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An integral projection model for gizzard shad (Dorosoma cepedianum) utilizing density-dependent age-0 survival

Gizzard shad (Dorosoma cepedianum) is a common freshwater fish species found throughout the central and eastern portions of North America. Within these regions, gizzard shad play several critical roles in the freshwater community such as serving as prey for other fish species and translocating nutrients from substrates into the water column. Because of this, it is important to understand gizzard shad population dynamics. Here, we introduce an integral projection model (IPM) for gizzard shad that incorporates empirical information from sources including Long Term Resource Monitoring (LTRM) upper Mississippi River restoration data. IPMs are a generalization of stage-based, matrix population models that have been used to describe a wide range of organisms, and as such are a natural choice for gizzard shad because many aspects of their life cycle have been studied. We tested model outputs against empirical patterns reported for gizzard shad from a different location along the Illinois River (La Grange Reach). Results of our work indicate that our model could serve as an important tool for predicting patterns within gizzard shad populations.

Ecological Modelling

Comparing cropland net primary production estimates from inventory, a satellite-based model, and a process-based model in the Midwest of the United States

Accurately quantifying the spatial and temporal variability of net primary production (NPP) for croplands is essential to understand regional cropland carbon dynamics. We compared three NPP estimates for croplands in the Midwestern United States: inventory-based estimates using crop yield data from the U.S. Department of Agriculture (USDA) National Agricultural Statistics Service (NASS); estimates from the satellite-based Moderate Resolution Imaging Spectroradiometer (MODIS) NPP product; and estimates from the General Ensemble biogeochemical Modeling System (GEMS) process-based model. The three methods estimated mean NPP in the range of 469–687 g C m −2 yr −1 and total NPP in the range of 318–490 Tg C yr −1 for croplands in the Midwest in 2007 and 2008. The NPP estimates from crop yield data and the GEMS model showed the mean NPP for croplands was over 650 g C m −2 yr −1 while the MODIS NPP product estimated the mean NPP was less than 500 g C m −2 yr −1 . MODIS NPP also showed very different spatial variability of the cropland NPP from the other two methods. We found these differences were mainly caused by the difference in the land cover data and the crop specific information used in the methods. Our study demonstrated that the detailed mapping of the temporal and spatial change of crop species is critical for estimating the spatial and temporal variability of cropland NPP. We suggest that high resolution land cover data with species–specific crop information should be used in satellite-based and process-based models to improve carbon estimates for croplands.

Illinois, Indiana, Iowa, Kansas, Michigan, Minneso

Landscape-level impact of tropical forest loss and fragmentation on bird occurrence in eastern Guatemala

Tropical forest destruction and fragmentation of habitat patches may reduce population persistence at the landscape level. Given the complex nature of simultaneously evaluating the effects of these factors on biotic populations, statistical presence/absence modelling has become an important tool in conservation biology. This study uses logistic regression to evaluate the independent effects of tropical forest cover and fragmentation on bird occurrence in eastern Guatemala. Logistic regression models were constructed for 10 species with varying response to habitat alteration. Predictive variables quantified forest cover, fragmentation and their interaction at three different radii (200, 500 and 1000 m scales) of 112 points where presence of target species was determined. Most species elicited a response to the 1000 m scale, which was greater than most species’ reported territory size. Thus, their presence at the landscape scale is probably regulated by extra-territorial phenomena, such as dispersal. Although proportion of forest cover was the most important predictor of species’ presence, there was strong evidence of area-independent and -dependent fragmentation effects on species presence, results that contrast with other studies from northernmost latitudes. Species’ habitat breadth was positively correlated with AIC model values, indicating a better fit for species more restricted to tropical forest. Species with a narrower habitat breadth also elicited stronger negative responses to forest loss. Habitat breadth is thus a simple measure that can be directly related to species’ vulnerability to landscape modification. Model predictive accuracy was acceptable for 4 of 10 species, which were in turn those with narrower habitat breadths.

