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Uncertainty in biological monitoring: a framework for data collection and analysis to account for multiple sources of sampling bias

Biological monitoring programmes are increasingly relying upon large volumes of citizen-science data to improve the scope and spatial coverage of information, challenging the scientific community to develop design and model-based approaches to improve inference. Recent statistical models in ecology have been developed to accommodate false-negative errors, although current work points to false-positive errors as equally important sources of bias. This is of particular concern for the success of any monitoring programme given that rates as small as 3% could lead to the overestimation of the occurrence of rare events by as much as 50%, and even small false-positive rates can severely bias estimates of occurrence dynamics. We present an integrated, computationally efficient Bayesian hierarchical model to correct for false-positive and false-negative errors in detection/non-detection data. Our model combines independent, auxiliary data sources with field observations to improve the estimation of false-positive rates, when a subset of field observations cannot be validated a posteriori or assumed as perfect. We evaluated the performance of the model across a range of occurrence rates, false-positive and false-negative errors, and quantity of auxiliary data. The model performed well under all simulated scenarios, and we were able to identify critical auxiliary data characteristics which resulted in improved inference. We applied our false-positive model to a large-scale, citizen-science monitoring programme for anurans in the north-eastern United States, using auxiliary data from an experiment designed to estimate false-positive error rates. Not correcting for false-positive rates resulted in biased estimates of occupancy in 4 of the 10 anuran species we analysed, leading to an overestimation of the average number of occupied survey routes by as much as 70%. The framework we present for data collection and analysis is able to efficiently provide reliable inference for occurrence patterns using data from a citizen-science monitoring programme. However, our approach is applicable to data generated by any type of research and monitoring programme, independent of skill level or scale, when effort is placed on obtaining auxiliary information on false-positive rates.

Methods in Ecology and Evolution

Modelling effects of chemical exposure on birds wintering in agricultural landscapes: The western burrowing owl ( Athene cunicularia hypugaea ) as a case study

We describe an ecotoxicological model that simulates the sublethal and lethal effects of chronic, low-level, chemical exposure on birds wintering in agricultural landscapes. Previous models estimating the impact on wildlife of chemicals used in agro-ecosystems typically have not included the variety of pathways, including both dermal and oral, by which individuals are exposed. The present model contains four submodels simulating (1) foraging behavior of individual birds, (2) chemical applications to crops, (3) transfers of chemicals among soil, insects, and small mammals, and (4) transfers of chemicals to birds via ingestion and dermal exposure. We demonstrate use of the model by simulating the impacts of a variety of commonly used herbicides, insecticides, growth regulators, and defoliants on western burrowing owls ( Athene cunicularia hypugaea ) that winter in agricultural landscapes in southern Texas, United States. The model generated reasonable movement patterns for each chemical through soil, water, insects, and rodents, as well as into the owl via consumption and dermal absorption. Sensitivity analysis suggested model predictions were sensitive to uncertainty associated with estimates of chemical half-lives in birds, soil, and prey, sensitive to parameters associated with estimating dermal exposure, and relatively insensitive to uncertainty associated with details of chemical application procedures (timing of application, amount of drift). Nonetheless, the general trends in chemical accumulations and the relative impacts of the various chemicals were robust to these parameter changes. Simulation results suggested that insecticides posed a greater potential risk to owls of both sublethal and lethal effects than do herbicides, defoliants, and growth regulators under crop scenarios typical of southern Texas, and that use of multiple indicators, or endpoints provided a more accurate assessment of risk due to agricultural chemical exposure. The model should prove useful in helping prioritize the chemicals and transfer pathways targeted in future studies and also, as these new data become available, in assessing the relative danger to other birds of exposure to different types of agricultural chemicals.

