THE VALUE OF SHORT‐RUN CLIMATE FORECASTS IN MANAGING THE COASTAL COHO SALMON (ONCORHYNCHUS KISUTCH) FISHERY IN WASHINGTON STATE

ABSTRACT. . In recent years our understanding of the intricate connections between climate variability, marine and freshwater environmental conditions and the responses of fish stocks has improved considerably. With predictable relationships between the environment and stock abundance, fishery managers should be able to forecast variation in stock survival and recruitment. Such forecasts present an opportunity for increasing the economic value of fisheries and for achieving other management objectives, such as stock conservation and maintenance of population diversity. After describing a 4‐step framework for addressing the question ‘What is a forecast worth?’ in a fishery decision‐making context, we introduce the management system for Washington's coastal coho salmon (Oncorhynchus kisutch) fishery. Then we apply the 4‐step framework to estimate the value of improved run size forecasts in the annual harvest management of coho salmon in Washington State. Our principal analytical tool is a stochastic simulation model that incorporates the main characteristics of the fishery. The paper concludes with a discussion of opportunities and constraints to the use of climate‐based forecasts in fishery management on various spatial and temporal scales, and we consider the challenges associated with forecasting variations in fish stock size caused by shifts in climate and related ocean conditions.     

A JUVENILE‐ADULT DISCRETE‐TIME PRODUCTION MODEL OF EXPLOITED FISHERY SYSTEMS

Abstract. Previous mathematical modeling of the population dynamics of Georges Bank Atlantic cod fishery employed discrete‐time models without age‐structure. To make use of a much wider variety of data on fisheries and fish stocks than was possible with an unstructured model, we introduce a juvenile‐adult age‐structured production exploited fishery model with a very general recruitment function. We use the age‐structured model to study the interaction between fish exploitation levels and recruitment dynamics. As case studies, we use our model results and historical fish population data from Georges Bank to investigate the impact of recent harvesting levels on the sustainability of cod fishery. We show that a constant harvesting policy with the same harvesting rate of 2007 would lead to the recovery and sustainability of Georges Bank cod fishery.     

EQUILIBRIUM ABUNDANCE OF SALMON STOCKS IN A LIFE CYCLE MODEL WITH INTERACTING HATCHERY AND NATURAL SUBSTOCKS

This paper develops the equilibrium solutions for an age-structured life cycle model where spawning stock is split between natural and hatchery spawners. Mixing is allowed between the stocks through natural stock take by the hatchery and release of eggs or fry by the hatchery when its capacity is exceeded. The natural stock is assumed to have density-dependent egg-smolt survival while the hatchery stock has linear survival. The model can be applied to any hatchery reared fish stock but is most appropriate for salmon, where hatchery and naturally spawned fish mix completely later in life. Questions about the mix between the hatchery and natural stocks can be addressed by computing the fraction of naturally and hatchery derived stock among the natural and hatchery spawners as well as among the total adult run. Columbia River chinook stock are used as an example for which equilibria and mixing fractions are computed. A Monte Carlo sensitivity study on model parameters showed that the natural stock survival from smoltification to age 1 and the natural basin smolt carrying capacity are most important in controlling the equilibrium age-1 naturally spawned stock. Changing hatchery capacity over two orders of magnitude showed a 50 percent change in the fraction of naturally derived fish in the natural spawning stock, while the relative size of natural and hatchery stocks changed over two orders of magnitude. The model can serve as a tool for quickly assessing the effects of spawning habitat modification and hatchery supplementation practices on long-term stock mixing and stock abundance.     