Cerro San Gil Watershed Protection Reserve

Accounting for location uncertainty in azimuthaltelemetry data improves ecological inference

Background Characterizing animal space use is critical for understanding ecological relationships. Animal telemetry technology has revolutionized the fields of ecology and conservation biology by providing high quality spatial data on animal movement. Radio-telemetry with very high frequency (VHF) radio signals continues to be a useful technology because of its low cost, miniaturization, and low battery requirements. Despite a number of statistical developments synthetically integrating animal location estimation and uncertainty with spatial process models using satellite telemetry data, we are unaware of similar developments for azimuthal telemetry data. As such, there are few statistical options to handle these unique data and no synthetic framework for modeling animal location uncertainty and accounting for it in ecological models. We developed a hierarchical modeling framework to provide robust animal location estimates from one or more intersecting or non-intersecting azimuths. We used our azimuthal telemetry model (ATM) to account for azimuthal uncertainty with covariates and propagate location uncertainty into spatial ecological models. We evaluate the ATM with commonly used estimators (Lenth (1981) maximum likelihood and M-Estimators) using simulation. We also provide illustrative empirical examples, demonstrating the impact of ignoring location uncertainty within home range and resource selection analyses. We further use simulation to better understand the relationship among location uncertainty, spatial covariate autocorrelation, and resource selection inference. Results We found the ATM to have good performance in estimating locations and the only model that has appropriate measures of coverage. Ignoring animal location uncertainty when estimating resource selection or home ranges can have pernicious effects on ecological inference. Home range estimates can be overly confident and conservative when ignoring location uncertainty and resource selection coefficients can lead to incorrect inference and over confidence in the magnitude of selection. Furthermore, our simulation study clarified that incorporating location uncertainty helps reduce bias in resource selection coefficients across all levels of covariate spatial autocorrelation. Conclusion The ATM can accommodate one or more azimuths when estimating animal locations, regardless of how they intersect; this ensures that all data collected are used for ecological inference. Our findings and model development have important implications for interpreting historical analyses using this type of data and the future design of radio-telemetry studies.

Movement Ecology

Evaluating land cover influences on model uncertainties—A case study of cropland carbon dynamics in the Mid-Continent Intensive Campaign region

Quantifying spatial and temporal patterns of carbon sources and sinks and their uncertainties across agriculture-dominated areas remains challenging for understanding regional carbon cycles. Characteristics of local land cover inputs could impact the regional carbon estimates but the effect has not been fully evaluated in the past. Within the North American Carbon Program Mid-Continent Intensive (MCI) Campaign, three models were developed to estimate carbon fluxes on croplands: an inventory-based model, the Environmental Policy Integrated Climate (EPIC) model, and the General Ensemble biogeochemical Modeling System (GEMS) model. They all provided estimates of three major carbon fluxes on cropland: net primary production (NPP), net ecosystem production (NEP), and soil organic carbon (SOC) change. Using data mining and spatial statistics, we studied the spatial distribution of the carbon fluxes uncertainties and the relationships between the uncertainties and the land cover characteristics. Results indicated that uncertainties for all three carbon fluxes were not randomly distributed, but instead formed multiple clusters within the MCI region. We investigated the impacts of three land cover characteristics on the fluxes uncertainties: cropland percentage, cropland richness and cropland diversity. The results indicated that cropland percentage significantly influenced the uncertainties of NPP and NEP, but not on the uncertainties of SOC change. Greater uncertainties of NPP and NEP were found in counties with small cropland percentage than the counties with large cropland percentage. Cropland species richness and diversity also showed negative correlations with the model uncertainties. Our study demonstrated that the land cover characteristics contributed to the uncertainties of regional carbon fluxes estimates. The approaches we used in this study can be applied to other ecosystem models to identify the areas with high uncertainties and where models can be improved to reduce overall uncertainties for regional carbon flux estimates.

Mid-Continent Intensive Campaign region

Bird migration and energetics simulations incorporating oil spill effects

Oil spills are well-known for causing acute mortality of birds, but sublethal and delayed impacts are less understood. Focusing on the mallard ( Anas platyrhynchos ), we used simulation modeling to explore how sublethal oiling may affect avian survival and breeding ground body condition. We used empirically informed migration and energetics simulations to model hypothetical spills occurring in northern Arkansas, USA occurring in either January to simulate thermoregulatory stress or March to simulate pre-migration effects. We modeled trace and lightly oiled female mallards (≤5% or 6 to 20% of feather area oiled, respectively), incorporating oiling-induced energetic effects on thermoregulation, flight, and energetic gain. We found that mortality was generally higher for simulated spills occurring in January versus March. In the simulations, mallards lost body mass due to oiling, but surviving individuals could partially recover body mass before arriving at the breeding grounds. Including oiling-induced energetic gain effects in simulations increased mortality as well as increased overall variability of simulation results. This modeling effort identified an important gap in knowledge regarding oiled bird energetics, specifically a need to better quantify oiling-induced energetic gain changes. Although the model is currently limited to a specific species and geographic area, it serves as a proof-of-concept for future research and modeling efforts aimed at understanding more broadly the impacts of oil spills on avian populations.