Texas

Spatially explicit modelling of floodplain forest succession: Interactions among flood inundation, forest successional processes, and other disturbances in the Upper Mississippi River floodplain, USA

Simulation models are often used to identify hydrologic regimes suitable for different riparian or floodplain tree species. However, most existing models pay little attention to forest successional processes or other disturbances that may interact with the hydrologic regime of river systems to alter forest dynamics in space and time. In this study, we introduce a flood disturbance module to the LANDIS-II forest succession modelling framework to enable investigations into how inundation interacts with other disturbances and successional processes to alter floodplain forest cover and community dynamics. We illustrate the functionality of the model using a case study with multiple scenarios in the Upper Mississippi River floodplain, USA. We found that model predictions of total forest cover and the abundance of specific forest community types were generally related to uncertainty in the susceptibility of different species and age classes to inundation. By simulation year 100, increases or decreases in total forest cover and forest type distributions were roughly proportional to the initial differences in the susceptibility of species and age classes to inundation. The largest decrease in total forest cover was associated with a scenario that included disturbance by the emerald ash borer ( Agrilus planipennis ) and when using susceptibility parameters corresponding to the weakest flood tolerance. In contrast, changes in the composition of aboveground biomass were not sensitive to differences in susceptibility, and generally showed shifts toward later successional species with higher shade tolerance and longer lifespans for all scenarios. Our findings suggest that flood inundation interacts with other disturbances (e.g., insect outbreaks) and forest successional processes to alter forest abundance, distribution, and species composition in this system. Our modelling framework should allow for future studies that examine such interactions in other systems, and in the context of alternative hydrologic scenarios and other disturbance regimes.

Minnesota, Wisconsin

Development and validation of a spatially-explicit agent-based model for space utilization by African savanna elephants (Loxodonta africana) based on determinants of movement

African elephants (Loxodonta africana) are well-studied and inhabit diverse landscapes that are being transformed by both humans and natural forces. Most tools currently in use are limited in their ability to predict how elephants will respond to novel changes in the environment. Individual-, or agent-based modeling (ABM), may extend current methods in addressing and predicting spatial responses to environmental conditions over time. We developed a spatially explicit agent-based model to simulate elephant space use and validated the model with movement data from elephants in Kruger National Park (KNP) and Chobe National Park (CNP). We simulated movement at an hourly scale, as this scale can reflect switches in elephant behavior due to changes in internal states and short-term responses to the local availability and distribution of critical resources, including forage, water, and shade. Known internal drivers of elephant movement, including perceived temperature and the time since an individual last visited a water source, were linked to the external environment through behavior-based movement rules. Simulations were run on model landscapes representing the wet season and the hot, dry season for both parks. The model outputs, including home range size, daily displacement distance, net displacement distance, and maximum distance traveled from a permanent water source, were evaluated through qualitative and quantitative comparisons to actual elephant movement data from both KNP and CNP. The ABM was successful in reproducing the differences in daily displacements between seasons in each park, and in distances traveled from a permanent water source between parks and seasons. Other movement characteristics, including differences in home range sizes and net daily displacements, were partially reproduced. Out of the all the statistical comparisons made between the empirical and simulated movement patterns, the majority were classified as discrepancies of medium or small effect size. We have shown that a resource-driven model with relatively simple decision rules generates trajectories with movement characteristics that are mostly comparable to those calculated from empirical data. Simulating hourly movement (as our model does) may be useful in predicting how finer-scale patterns of space use, such as those created by foraging movements, are influenced by finer spatio-temporal changes in the environment.

Chobe National Park, Kruger National Park

A dynamic landscape model for fish in the Everglades and its application to restoration

A model (ALFISH) for fish functional groups in freshwater marshes of the greater Everglades area of southern Florida has been developed. Its main objective is to assess the spatial pattern of fish densities through time across freshwater marshes. This model has the capability of providing a dynamic measure of the spatially-explicit food resources available to wading birds. ALFISH simulates two functional groups, large and small fish, where the larger ones can prey on the small fish type. Both functional groups are size-structured. The marsh landscape is modeled as 500×500 m spatial cells on a grid across southern Florida. A hydrology model predicts water levels in the spatial cells on 5-day time steps. Fish populations spread across the marsh during flooded conditions and either retreat into refugia (alligator ponds), move to other spatial cells, or die if their cell dries out. ALFISH has been applied to the evaluation of alternative water regulation scenarios under the Central and South Florida Comprehensive Project Review Study. The objective of this Review Study is to compare alternative methods for restoring historical ecological conditions in southern Florida. ALFISH has provided information on which plans are most are likely to increase fish biomass and its availability to wading bird populations.