THE INCOMPLETE‐INFORMATION SPLIT‐STREAM FISH WAR: EXAMINING THE IMPLICATIONS OF COMPETING RISKS

ABSTRACT. . It is now widely recognized that climactic regime shifts, which aperiodically alter a harvested fish stock's biomass and spatial distribution, may lead to distorted fisheries management decisions which negatively impact the fishery, both biologically and economically. This is particularly true for trans‐boundary migratory stocks, where optimal management relies on coordination among independent nation‐states. Unanticipated changes in stock distribution and abundance can upset expectations of national authorities, leading them to sanction inappropriate harvesting levels by their separately managed fleets targeting the same breeding fish stock. Our theoretical studies are based on a spatially‐distributed stochastic model, which we have called the “split‐stream model,‘ where two separately managed fleets harvest simultaneously at two separate sites. Our key assumption is that competing fleet managers, when harvesting noncooperatively, hold incomplete and asymmetric private information of current stock recruitment and spatial distribution. When subsequently negotiating to coordinate their harvests, they agree that they will share their information and then bargain over partition of the gains from their cooperation. This bargaining process takes into account the fleet's relative competitive strengths, particularly due to private information asymmetries. In this present article we introduce a more complex information structure than had been assumed in our earlier work (McKelvey and Golubtsov [2002], McKelvey, Miller and Golubtsov [2003], Mckelvey et al. [2004]). Specifically, both stock‐growth and stock‐split parameters vary stochastically and asynchronously. Thus, when harvesting noncooperatively, each fleet may possess private knowledge which is unavailable to the other. We examine the interplay of the harvesting game's information structure with other fishery characteristics, such as the fleets' economics and operating characteristics and their attitudes toward risk, to determine the implications of such structure for the outcome of the harvesting game. All of these changes are made to capture new conceptual phenomena and expand the range of applicability of the model.     

Game chromatic index of k‐degenerate graphs

We consider the following edge coloring game on a graph G. Given t distinct colors, two players Alice and Bob, with Alice moving first, alternately select an uncolored edge e of G and assign it a color different from the colors of edges adjacent to e. Bob wins if, at any stage of the game, there is an uncolored edge adjacent to colored edges in all t colors; otherwise Alice wins. Note that when Alice wins, all edges of G are properly colored. The game chromatic index of a graph G is the minimum number of colors for which Alice has a winning strategy. In this paper, we study the edge coloring game on k‐degenerate graphs. We prove that the game chromatic index of a k‐degenerate graph is at most Δ + 3k − 1, where Δ is the maximum vertex degree of the graph. We also show that the game chromatic index of a forest of maximum degree 3 is at most 4 when the forest contains an odd number of edges. © 2001 John Wiley & Sons, Inc. J Graph Theory 36: 144–155, 2001     

Evaluating impacts of fish stock enhancement and biodiversity conservation actions on the livelihoods of small‐scale fishers on the Beijiang River,China

Inappropriate development and overexploitation have seriously degraded aquatic resources in China. Stakeholders identified three fish stock enhancement and biodiversity conservation scenarios for the Beijiang River: S1, increased fish restocking; S2, no fishing season and habitat conservation; and S3, strict pollution control. Potential impacts of these actions on the livelihoods of fishers were evaluated using applied economic modeling. Baseline costs and benefits came from logbooks from 30 fishers and a survey of 90 households in three villages. The financial net benefit for a household was US$1583 (￥11,160) annually, representing a 142% and 387% return on capital and operating costs, respectively. Larger catches associated with S1 and S2 generated a net benefit of US$1651 and US$1822, respectively. Strict pollution control resulting in higher catches (+20%) and lower operating costs (?20%) would increase the net benefit by 15.9% to US$1835 annually. Pollution control would benefit other resource users and is a prerequisite for ecological restoration. Recommendations for Resource Managers

• Stringent pollution control measures are essential to conserve aquatic biodiversity and enhance the livelihoods of fishers but will require considerable public and private sector investment.
• Enhanced fish stocks in the Beijiang River could benefit poor livelihoods but may not be sufficient to lift households out of poverty, aged fishers require government assistance to diversify their livelihoods, access alternative urban employment, and survive with dignity.
• Adopting the economic modeling approach presented here could enable responsible authorities to simultaneously evaluate fish stock enhancement and biodiversity conservation options.
• Broader application of the approach presented here could help ensure small‐scale inland fisheries are managed sustainably and aquatic ecosystems are restored and protected by 2020, in line with Target 6 of the United Nations’ Convention on Biological Diversity Strategic Plan for Biodiversity.