Prairie Pothole region

Simulating 2,368 temperate lakes reveals weak coherence in stratification phenology

Changes in water temperatures resulting from climate warming can alter the structure and function of aquatic ecosystems. Lake-specific physical characteristics may play a role in mediating individual lake responses to climate. Past mechanistic studies of lake-climate interactions have simulated generic lake classes at large spatial scales or performed detailed analyses of small numbers of real lakes. Understanding the diversity of lake responses to climate change across landscapes requires a hybrid approach that couples site-specific lake characteristics with broad-scale environmental drivers. This study provides a substantial advancement in lake ecosystem modeling by combining open-source tools with freely available continental-scale data to mechanistically model daily temperatures for 2,368 Wisconsin lakes over three decades (1979-2011). The model accurately predicted observed surface layer temperatures (RMSE: 1.74°C) and the presence/absence of stratification (81.1% agreement). Among-lake coherence was strong for surface temperatures and weak for the timing of stratification, suggesting individual lake characteristics mediate some - but not all - ecologically relevant lake responses to climate.

Ecological Modelling

Foraging and predation risk for larval cisco (Coregonus artedi) in Lake Superior: A modelling synthesis of empirical survey data

The relative importance of predation and food availability as contributors to larval cisco ( Coregonus artedi ) mortality in Lake Superior were investigated using a visual foraging model to evaluate potential predation pressure by rainbow smelt ( Osmerus mordax ) and a bioenergetic model to evaluate potential starvation risk. The models were informed by observations of rainbow smelt, larval cisco, and zooplankton abundance at three Lake Superior locations during the period of spring larval cisco emergence and surface-oriented foraging. Predation risk was highest at Black Bay, ON, where average rainbow smelt densities in the uppermost 10 m of the water column were >1000 ha &minus;1 . Turbid conditions at the Twin Ports, WI-MN, affected larval cisco predation risk because rainbow smelt remained suspended in the upper water column during daylight, placing them alongside larval cisco during both day and night hours. Predation risk was low at Cornucopia, WI, owing to low smelt densities (<400 ha &minus;1 ) and deep light penetration, which kept rainbow smelt near the lakebed and far from larvae during daylight. In situ zooplankton density estimates were low compared to the values used to develop the larval coregonid bioenergetics model, leading to predictions of negative growth rates for 10 mm larvae at all three locations. The model predicted that 15 mm larvae were capable of attaining positive growth at Cornucopia and the Twin Ports where low water temperatures (2&ndash;6 &deg;C) decreased their metabolic costs. Larval prey resources were highest at Black Bay but warmer water temperatures there offset the benefit of increased prey availability. A sensitivity analysis performed on the rainbow smelt visual foraging model showed that it was relatively insensitive, while the coregonid bioenergetics model showed that the absolute growth rate predictions were highly sensitive to input parameters (i.e., 20% parameter perturbation led to order of magnitude differences in model estimates). Our modelling indicated that rainbow smelt predation may limit larval cisco survival at Black Bay and to a lesser extent at Twin Ports, and that starvation may be a major source of mortality at all three locations. The framework we describe has the potential to further our understanding of the relative importance of starvation and predation on larval fish survivorship, provided information on prey resources available to larvae are measured at sufficiently fine spatial scales and the models provide a realistic depiction of the dynamic processes that the larvae experience.