Ecological Modelling

Elucidating spatially explicit behavioral landscapes in the Willow Flycatcher

Animal resource selection is a complex, hierarchical decision-making process, yet resource selection studies often focus on the presence and absence of an animal rather than the animal's behavior at resource use locations. In this study, we investigate foraging and vocalization resource selection in a population of Willow Flycatchers, Empidonax traillii adastus , using Bayesian spatial generalized linear models. These models produce “behavioral landscapes” in which space use and resource selection is linked through behavior. Radio telemetry locations were collected from 35 adult Willow Flycatchers ( n = 14 males, n = 13 females, and n = 8 unknown sex) over the 2003 and 2004 breeding seasons at Fish Creek, Utah. Results from the 2-stage modeling approach showed that habitat type, perch position, and distance from the arithmetic mean of the home range (in males) or nest site (in females) were important factors influencing foraging and vocalization resource selection. Parameter estimates from the individual-level models indicated high intraspecific variation in the use of the various habitat types and perch heights for foraging and vocalization. On the population level, Willow Flycatchers selected riparian habitat over other habitat types for vocalizing but used multiple habitat types for foraging including mountain shrub, young riparian, and upland forest. Mapping of observed and predicted foraging and vocalization resource selection indicated that the behavior often occurred in disparate areas of the home range. This suggests that multiple core areas may exist in the home ranges of individual flycatchers, and demonstrates that the behavioral landscape modeling approach can be applied to identify spatially and behaviorally distinct core areas. The behavioral landscape approach is applicable to a wide range of animal taxa and can be used to improve our understanding of the spatial context of behavior and resource selection.

Utah

Modeling soil porewater salinity in mangrove forests (Everglades, Florida, USA) impacted by hydrological restoration and a warming climate

Hydrology is a critical driver controlling mangrove wetlands structural and functional attributes at different spatial and temporal scales. Yet, human activities have negatively affected hydrology, causing mangrove diebacks and coverage loss worldwide. In fact, the assessment of mangrove water budgets, impacted by natural and human disturbances, is limited due to a lack of long-term data and information that hinders our understanding of how changes in hydroperiod and salinity control mangrove productivity and spatial distribution. In this study, we implemented a mass balance-based hydrological model (RHYMAN) that explicitly considers groundwater discharge in the Shark River estuary (SRE, southwestern Everglades) located in a karstic geomorphic setting and influenced by regional hydrological restoration. We used long-term hydroperiod and porewater salinity (PWS) datasets obtained from 2004 to 2016 for model calibration and validation and to determine spatiotemporal variability in water levels and PWS at three riverine mangrove sites (downstream, SRS-6; midstream, SRS-5; upstream, SRS-4) along SRE. Model results agree with a distinct PWS pattern along the estuarine salinity gradient where the highest PWS occurs at SRS-6 (mean: 25, range: 22–30 ppt), followed by SRS-5 (17, 14–25 ppt) and SRS-4 (5, 3–13 ppt). A commensurate increase in PWS over a thirteen-year period indicates a long-term reduction in freshwater inflow coupled with sea-level rise (SLR). Increasing freshwater scenario simulation results show a significant reduction (17–27%) in PWS along the estuary in contrast with a high SLR scenario when salinity increases up to 1.1 to 2.5 times that of control values. Model results show that freshwater inflow and SLR are key drivers controlling mangrove wetlands PWS in this karstic coastal region. Given its relatively simple structure, this mass balance-based hydrological model could be used in other environmental settings to evaluate potential habitat and regime shifts due to changes in hydrology and PWS under regional hydrological restoration management.

