling↗

The economics of territory selection

Territorial behavior is a fundamental and conspicuous behavior within numerous species, but the mechanisms driving territory selection remain uncertain. Theory and empirical precedent indicate that many animals select territories economically to satisfy resource requirements for survival and reproduction, based on benefits of food resources and costs of competition and travel. Costs of competition may vary by competitive ability, and costs of predation risk may also drive territory selection. Habitat structure, resource requirements, conspecific density, and predator distribution and abundance are likely to further influence territorial behavior. We developed a mechanistic, spatially-explicit, individual-based model to better understand how and why animals select particular territories. The model was based on optimal selection of individual patches for inclusion in a territory according to their net value, i.e., benefits (food resources) minus costs (travel, competition, predation risk). Simulations produced predictions for what may be observed empirically if such optimization drives placement and characteristics of territories. Simulations consisted of sequential, iterative selection of territories by simulated animals that interacted to defend and maintain territories. Results explain why certain patterns in space use are commonly observed, and when and why these patterns will differ from the norm. For example, more clumped or abundant food resources are predicted to result, on average, in smaller territories with more overlap. Strongly different resource requirements for individuals or groups in a population will directly affect space use and are predicted to cause different responses under identical conditions. Territories are predicted to decrease in size with increasing population density, which can enable a population’s density of territories to change at faster rates than their spatial distribution. Due to competition, less competitive territory-holders are generally predicted to have larger territories in order to accumulate sufficient resources, which could produce an ideal despotic distribution of territories. Interestingly, territory size and overlap are predicted to show a parabolic response to increases in predator densities, and territories are predicted to be larger where predators are more clumped in distribution. Our model’s predictions are consistent with many empirical observations, providing support for optimal patch selection as a mechanism for the economical territories of animals commonly observed in nature.

Ecological Modelling↗

Modeling structural mechanics of oyster reef self-organization including environmental constraints and community interactions

Self-organization is a process of establishing and reinforcing local structures through feedbacks between internal population dynamics and external factors. In reef-building systems, substrate is collectively engineered by individuals that also occupy it and compete for space. Reefs are constrained spatially by the physical environment, and by mortality, which reduces production but exposes substrate for recruits. Reef self-organization therefore depends on efficient balancing of production and occupancy of substrate. To examine this, we develop a three-dimensional individual-based model (IBM) of oyster reef mechanics. Shell substrate is grown by individuals as valves, accumulates at the reef level, and degrades following mortality. Single restoration events and subsequent dynamics are simulated for a case study in South Carolina (USA). Variability in model processes is included on recruitment, spatial environmental constraints, and predation, over multiple independent runs and five predator community scenarios. The main goal for this study is to summarize trends in dynamics that are robust across this uncertainty, and from these generate new hypotheses and predictions for future studies. Simulation results demonstrate three phases following restoration: initial transient dynamics with considerable shell loss, followed by growth and saturation of the live population, and then saturation of settlement habitat several years later. Over half of simulations recoup initial shell losses as populations grow, while others continue in decline. The balance between population density, substrate supporting the reef, and exposed surfaces for settlement is mediated by overall population size and size structure, presence of predators, and relative amounts of live individuals and intact dead shells. The efficiency of settlement substrate production improves through time as population size structure becomes more complex, and the population of dead valves accumulates.

Ecological Modelling↗

Visualizing wading bird optimal foraging decisions with aggregation behaviors using individual-based modeling

Foragers on patchy landscapes must efficiently balance time between searching for and consuming resources to meet their daily energetic requirements. Spatial aggregation foraging behaviors may improve foraging efficiency by sharing information on locations of resource hotspots. Wading birds are an example of patch foragers that form colonial aggregations during the breeding season to obtain sufficient prey energy to sustain themselves and their offspring each day. Here, we describe a spatially-explicit simulation model of wading bird optimal foraging that represents information sharing through visual cues. The overall purpose of the model is to describe how wading bird daily foraging and reproductive success may change with alternative water control management practices that determine spatial availability of prey for wading birds on the landscape, throughout their breeding seasons. Wading birds are simulated as individuals that operate independently, sampling and selecting among patches based on a prey density tolerance threshold, but also use information from other birds to inform their selection decisions. Foraging success is evaluated against the fundamental objectives of (a) fulfilling daily energetic demands and (b) minimizing predation exposure, by tracking individual daily energetic intake and time spent foraging. In this way, the model approximates population level dynamics of wading bird aggregations that emerge through collective decision making of birds simulated at the lower individual level. Key results of this study suggest that aggregation behaviors may improve population-level foraging success rates, and the optimal settling threshold may modulate when resources become more scarce or difficult to find. Thus, the model addresses ecological theory on the advantages of foraging in groups versus independently. This technique is appropriate for evaluating wading bird populations that forage on patchy landscapes, such as seasonally-pulsed wetlands, wherever sufficient information is available to describe (1) foraging behavior (e.g., feeding rate, flight speeds, patch selection decisions), (2) key landscape characteristics, (3) spatial distributions of prey densities among foraging patches, and (4) changes in prey densities through time. The model was designed to predict qualitative, testable spatial patterns of wading bird foraging movements which can be compared with empirical observations and empirically-derived habitat suitability models. These techniques can also be applied to other bird species, such as shorebirds, or more generally to any species that transits between discrete foraging patches.