Lake Superior

Available data support protection of the Southwestern Willow Flycatcher under the Endangered Species Act

Zink (2015) argued there was no evidence for genetic, morphological, or ecological differentiation between the federally endangered Southwestern Willow Flycatcher ( Empidonax traillii extimus ) and other Willow Flycatcher subspecies. Using the same data, we show there is a step-cline in both the frequency of a mtDNA haplotype and in plumage variation roughly concordant with the currently recognized boundary between E. t. extimus and E. t adastus, the subspecies with which it shares the longest common boundary. The geographical pattern of plumage variation is also concordant with previous song analyses differentiating those 2 subspecies and identified birds in one low-latitude, high-elevation site in Arizona as the northern subspecies. We also demonstrate that the ecological niche modeling approach used by Zink yields the same result whether applied to the 2 flycatcher subspecies or to 2 unrelated species, E. t. extimus and Yellow Warbler ( Setophaga petechia ). As a result, any interpretation of those results as evidence for lack of ecological niche differentiation among Willow Flycatcher subspecies would also indicate no differentiation among recognized species and would therefore be an inappropriate standard for delineating subspecies. We agree that many analytical techniques now available to examine genetic, morphological, and ecological differentiation would improve our understanding of the distinctness (or lack thereof) of Willow Flycatcher subspecies, but we argue that currently available evidence supports protection of the Southwestern Willow Flycatcher under the Endangered Species Act.

The Condor

Capturing interactions between nitrogen and hydrological cycles under historical climate and land use: Susquehanna watershed analysis with the GFDL land model LM3-TAN

We developed a process model LM3-TAN to assess the combined effects of direct human influences and climate change on terrestrial and aquatic nitrogen (TAN) cycling. The model was developed by expanding NOAA's Geophysical Fluid Dynamics Laboratory land model LM3V-N of coupled terrestrial carbon and nitrogen (C-N) cycling and including new N cycling processes and inputs such as a soil denitrification, point N sources to streams (i.e., sewage), and stream transport and microbial processes. Because the model integrates ecological, hydrological, and biogeochemical processes, it captures key controls of the transport and fate of N in the vegetation–soil–river system in a comprehensive and consistent framework which is responsive to climatic variations and land-use changes. We applied the model at 1/8° resolution for a study of the Susquehanna River Basin. We simulated with LM3-TAN stream dissolved organic-N, ammonium-N, and nitrate-N loads throughout the river network, and we evaluated the modeled loads for 1986–2005 using data from 16 monitoring stations as well as a reported budget for the entire basin. By accounting for interannual hydrologic variability, the model was able to capture interannual variations of stream N loadings. While the model was calibrated with the stream N loads only at the last downstream Susquehanna River Basin Commission station Marietta (40°02' N, 76°32' W), it captured the N loads well at multiple locations within the basin with different climate regimes, land-use types, and associated N sources and transformations in the sub-basins. Furthermore, the calculated and previously reported N budgets agreed well at the level of the whole Susquehanna watershed. Here we illustrate how point and non-point N sources contributing to the various ecosystems are stored, lost, and exported via the river. Local analysis of six sub-basins showed combined effects of land use and climate on soil denitrification rates, with the highest rates in the Lower Susquehanna Sub-Basin (extensive agriculture; Atlantic coastal climate) and the lowest rates in the West Branch Susquehanna Sub-Basin (mostly forest; Great Lakes and Midwest climate). In the re-growing secondary forests, most of the N from non-point sources was stored in the vegetation and soil, but in the agricultural lands most N inputs were removed by soil denitrification, indicating that anthropogenic N applications could drive substantial increase of N 2 O emission, an intermediate of the denitrification process.

Pennsylvania

Spatially integrating microbiology and geochemistry to reveal complex environmental health issues: Anthrax in the contiguous United States