Florida

Exploring the effect of drought extent and interval on the Florida snail kite: Interplay between spatial and temporal scales

The paper aims at exploring the viability of the Florida snail kite population under various drought regimes in its wetland habitat. The population dynamics of snail kites are strongly linked with the hydrology of the system due to the dependence of this bird species on one exclusive prey species, the apple snail, which is negatively affected by a drying out of habitat. Based on empirical evidence, it has been hypothesised that the viability of the snail kite population critically depends not only on the time interval between droughts, but also on the spatial extent of these droughts. A system wide drought is likely to result in reduced reproduction and increased mortality, whereas the birds can respond to local droughts by moving to sites where conditions are still favourable. This paper explores the implications of this hypothesis by means of a spatially-explicit individual-based model. The specific aim of the model is to study in a factorial design the dynamics of the kite population in relation to two scale parameters, the temporal interval between droughts and the spatial correlation between droughts. In the model high drought frequencies led to reduced numbers of kites. Also, habitat degradation due to prolonged periods of inundation led to lower predicted numbers of kites. Another main result was that when the spatial correlation between droughts was low, the model showed little variability in the predicted numbers of kites. But when droughts occurred mostly on a system wide level, environmental stochasticity strongly increased the stochasticity in kite numbers and in the worst case the viability of the kite population was seriously threatened.

Ecological Modelling

Modeling species invasions in Ecopath with Ecosim: an evaluation using Laurentian Great Lakes models

Invasive species affect the structure and processes of ecosystems they invade. Invasive species have been particularly relevant to the Laurentian Great Lakes, where they have played a part in both historical and recent changes to Great Lakes food webs and the fisheries supported therein. There is increased interest in understanding the effects of ecosystem changes on fisheries within the Great Lakes, and ecosystem models provide an essential tool from which this understanding can take place. A commonly used model for exploring fisheries management questions within an ecosystem context is the Ecopath with Ecosim (EwE) modeling software. Incorporating invasive species into EwE models is a challenging process, and descriptions and comparisons of methods for modeling species invasions are lacking. We compared four methods for incorporating invasive species into EwE models for both Lake Huron and Lake Michigan based on the ability of each to reproduce patterns in observed data time series. The methods differed in whether invasive species biomass was forced in the model, the initial level of invasive species biomass at the beginning of time dynamic simulations, and the approach to cause invasive species biomass to increase at the time of invasion. The overall process of species invasion could be reproduced by all methods, but fits to observed time series varied among the methods and models considered. We recommend forcing invasive species biomass when model objectives are to understand ecosystem impacts in the past and when time series of invasive species biomass are available. Among methods where invasive species time series were not forced, mediating the strength of predator–prey interactions performed best for the Lake Huron model, but worse for the Lake Michigan model. Starting invasive species biomass at high values and then artificially removing biomass until the time of invasion performed well for both models, but was more complex than starting invasive species biomass at low values. In general, for understanding the effect of invasive species on future fisheries management actions, we recommend initiating invasive species biomass at low levels based on the greater simplicity and realism of the method compared to others.

Ecological Modelling

Evaluations of Lagrangian egg drift models: From a laboratory flume to large channelized rivers

To help better interpret computational models in predicting drift of carp eggs in rivers, we present a series of model assessments for the longitudinal egg dispersion. Two three-dimensional Lagrangian particle tracking models, SDrift and FluEgg, are evaluated in a series of channels with increasing complexity. The model evaluation demonstrates that both models are able to accommodate channel complexity and provide a wide range of dispersion coefficients: K l = 0 (1 − 100) Hu ∗ with H being water depth and u ∗ being shear velocity. In a straight channel with K l = 0 (1) Hu ∗ SDrift predicts weaker longitudinal dispersion than FluEgg in the early stage as a result of weak vertical mixing associated with smooth wall turbulence. With sufficient time, SDrift and FluEgg predict similar egg dispersion, accounting for the differential advection due to the vertical velocity profile. In an idealized curved channel with K l = 0 (10) Hu ∗ , dispersion is driven by both vertical and transverse velocity profiles. SDrift yields slightly larger dispersion coefficients than FluEgg. In a real river with channel-training structures and having K l = 0 (100) Hu ∗ SDrift predicts a stronger longitudinal dispersion than FluEgg due to substantial local turbulent eddies and velocity gradients. To summarize, FluEgg shows good performance in capturing dispersion due to vertical velocity profiles and cross-channel velocity gradients. SDrift shows excellent model capabilities of revealing various dispersion mechanisms in addition to the vertical and cross-channel velocity variations. They include the initial turbulent diffusion stage with growing dispersion coefficients and strong dispersion due to in-stream hydraulic structures and localized turbulence.