     

FORECASTING TREE SIZE DISTRIBUTION BY NUMERICAL APPROXIMATION OF THE FORWARD KOLMOGOROV EQUATION

ABSTRACT. Forest management planning of uneven‐aged stands involves forecasting of the tree size distribution. The temporal development of the size distribution in a forest stand may be described by the forward Kolmogorov equation. The objective of this study is to illustrate that numerical approximation of the solution to the equation provides a reasonably accurate way of forecasting future tree size distribution, especially for stands with non‐normal size distribution. Furthermore, a method for the practical application is devised. The analyses compare observed and forecasted tree size distributions for two forest stands, 1) an unthinned stand of Sitka spruce (Picea sitchensis (Bong.) Carr.), and 2) an uneven‐aged stand of beech (Fagus sylvatica L.) managed under the selection system. The analyses show that the size distribution in the uneven‐aged stand may be forecasted correctly for a 20 25 year period, while for the even‐aged stand the method seems to fail after 10 to 15 years.     

HABITAT PATCH OCCUPANCY DYNAMICS OF GLAUCOUS‐WINGED GULLS (LARUS GLAUCESCENS) II: A CONTINUOUS‐TIME MODEL

ABSTRACT. The diurnal distribution and abundance dynamics of loafing Glaucous‐winged Gulls (Larus glaucescens) were examined at Protection Island National Wildlife Refuge, Strait of Juan de Fuca, Washington. Asynchronous movement of gulls among three habitat patches dedicated to loafing was modeled as a function of environmental variables using differential equations. Multiple time scale analysis led to the derivation of algebraic models for habitat patch occupancy dynamics. The models were parameterized with hourly census data collected from each habitat patch, and the resulting model predictions were compared with observed census data. A four‐compartment model explained 41% of the variability in the data. Models that predict the dynamics of organism distribution and abundance enhance understanding of the temporal and spatial organization of ecological systems, as well as the decision‐making process in natural resource management.     

OPTIMAL HARVEST FEEDBACK RULE ACCOUNTING FOR THE FISHING‐UP EFFECT AND SIZE‐DEPENDENT PRICING

Abstract Fishing leads to truncation of a population's age and size structure. However, large‐sized fish are usually more valuable per unit weight than small ones. Nevertheless, these size‐related factors have mostly been ignored in bioeconomic modeling. Here, we present a simple extension to the Gordon–Schaefer model that accounts for variations in mean individual catch weight, and derive the feedback rule for optimal harvest in this setting. As the Gordon–Schaefer model has no population structure, size effects have to be accounted for indirectly. Here we assume a simple negative relationship between fishing effort and mean individual weight, and a positive relationship between mean catch weight and price. The aim is to emulate alterations of size structure in fish populations due to fishing and the influence of size on price per weight unit and eventually, net revenues. This demonstrates, on a general level, how such size‐dependent effects change the patterns of optimal harvest paths and sustainable revenue in single fish stocks. The model shows clear shifts toward lower levels of optimal effort and yield compared to classical models without size effects. This suggests that ignoring body size could lead to misleading assumptions and policies, potentially causing rent dissipation and suboptimal utilization of renewable resources.     

A MODEL OF CHINOOK SALMON POPULATION DYNAMICS INCORPORATING SIZE‐SELECTIVE EXPLOITATION AND INHERITANCE OF POLYGENIC CORRELATED TRAITS

Abstract Concern regarding the potential for selective fisheries to degrade desirable characteristics of exploited fish populations is growing worldwide. Although the occurrence of fishery‐induced evolution in a wild population has not been irrefutably documented, considerable theoretical and empirical evidence for that possibility exists. Environmental conditions influence survival and growth in many species and may mask comparatively subtle trends induced by selective exploitation, especially given the evolutionarily short time series of data available from many fisheries. Modeling may be the most efficient investigative tool under such conditions. Motivated by public concern that large‐mesh gillnet fisheries may be altering Chinook salmon in western Alaska, we constructed a stochastic model of the population dynamics of Chinook salmon. The model contained several individually based components and incorporated size‐selective exploitation, assortative mating, size‐dependent female fecundity, density‐dependent survival, and the heritability of size and age. Substantial reductions in mean size and age were observed under all scenarios. Concurrently reducing directional selection and increasing spawning abundance was most effective in stimulating population recovery. Use of this model has potential to improve our ability to investigate the consequences of selective exploitation and aid development of improved management strategies to more effectively sustain fish and fisheries into the future.     

DO ACCURATE STOCK ESTIMATES INCREASE HARVEST AND REDUCE VARIABILITY IN FISHERIES YIELDS?