Maxent models were run using the B. anthracis presence data and/or the animal outbreak presence data. Models run using the animal outbreak data alone utilized two scales: the Outbreak State scale which included only states reporting animal anthrax outbreaks from 2001 to 2013 and the National scale which included all states in the contiguous United States. Three iterations of the environmental data were used and included the Sample Location dataset which utilized the environmental variable data with assigned latitude and longitude locations from the USGS NASGLP project; the Normalized dataset which scaled the environmental variables so that the values fell between 0 and 1; and the Interpolated dataset which provided an interpolation of the environmental variables averaged for each county and assigned to a point for that county at the centroid (rather than using the NASGLP latitude and longitude location). Two metrics were used to measure model performance including the widely used area under the curve (AUC) and an alternative method, the True Skill Statistic (TSS). The AUC gives the probability that a randomly chosen presence location has been correctly ranked higher than the absence/background site. AUC values at 0.5 or lower mean the ranking is no better than random, while the AUC values nearer to 1 mean the model is a better predictor. The TSS provides a comparison of how well the background predictions made by the model match the model results at the test dataset (presence) locations. TSS values near +1 means the model approaches perfect agreement, while values near −1 indicate the model is no better than random. Maxent models to determine the influence of environmental factors on the B. anthracis distribution using the PCR data yielded a low TSS, which suggested the model might be underfitting the data. This was not surprising due to the difficulty in recovering B. anthracis in soil samples as well as the samples themselves being discrete in nature and only capturing a snapshot in time. Therefore, the distribution of B. anthracis and its niche in the contiguous United States could not be determined in this study. However, efforts to investigate environmental factors that would have a higher potential of supporting an anthrax outbreak in wildlife and livestock yielded better results. Results showed that most of the Maxent models in this study performed best when using the Outbreak State scale. When the models were scaled up to the National scale, model performance declined, except for the Normalized variable dataset. At the Outbreak State scale, a large proportion of the area was predicted to be of higher probability for wildlife/livestock anthrax outbreaks, and the statistical measures assumed the model was underfitting the data. The model with the highest AUC and TSS scores for this study was the Outbreak State scale using Sample Location dataset (AUC = 0.918 and TSS = 0.82). Some of the variables found to be closely related to the occurrence of B. anthracis in this study included pH, drainage potential, and concentration of elements including Na, Ca, Sr, and Mg, which have also been found to be related to animal outbreaks or to the occurrence of B. anthracis in previous studies. The models in the current study indicated possible regions that have not had recent wildlife/livestock anthrax outbreaks but contained environmental conditions that could potentially support an outbreak if one were to occur (Michigan and Maine). This work provides an extension to the use of ecological niche modeling to outbreak potential in livestock/wildlife in the United States because it utilizes additional soil geochemistry data and has shown that further validation techniques, such as the TSS, should be considered in addition to AUC. Results from this study could be used by animal and public health officials to identify areas with a higher potential for anthrax outbreak in wildlife and livestock due to naturally occurring soil and environmental conditions.

Book chapter

An evaluation of multistate occupancy models for estimating relative abundance and population trends

Detecting spatiotemporal changes in the abundances of organisms is key to effectively conserving species. While indices of abundance have long been used, there has been a shift toward model-based estimators that account for the detection process. Popular approaches including traditional occupancy models and N-mixture models entail tradeoffs. The traditional occupancy approach requires the researcher coarsen the characterization of abundance to the probability that a site is occupied or unoccupied. Conversely, N-mixture models make use of variation in counts, but perform poorly when individuals have low detectability or move into or out of sites between visits. Multistate occupancy models that differentiate relatively abundant from non-abundant states have the potential to fill this gap but have been underexplored. We conducted a simulation study to test whether multistate occupancy models could capture spatial abundance patterns and detect population declines in the face of low individual detection probability ( p ≤ 0.3) and unmodeled heterogeneity (e.g., that arising from individual movement). We considered 10,773 scenarios to examine the effects of differing amounts of heterogeneity as well as alternative study designs, population parameters, and modeling choices. We tracked bias in the proportion of sites estimated to be in the abundant state for single-season models, and power to detect a declining trend across multiple years. We also evaluated data diagnostic metrics to provide guidance to users. Multistate occupancy models were able to differentiate sites with higher abundances from sites with lower abundances when there were at least medium levels of spatial heterogeneity in true abundances. If different sites were randomly selected each year, power to detect even large population declines (65%) was poor (power < 0.8). However, if the same sites were surveyed each year, and a dynamic multistate occupancy was used, multistate occupancy models could detect (power ≥ 0.8) relatively small declines (5-40%) in 20% of scenarios, and frequently detect large declines of 45-60% (mean power = 0.92). Conservation decisions rely on detecting change reliably, rarely needing absolute abundance information. Multistate occupancy models can improve our ability to detect changing abundance while accommodating low individual detection probability and heterogeneity in count monitoring data.