Missouri

Resolving model parameter values from carbon and nitrogen stock measurements in a wide range of tropical mature forests using nonlinear inversion and regression trees

Objectively assessing the performance of a model and deriving model parameter values from observations are critical and challenging in landscape to regional modeling. In this paper, we applied a nonlinear inversion technique to calibrate the ecosystem model CENTURY against carbon (C) and nitrogen (N) stock measurements collected from 39 mature tropical forest sites in seven life zones in Costa Rica. Net primary productivity from the Moderate-Resolution Imaging Spectroradiometer (MODIS), C and N stocks in aboveground live biomass, litter, coarse woody debris (CWD), and in soils were used to calibrate the model. To investigate the resolution of available observations on the number of adjustable parameters, inversion was performed using nine setups of adjustable parameters. Statistics including observation sensitivity, parameter correlation coefficient, parameter sensitivity, and parameter confidence limits were used to evaluate the information content of observations, resolution of model parameters, and overall model performance. Results indicated that soil organic carbon content, soil nitrogen content, and total aboveground biomass carbon had the highest information contents, while measurements of carbon in litter and nitrogen in CWD contributed little to the parameter estimation processes. The available information could resolve the values of 2-4 parameters. Adjusting just one parameter resulted in under-fitting and unacceptable model performance, while adjusting five parameters simultaneously led to over-fitting. Results further indicated that the MODIS NPP values were compressed as compared with the spatial variability of net primary production (NPP) values inferred from inverse modeling. Using inverse modeling to infer NPP and other sensitive model parameters from C and N stock observations provides an opportunity to utilize data collected by national to regional forest inventory systems to reduce the uncertainties in the carbon cycle and generate valuable databases to validate and improve MODIS NPP algorithms.

Ecological Modelling

An agent-based model to quantify energetics, movement and habitat selection of mid-continent mallards in the Mississippi Alluvial Valley

The dynamics of wintering waterfowl populations at the landscape scale are the result of complex interactions of environmental, behavioral and energetic drivers. Agent-based models provide a method to directly link these factors in a spatially explicit framework and allow the emergence of patterns from the aggregation of individual agent actions. We adapted the Spatially-explicit Waterbird Agent-based Model Program (SWAMP), originally developed for waterfowl in central California, to simulate a basin-scale population of mallards ( A. platyrhynchos ) wintering in the Mississippi Alluvial Valley over a four-month period (November–February). Simulated agents move within the landscape, foraging on areas made available based on a probabilistic inundation status, and converting food resources to endogenous energy. The model uses a high-resolution map of eastern Arkansas waterfowl habitats and incorporates a hierarchical habitat selection system that enables mallards to relocate at increasing scales in response to changing food availability. We validated the performance of modeled mallard body condition and behavioral metrics under a range of environmental conditions against expected outcomes derived from empirical data and found that the simulation produced realistic representations of changes in flight distances, energy expenditure, lipid storage, and foraging habitat use in response to depleting food resources over time. We discuss the model's applicability as a tool to quantify waterfowl response to a range of environmental conditions and to evaluate scenarios of landscape composition and configuration in the context of waterfowl population management.

Ecological Modelling

Allowing macroalgae growth forms to emerge: Use of an agent-based model to understand the growth and spread of macroalgae in Florida coral reefs, with emphasis on Halimeda tuna