Ecological Modelling↗

Foraging of wading birds on a patchy landscape: Simulating effects of social information, interference competition, and patch selection on prey intake and individual distribution

Foragers on patchy landscapes must acquire sufficient resources despite uncertainty in the location and amount of the resources. Optimal Foraging Theory posits that foragers deal with this uncertainty by using strategies that optimize resource intake within foraging periods. For species such as wading birds, this optimization is closely linked to their survival and reproductive success. Understanding the influence of patch selection on individual resource intake and foraging distribution is therefore crucial. In this study, we simulated how resource distribution, interference competition, and social cues—such as aggregation behaviors—influence resource intake and foraging spatial distribution. We employed an individual-based model simulating wading bird foraging behaviors, with 900 individuals simultaneously foraging across a landscape with unknown resource distribution. Birds employed one of three patch-finding strategies: random, cue-searching, or hybrid, which uses both searching strategies. Each bird decided whether to remain in a patch based on a prey density threshold. We compared the daily resource intake and foraging distribution of birds across different modeled patch-finding strategies, resource distribution patterns, and the presence or absence of interference competition. Wading birds exhibiting aggregation behavior displayed increased intake rates when resources were concentrated and interference minimal. Aggregation behavior led to a closer match with the ideal free distribution when the prey density threshold was optimal. These findings provide theoretical support that aggregation behavior is effective in scenarios where resources are concentrated in a few patches, social cues are used by relatively few individuals, and interference competition is limited.

Ecological Modelling↗

The occupancy-abundance relationship and sampling designs using occupancy to monitor populations of Asian bears

Designing a population monitoring program for Asian bears presents challenges associated with their low densities and detectability, generally large home ranges, and logistical or resource constraints. The use of an occupancy-based method to monitor bear populations can be appropriate under certain conditions given the mechanistic relationship between occupancy and abundance. The form of the occupancy–abundance relationship is dependent on species-specific characteristics such as home range size and population density, as well as study area size. To assess the statistical power of tests to detect population change of Asian bears, we conducted a study using a range of scenarios by simulating spatially explicit individual-based capture-recapture data from a demographically open model. Simulations assessed the power to detect changes in population density via changes in site-level occupancy or abundance through time, estimated using a standard occupancy model or a Royle-Nichols model, both with point detectors (representing camera traps). We used IUCN Red List criteria as a guide in selection of two population decline scenarios (20% and 50%), but we chose a shorter time horizon (10 years = 1 bear generation), meaning that declines were steeper than used for IUCN criteria (3 generations). Our simulations detected population declines of 50% with high power (>0.80) and low false positive rates (FPR: incorrectly detecting a decline) (<0.10) when detectors were spaced at > 0.67 times the home range diameter (home-range spacing ratio: HRSR, a measure of spatial correlation), such that bears would tend to overlap no more than two detectors. There was high (0.85) correlation between realized occupancy and N in these scenarios. The FPR increased as the HRSR decreased because of spatial correlation in the occupancy process induced when individual home ranges overlap multiple detectors. The mean statistical power to detect more gradual population declines (20% in 10 years) with HRSR > 0.67 was low for occupancy models 0.22 (maximum power 0.67) and Royle-Nichols models (0.24; maximum power 0.67), suggesting that declines of this magnitude may not be described reliably with 10 years of monitoring. Our results demonstrated that under many realistic scenarios that we explored, false positive rates were unacceptably high. We highlight that when designing occupancy studies, the spacing between point detectors be at least 0.67 times the diameter of the home range size of the larger sex (e.g., males) when the assumptions of the spatial capture-recapture model used for simulation are met.