Ecological Modelling

Balancing predation and egg harvest in a colonial seabird: A simulation model

We developed an individual-based model to study the effects of different regimes of harvesting eggs and natural predation on reproductive success in a colony of the glaucous-winged gull ( Larus glaucescens ) in Glacier Bay National Park, Alaska. The model incorporates the sequence of egg laying, relaying, and incubation to hatching for individual nests and calculates hatching success, incubation length, and the total number of eggs laid (as a result of re-nesting and relaying) in all nests in the colony. Stochasticity is incorporated in the distribution of nest lay dates, predation rates, and nests attacked during predation and harvest events. We estimated parameter values by fitting the model to data collected at a small colony during 1999 and 2000 using maximum likelihood. We then simulated harvests and analyzed model predictions. Model outputs indicate that harvesting early, and at one time, provides a predictable take of eggs with the least impact to gulls.

Ecological Modelling

Exploring behavior of an unusual megaherbivore: A spatially explicit foraging model of the hippopotamus

Herbivore foraging theories have been developed for and tested on herbivores across a range of sizes. Due to logistical constraints, however, little research has focused on foraging behavior of megaherbivores. Here we present a research approach that explores megaherbivore foraging behavior, and assesses the applicability of foraging theories developed on smaller herbivores to megafauna. With simulation models as reference points for the analysis of empirical data, we investigate foraging strategies of the common hippopotamus (Hippopotamus amphibius). Using a spatially explicit individual based foraging model, we apply traditional herbivore foraging strategies to a model hippopotamus, compare model output, and then relate these results to field data from wild hippopotami. Hippopotami appear to employ foraging strategies that respond to vegetation characteristics, such as vegetation quality, as well as spatial reference information, namely distance to a water source. Model predictions, field observations, and comparisons of the two support that hippopotami generally conform to the central place foraging construct. These analyses point to the applicability of general herbivore foraging concepts to megaherbivores, but also point to important differences between hippopotami and other herbivores. Our synergistic approach of models as reference points for empirical data highlights a useful method of behavioral analysis for hard-to-study megafauna. ?? 2003 Elsevier B.V. All rights reserved.

Ecological Modelling

Compartment-based hydrodynamics and water quality modeling of a northern Everglades wetland, Florida, USA

The last remaining large remnant of softwater wetlands in the US Florida Everglades lies within the Arthur R. Marshall Loxahatchee National Wildlife Refuge. However, Refuge water quality today is impacted by pumped stormwater inflows to the eutrophic and mineral-enriched 100-km canal, which circumscribes the wetland. Optimal management is a challenge and requires scientifically based predictive tools to assess and forecast the impacts of water management on Refuge water quality. In this research, we developed a compartment-based numerical model of hydrodynamics and water quality for the Refuge. Using the numerical model, we examined the dynamics in stage, water depth, discharge from hydraulic structures along the canal, and exchange flow among canal and marsh compartments. We also investigated the transport of chloride, sulfate and total phosphorus from the canal to the marsh interior driven by hydraulic gradients as well as biological removal of sulfate and total phosphorus. The model was calibrated and validated using long-term stage and water quality data (1995-2007). Statistical analysis indicates that the model is capable of capturing the spatial (from canal to interior marsh) gradients of constituents across the Refuge. Simulations demonstrate that flow from the eutrophic and mineral-enriched canal impacts chloride and sulfate in the interior marsh. In contrast, total phosphorus in the interior marsh shows low sensitivity to intrusion and dispersive transport. We conducted a rainfall-driven scenario test in which the pumped inflow concentrations of chloride, sulfate and total phosphorus were equal to rainfall concentrations (wet deposition). This test shows that pumped inflow is the dominant factor responsible for the substantially increased chloride and sulfate concentrations in the interior marsh. Therefore, the present day Refuge should not be classified as solely a rainfall-driven or ombrotrophic wetland. The model provides an effective screening tool for studying the impacts of various water management alternatives on water quality across the Refuge, and demonstrates the practicality of similarly modeling other wetland systems. As a general rule, modeling provides one component of a multi-faceted effort to provide technical support for ecosystem management decisions.