The growth patterns of macroalgae in three-dimensional space can provide important information regarding the environments in which they live, and insights into changes that may occur when those environments change due to anthropogenic and/or natural causes. To decipher these patterns and their attendant mechanisms and influencing factors, a spatially explicit model has been developed. The model SPREAD (SPatially-explicit Reef Algae Dynamics), which incorporates the key morphogenetic characteristics of clonality and morphological plasticity, is used to investigate the influences of light, temperature, nutrients and disturbance on the growth and spatial occupancy of dominant macroalgae in the Florida Reef Tract. The model species, Halimeda and Dictyota spp., are modular organisms, with an 'individual' being made up of repeating structures. These species can also propagate asexually through clonal fragmentation. These traits lead to potentially indefinite growth and plastic morphology that can respond to environmental conditions in various ways. The growth of an individual is modeled as the iteration of discrete macroalgal modules whose dynamics are affected by the light, temperature, and nutrient regimes. Fragmentation is included as a source of asexual reproduction and/or mortality. Model outputs are the same metrics that are obtained in the field, thus allowing for easy comparison. The performance of SPREAD was tested through sensitivity analysis and comparison with independent field data from four study sites in the Florida Reef Tract. Halimeda tuna was selected for initial model comparisons because the relatively untangled growth form permits detailed characterization in the field. Differences in the growth patterns of H. tuna were observed among these reefs. SPREAD was able to closely reproduce these variations, and indicate the potential importance of light and nutrient variations in producing these patterns. ?? 2008 Elsevier B.V.

Ecological Modelling

Defining the next generation modeling of coastal ecotone dynamics in response to global change

Coastal ecosystems are especially vulnerable to global change; e.g., sea level rise (SLR) and extreme events. Over the past century, global change has resulted in salt-tolerant (halophytic) plant species migrating into upland salt-intolerant (glycophytic) dominated habitats along major rivers and large wetland expanses along the coast. While habitat transitions can be abrupt, modeling the specific drivers of abrupt change between halophytic and glycophytic vegetation is not a simple task. Correlative studies, which dominate the literature, are unlikely to establish ultimate causation for habitat shifts, and do not generate strong predictive capacity for coastal land managers and climate change adaptation exercises. In this paper, we first review possible drivers of ecotone shifts for coastal wetlands, our understanding of which has expanded rapidly in recent years. Any exogenous factor that increases growth or establishment of halophytic species will favor the ecotone boundary moving upslope. However, internal feedbacks between vegetation and the environment, through which vegetation modifies the local microhabitat (e.g., by changing salinity or surface elevation), can either help the system become resilient to future changes or strengthen ecotone migration. Following this idea, we review a succession of models that have provided progressively better insight into the relative importance of internal positive feedbacks versus external environmental factors. We end with developing a theoretical model to show that both abrupt environmental gradients and internal positive feedbacks can generate the sharp ecotonal boundaries that we commonly see, and we demonstrate that the responses to gradual global change (e.g., SLR) can be quite diverse.

Ecological Modelling

A population model for a long-lived, resprouting chaparral shrub: Adenostoma fasciculatum

Extensive stands of Adenostoma fasciculatum H.&A. (chamise) in the chaparral of California are periodically rejuvenated by fire. A population model based on size-specific demographic characteristics (thinning and fire-caused mortality) was developed to generate probable age distributions within size classes and survivorship curves for typical stands. The model was modified to assess the long term effects of different mortality rates on age distributions. Under observed mean mortality rates (28.7%), model output suggests some shrubs can survive more than 23 fires. A 10% increase in mortality rate by size class slightly shortened the survivorship curve, while a 10% decrease in mortality rate by size class greatly elongated the curve. This approach may be applicable to other long-lived plant species with complex life histories.

Ecological Modelling

Simulating the impact of cholinesterase-inhibiting pesticides on non-target wildlife in irrigated crops

We present a simulation model for risk assessment of the impact of insecticide inhibitors of cholinesterase (ChE) applied in irrigated agricultural fields on non-target wildlife. The model, which we developed as a compartment model based on difference equations (??t = 1 h), consists of six submodels describing the dynamics of (1) insecticide application, (2) insecticide movement into floodable soil, (3) irrigation and rain, (4) insecticide dissolution in water, (5) foraging and insecticide intake from water, and (6) ChE inhibition and recovery. To demonstrate application of the model, we simulated historical and "worst-case" scenarios of the impact of ChE-inhibiting insecticides on white-winged doves (Zenaida asiatica) inhabiting natural brushland adjacent to cotton and sugarcane fields in the Lower Rio Grande Valley of Texas, USA. Only when a rain event occurred just after insecticide application did predicted levels of ChE inhibition surpass the diagnostic level of 20% exposure. The present model should aid in assessing the effect of ChE-inhibiting insecticides on ChE activity of different species that drink contaminated water from irrigated agricultural fields, and in identifying specific situations in which the juxtaposition of environmental conditions and management schemes could result in a high risk to non-target wildlife. ?? 2007 Elsevier B.V. All rights reserved.