Abstract Fisheries managers normally make decisions based on stock abundance estimates subject to process, observation, and model uncertainties. Considerable effort is invested in gathering information about stock size to decrease these uncertainties. However, few studies have evaluated benefits from collecting such information in terms of yield and stability of annual harvest. Here, we develop a strategic age‐structured population model for a long‐lived fish with stochastic recruitment, resembling the Norwegian spring‐spawning herring (NSSH, Clupea harengus L.). We evaluate how uncertainties in population estimates influence annual yield, spawning stock biomass (SSB), and variation in annual harvest, using both the proportional threshold harvesting (PTH) and the current harvest control rule for NSSH as harvest strategies. Results show that the consequences of a biased estimate are sensitive to the harvest strategy employed. If the harvest strategy is suitably chosen, the benefits of accurate information are low, and less information about the stock is necessary to maintain high average yield. Reduced harvest intensity effectively removes the need for accurate stock estimates. PTH (a variant of the constant escapement strategy) with low harvest ratio and the current NSSH harvest control rule both provide remarkable stability in yield and SSB. However, decreased uncertainty will often decrease year‐to‐year variation in harvest and the frequency of fishing moratoria.     

SHARING THE NORTHEAST ARCTIC COD: POSSIBLE EFFECTS OF CLIMATE CHANGE

ABSTRACT. The Northeast Arctic cod inhabits the exclusive economic zones of Norway and Russia and migrates extensively between these zones. The stock is shared evenly between the two countries, with a small allocation to third countries. Higher temperatures in the Barents Sea and the Norwegian Sea are expected to affect the stock, probably increasing its size and leading to a larger share inhabiting the Russian economic zone. It is also conceivable that some spawning will begin to take place off the coast of Russia in addition to the spawning that now occurs exclusively in Norwegian waters. This paper looks at the implication of this for the division of the stock between the two countries. It is found that a greater presence of the stock in the Russian zone would strengthen rather than weaken the Norwegian bargaining position if the unit cost of fish is not sensitive to the size of the stock. If, on the other hand, the fishing costs are proportional to fishing mortality, Norway's position would be weakened almost on par with the fall in its share of the stock. The paper uses a Beverton‐Holt year class model with a Ricker recruitment function. The recruitment function is hump‐shaped, implying that a too large spawning stock is harmful for recruitment. Strong density‐dependence in the survival of eggs and larvae is a possible reason for this. It is shown that, for a stock being limited by carrying capacity at the pre‐recruit stage rather than the post‐recruit stage, one may expect a strongly asymmetric curve for sustainable yield as a function of total biomass. The biomass of an exploited population might possibly exceed the biomass of a pristine population under those circumstances.     

A STATE‐SPACE MODEL FOR UNIVARIATE ORDINAL‐VALUED TIME SERIES

Abstract A state‐space model was developed to analyze a univariate time series of ordinal‐valued flower phenology data. Flower abundance, recorded as either “none,”“some,” or “much,” was observed each month on trees in a lowland Costa Rican rain forest to investigate flowering patterns of different species. Data from a single Capparis pittieri and daily rain measurements are used to demonstrate the model. A method to obtain maximum likelihood estimates of the parameters and the use of predicted probability differences to assess goodness of fit are described. Opportunities for improving the model are also discussed.     

THE IMPACT OF AGE-, SPACE-AND STOCHASTIC-STRUCTURE ON THE EXTINCTION PROBABILITY OF A CHINOOK SALMON METAPOPULATION

ABSTRACT. Using a mechanistic model, based on chinook life history, incorporating environmental and demographic stochasticity, we investigate how the probability of extinction is controlled by age, space and stochastic structure. Environmental perturbations of age dependent survivorships, combined with mixing of year classes in the spawning population, can lower the probability of extinction dramatically. This is an analog of the more familiar metapopulation result where dispersal between asynchronously fluctuating populations enhances persistence. For a two-river chinook metapopulation, dispersal between rivers with asynchronous environmental perturbations also dramatically enhances persistence, and anti-synchronous population fluctuations provide an even greater persistence probability. Anti-synchronous fluctuations would most likely occur in pristine habitat with naturally high levels of heterogeneity. Fifty percent dispersal between two populations provides the greatest insurance against extinction, a rate unrealistically high for salmon. In contrast, dispersal between exactly correlated populations with large amplitude environmental perturbations does not help persistence, no matter how high the dispersal rate. This is in spite of weak asynchrony provided by demographic stochasticity. Dispersal between rivers, one degraded and the other pristine, can substantially increase the probability of metapopulation extinction. Population structure, combined with asynchronous environmental perturbations and dispersal (or age class mixing) lowers the probability of chinook extinction dramatically but is almost useless when survivorships are impaired.     