Florida

Optimal control of Atlantic population Canada geese

Management of Canada geese ( Branta canadensis ) can be a balance between providing sustained harvest opportunity while not allowing populations to become overabundant and cause damage. In this paper, we focus on the Atlantic population of Canada geese and use stochastic dynamic programming to determine the optimal harvest strategy over a range of plausible models for population dynamics. There is evidence to suggest that the population exhibits significant age structure, and it is possible to reconstruct age structure from surveys. Consequently the harvest strategy is a function of the age composition, as well as the abundance, of the population. The objective is to maximize harvest while maintaining the number of breeding adults in the population between specified upper and lower limits. In addition, the total harvest capacity is limited and there is uncertainty about the strength of density-dependence. We find that under a density-independent model, harvest is maximized by maintaining the breeding population at the highest acceptable abundance. However if harvest capacity is limited, then the optimal long-term breeding population size is lower than the highest acceptable level, to reduce the risk of the population growing to an unacceptably large size. Under the proposed density-dependent model, harvest is maximized by maintaining the breeding population at an intermediate level between the bounds on acceptable population size; limits to harvest capacity have little effect on the optimal long-term population size. It is clear that the strength of density-dependence and constraints on harvest significantly affect the optimal harvest strategy for this population. Model discrimination might be achieved in the long term, while continuing to meet management goals, by adopting an adaptive management strategy.

Ecological Modelling

Migration and energetics model predicts delayed migration and likely starvation in oiled waterbirds

Oil spills can inflict mortality and injury on bird populations; many of these deaths involve starvation resulting from thermoregulatory costs incurred by oiling of birds’ feathers. However, the fates and responses of sublethally oiled birds are poorly known. Due to this knowledge gap and the potential for birds to die far from the spill site, resource risk and injury assessors need tools to make informed estimates for delayed deaths and lost reproductive capacity in these birds. Focusing on the thermoregulatory cost of oiled feathers, we present a model addressing one facet of the effects of sublethal oiling on birds. Using mallard-like ducks as a model organism, we combined values from previous laboratory studies of oiled birds with a modified version of an existing temperature-influenced avian migration energetics model. Using this model, we examined the potential effects of oiling on general migration patterns, changes in energetic gains required to compensate for oiling, and starvation. We assessed all metrics across multiple oiling severities; we assessed starvation across both oiling severity and body condition. Median estimates for delays in spring migration were one to two months for trace and lightly oiled birds, and we predicted arrested spring migration in moderately oiled birds. Median estimates of required increases in energetic gains to offset costs of increased thermoregulation ranged from 20.3% to 88.6% depending on severity of oiling. We predicted starvation within four weeks for most combinations of oiling severity and body condition at the median predicted minimum wintering temperature of unoiled birds (-4.9°C). However, at the average winter temperature of the southernmost model latitude (10.8°C), we predicted only moderately oiled birds in less-than-excellent body condition had the potential to starve within a four-week time frame. Due to the potential for even trace oiling to delay spring migration and decrease body condition, the thermoregulatory costs of sublethal oiling during spring migration could reduce a bird's reproductive capacity. Future research integrating this initial energetics-based model into a spatially explicit, population scale migration model could provide additional insight into the potential effects of sublethal oiling on reproduction and survival. Such an integrated model could strengthen risk predictions and injury assessments for birds subjected to sublethal oiling.

Ecological Modelling

Merging empirical and mechanistic approaches to modeling aquatic visual foraging using a generalizable visual reaction distance model

Visual encounter distance models are important tools for predicting how light and water clarity mediate visual predator-prey interactions that affect the structure and function of aquatic ecosystems at multiple spatial, temporal, and organizational scales. The two main varieties of visual encounter distance models, mechanistic and empirical, are used for similar purposes but take fundamentally different approaches to model development and have different strengths and weaknesses in terms of predictive accuracy, physical and biological interpretability of parameters, ability to incorporate outside information, and utility for knowledge transfer. To overcome weaknesses of existing mechanistic and empirical models and bridge the gap between approaches, we developed a generalized visual reaction distance model that relaxes assumptions of a widely-used mechanistic model that are violated in real predator-prey interactions. We compared the performance of the generalized visual reaction distance model to a widely used mechanistic model and an empirical visual encounter distance model by fitting models to data from four predator-prey experiments. The generalized visual reaction distance model substantially outperformed the other models in all cases based on fit to reaction distance data and presents an attractive alternative to prior models based on comparatively high predictive accuracy, use of interpretable parameters, and ability to incorporate outside information—characteristics that facilitate knowledge transfer.

Ecological Modelling