Global Ecology and Conservation↗

Basis function models for animal movement

Advances in satellite-based data collection techniques have served as a catalyst for new statistical methodology to analyze these data. In wildlife ecological studies, satellite-based data and methodology have provided a wealth of information about animal space use and the investigation of individual-based animal–environment relationships. With the technology for data collection improving dramatically over time, we are left with massive archives of historical animal telemetry data of varying quality. While many contemporary statistical approaches for inferring movement behavior are specified in discrete time, we develop a flexible continuous-time stochastic integral equation framework that is amenable to reduced-rank second-order covariance parameterizations. We demonstrate how the associated first-order basis functions can be constructed to mimic behavioral characteristics in realistic trajectory processes using telemetry data from mule deer and mountain lion individuals in western North America. Our approach is parallelizable and provides inference for heterogenous trajectories using nonstationary spatial modeling techniques that are feasible for large telemetry datasets. Supplementary materials for this article are available online.

Journal of the American Statistical Association↗

Thinking beyond the closure assumption: Designing surveys for estimating biological truth with occupancy models

Occupancy models estimate distributions of imperfectly detected species, but violations of the closure assumption can bias results. However, researchers working with mobile animals may find it impossible to eliminate such violations. Here, we tested the hypothesis that occupancy models fit to realistic sampling data can generate unbiased occupancy estimates for an itinerant Wood Thrush ( Hylocichla mustelina ) population. In 2013 and 2014, we tracked movements of 41 breeding Wood Thrush males. We modelled territory shift probabilities using logistic exposure models and within-territory movements using continuous-time stochastic process models. We then constructed an individual-based model, simulated (1000 iterations) spatiotemporal locations for individuals and simulated sampling these populations using 162 different point count protocols with variable spatial (sampling radius and point placement method), and temporal (survey length, between-survey intervals and number of surveys) characteristics. We compared occupancy estimates with true values of instantaneous, daily and seasonal occupancy from the simulations. We parameterized continuous time stochastic process models based on movements within 34 unique territories and estimated a daily territory shift probability of 0.0099 (95% CI: 0.0060, 0.0152). Simulated data indicated that estimates of occupancy ranged from 0.18 (0.06, 1.00) to 0.80 (0.71, 0.89) depending on protocol characteristics. Occupancy estimates increased with increasing survey radius, survey length and between-survey interval. Protocols using shorter surveys and between-survey intervals were good estimators for instantaneous occupancy (low bias and mean-squared error) but poor estimators for daily and seasonal occupancy; longer surveys and intervals generated unbiased estimators of daily occupancy but underestimated seasonal occupancy. Logistic regression models that ignored imperfect detection outperformed occupancy models for estimating instantaneous occupancy but not daily or seasonal occupancy. For mobile animals, occupancy of sampling sites changes in space and time. Consequently, the spatial and temporal aspects of a sampling protocol have strong, but predictable, effects on occupancy model parameter estimates. Our results demonstrate that how these factors interact is critical for designing surveys that produce occupancy estimates representative of the biological process of interest to a researcher.

Methods in Ecology and Evolution↗

Growth potential and habitat requirements of endangered age-0 pallid sturgeon (Scaphirhynchus albus) in the Missouri River, USA, determined using a individual-based model framework

An individual-based model framework was used to evaluate growth potential of the federally endangered pallid sturgeon ( Scaphirhynchus albus ) in the Missouri River. The model, developed for age-0 sturgeon, combines information on functional feeding response, bioenergetics and swimming ability to regulate consumption and growth within a virtual foraging arena. Empirical data on water temperature, water velocity and prey density were obtained from three sites in the Missouri River and used as inputs in the model to evaluate hypotheses concerning factors affecting pallid sturgeon growth. The model was also used to evaluate the impacts of environmental heterogeneity and water velocity on individual growth variability, foraging success and dispersal ability. Growth was simulated for a period of 100 days using 100 individuals (first feeding; 19 mm and 0.035 g) per scenario. Higher growth was shown to occur at sites where high densities of Ephemeroptera and Chironomidae larvae occurred throughout the growing season. Highly heterogeneous habitats (i.e., wide range of environmental conditions) and moderate water velocities (0.3 m/s) were also found to positively affect growth rates. The model developed here provides an important management and conservation tool for evaluating growth hypotheses and(or) identifying habitats in the Missouri River that are favourable to age-0 pallid sturgeon growth.