Ecological Modelling

An epidemiological model of virus transmission in salmonid fishes of the Columbia River Basin

We have developed a dynamic epidemiological model informed by records of viral presence and genotypes to evaluate potential transmission routes maintaining a viral pathogen in economically and culturally important anadromous fish populations. In the Columbia River Basin, infectious hematopoietic necrosis virus (IHNV) causes severe disease, predominantly in juvenile steelhead trout ( Oncorhynchus mykiss ) and less frequently in Chinook salmon ( O. tshawytscha ). Mortality events following IHNV infection can be devastating for individual hatchery programs. Despite reports of high local mortality and extensive surveillance efforts, there are questions about how viral transmission is maintained. Modeling this system offers important insights into disease transmission in natural aquatic systems, as well as about the data requirements for generating accurate estimates about transmission routes and infection probabilities. We simulated six scenarios in which testing rates and the relative importance of different transmission routes varied. The simulations demonstrated that the model accurately identified routes of transmission and inferred infection probabilities accurately when there was testing of all cohort-sites. When testing records were incomplete, the model accurately inferred which transmission routes exposed particular cohort-sites but generated biased infection probabilities given exposure. After validating the model and generating guidelines for result interpretation, we applied the model to data from 14 annual cohorts (2000–2013) at 24 focal sites in a sub-region of the Columbia River Basin, the lower Columbia River (LCR), to quantify the relative importance of potential transmission routes in this focal sub-region. We demonstrate that exposure to IHNV via the return migration of adult fish is an important route for maintaining IHNV in the LCR sub-region, and the probability of infection following this exposure was relatively high at 0.16. Although only 1% of cohort-sites experienced self-exposure by infected juvenile fish, this transmission route had the greatest probability of infection (0.22). Increased testing and/or determining whether transmission can occur from cohort-sites without testing records (e.g., determining there was no testing record because there were no fish at the cohort-site) are expected to improve inference about infection probabilities. Increased use of secure water supplies and continued use of biosecurity protocols may reduce IHNV transmission from adult fish and juvenile fish within the site, respectively, to juvenile salmonids at hatcheries. Models and conclusions from this study are potentially relevant to understanding the relative importance of transmission routes for other important aquatic pathogens in salmonids, including the agents of bacterial kidney disease and coldwater disease, and the basic approach may be useful for other pathogens and hosts in other geographic regions.

Ecological Modelling

A fully-stochasticized, age-structured population model for population viability analysis of fish: Lower Missouri River endangered pallid sturgeon example

We develop a fully-stochasticized, age-structured population model suitable for population viability analysis (PVA) of fish and demonstrate its use with the endangered pallid sturgeon ( Scaphirhynchus albus ) of the Lower Missouri River as an example. The model incorporates three levels of variance: parameter variance (uncertainty about the value of a parameter itself) applied at the iteration level, temporal variance (uncertainty caused by random environmental fluctuations over time) applied at the time-step level, and implicit individual variance (uncertainty caused by differences between individuals) applied within the time-step level. We found that population dynamics were most sensitive to survival rates, particularly age-2+ survival, and to fecundity-at-length. The inclusion of variance (unpartitioned or partitioned), stocking, or both generally decreased the influence of individual parameters on population growth rate. The partitioning of variance into parameter and temporal components had a strong influence on the importance of individual parameters, uncertainty of model predictions, and quasiextinction risk (i.e., pallid sturgeon population size falling below 50 age-1+ individuals). Our findings show that appropriately applying variance in PVA is important when evaluating the relative importance of parameters, and reinforce the need for better and more precise estimates of crucial life-history parameters for pallid sturgeon.

Lower Missouri River