Ecology of Freshwater Fish↗

Effects of sample size, number of markers, and allelic richness on the detection of spatial genetic pattern

The influence of study design on the ability to detect the effects of landscape pattern on gene flow is one of the most pressing methodological gaps in landscape genetic research. To investigate the effect of study design on landscape genetics inference, we used a spatially-explicit, individual-based program to simulate gene flow in a spatially continuous population inhabiting a landscape with gradual spatial changes in resistance to movement. We simulated a wide range of combinations of number of loci, number of alleles per locus and number of individuals sampled from the population. We assessed how these three aspects of study design influenced the statistical power to successfully identify the generating process among competing hypotheses of isolation-by-distance, isolation-by-barrier, and isolation-by-landscape resistance using a causal modelling approach with partial Mantel tests. We modelled the statistical power to identify the generating process as a response surface for equilibrium and non-equilibrium conditions after introduction of isolation-by-landscape resistance. All three variables (loci, alleles and sampled individuals) affect the power of causal modelling, but to different degrees. Stronger partial Mantel r correlations between landscape distances and genetic distances were found when more loci were used and when loci were more variable, which makes comparisons of effect size between studies difficult. Number of individuals did not affect the accuracy through mean equilibrium partial Mantel r , but larger samples decreased the uncertainty (increasing the precision) of equilibrium partial Mantel r estimates. We conclude that amplifying more (and more variable) loci is likely to increase the power of landscape genetic inferences more than increasing number of individuals.

Molecular Ecology Resources↗

Learning to be different: Acquired skills, social learning, frequency dependence, and environmental variation can cause behaviourally mediated foraging specializations

Question: How does the ability to improve foraging skills by learning, and to transfer that learned knowledge, affect the development of intra-population foraging specializations? Features of the model: We use both a state-dependent life-history model implemented by stochastic dynamic programming (SDPM) and an individual-based model (IBM) to capture the dynamic nature of behavioural preferences in feeding. Variables in the SDPM include energy reserves, skill levels, energy and handling time per single prey item, metabolic rate, the rates at which skills are learned and forgotten, the effect of skills on handling time, and the relationship between energy reserves and fitness. Additional variables in the IBM include the probability of successful weaning, the logistic dynamics of the prey species with stochastic recruitment, the intensity of top-down control of prey by predators, the mean and variance in skill levels of new recruits, and the extent to which learned Information can be transmitted via matrilineal social learning. Key range of variables: We explore the effects of approaching the time horizon in the SDPM, changing the extent to which skills can improve with experience, increasing the rates of learning or forgetting of skills, changing whether the learning curve is constant, accelerating (T-shaped) or decelerating ('r'-shaped), changing both mean and maximum possible energy reserves, changing metabolic costs of foraging, and changing the rate of encounter with prey. Conclusions: The model results show that the following factors increase the degree of prey specialization observed in a predator population: (1) Experience handling a prey type can substantially improve foraging skills for that prey. (2) There is limited ability to retain complex learned skills for multiple prey types. (3) The learning curve for acquiring new foraging skills is accelerating, or J-shaped. (4) The metabolic costs of foraging are high relative to available energy reserves. (5) Offspring can learn foraging skills from their mothers (matrilineal social learning). (6) Food abundance is limited, such that average individual energy reserves are low Additionally, the following factors increase the likelihood of alternative specializations co-occurring in a predator population: (1) The predator exerts effective top-down control of prey abundance, resulting in frequency-dependent dynamics. (2) There is stochastic Variation in prey population dynamics, but this Variation is neither too extreme in magnitude nor too 'slow' with respect to the time required for an individual forager to learn new foraging skills. For a given predator population, we deduce that the degree of specialization will be highest for those prey types requiring complex capture or handling skills, while prey species that are both profitable and easy to capture and handle will be included in the diet of all individuals. Frequency-dependent benefits of selecting alternative prey types, combined with the ability of foragers to improve their foraging skills by learning, and transmit learned skills to offspring, can result in behaviourally mediated foraging specialization, and also lead to the co-existence of alternative specializations. The extent of such specialization is predicted to be a variable trait, increasing in locations or years when intra-specific competition is high relative to inter-specific competition. ?? 2009 M. Tim Tinker.

Evolutionary Ecology Research↗