Maximum Δ‐edge‐colorable subgraphs of class II graphs

A graph G is class II, if its chromatic index is at least Δ + 1. Let H be a maximum Δ‐edge‐colorable subgraph of G. The paper proves best possible lower bounds for |E(H)|/|E(G)|, and structural properties of maximum Δ‐edge‐colorable subgraphs. It is shown that every set of vertex‐disjoint cycles of a class II graph with Δ≥3 can be extended to a maximum Δ‐edge‐colorable subgraph. Simple graphs have a maximum Δ‐edge‐colorable subgraph such that the complement is a matching. Furthermore, a maximum Δ‐edge‐colorable subgraph of a simple graph is always class I. © 2011 Wiley Periodicals, Inc. J Graph Theory     

TOWARDS A STANDARD FOR THE INDIVIDUAL‐BASED MODELING OF PLANT POPULATIONS: SELF‐THINNING AND THE FIELD‐OF‐NEIGHBORHOOD APPROACH

ABSTRACT. In classical theoretical ecology there are numerous standard models which are simple, generally applicable, and have well‐known properties. These standard models are widely used as building blocks for all kinds of theoretical and applied models. In contrast, there is a total lack of standard individual‐based models (IBM's), even though they are badly needed if the advantages of the individual‐based approach are to be exploited more efficiently. We discuss the recently developed ‘field‐of‐neighborhood’ approach as a possible standard for modeling plant populations. In this approach, a plant is characterized by a circular zone of influence that grows with the plant, and a field of neighborhood that for each point within the zone of influence describes the strength of competition, i.e., growth reduction, on neighboring plants. Local competition is thus described phenomenologically. We show that a model of mangrove forest dynamics, KiWi, which is based on the FON approach, is capable of reproducing self‐thinning trajectories in an almost textbook‐like manner. In addition, we show that the entire biomass‐density trajectory (bdt) can be divided into four sections which are related to the skewness of the stem diameter distributions of the cohort. The skewness shows two zero crossings during the complete development of the population. These zero crossings indicate the beginning and the end of the self‐thinning process. A characteristic decay of the positive skewness accompanies the occurrence of a linear bdt section, the well‐known self‐thinning line. Although the slope of this line is not fixed, it is confined in two directions, with morphological constraints determining the lower limit and the strength of neighborhood competition exerted by the individuals marking the upper limit.     

HOW BIG AND HOW CLOSE? HABITAT PATCH SIZE AND SPACING TO CONSERVE A THREATENED SPECIES

Abstract. We present results of a spatially explicit, individual‐based stochastic dispersal model (HexSim) to evaluate effects of size and spacing of patches of habitat of Northern Spotted Owls (NSO; Strix occidentalis caurina) in Pacific Northwest, USA, to help advise recovery planning efforts. We modeled 31 artificial landscape scenarios representing combinations of NSO habitat cluster size (range 4–49 NSO pairs per cluster) and edge‐to‐edge cluster spacing (range 7–101 km), and an all‐habitat landscape. We ran scenarios using empirical estimates of NSO dispersal dynamics and distances and stage class vital rates (representing current population declines) and under adult survival rates adjusted to achieve an initially stationary population. Results suggested that long‐term (100‐yr) habitat occupancy rates are significantly higher with habitat clusters supporting ≥25 NSO pairs and ≤15 km spacing, and with overall landscapes of ≥35–40% habitat. Although habitat provision is key to NSO recovery, no habitat configuration provided for long‐term population persistence when coupled with currently observed vital rates. Results also suggested a key role of floaters (unpaired, nonterritorial, dispersing owls) in recolonizing vacant habitat, and that the floater population segment becomes increasingly depleted with greater population declines. We suggest additional areas of modeling research on this and other threatened species.     

ANALYSIS OF SENSITIVITY AND UNCERTAINTY IN AN INDIVIDUAL‐BASED MODEL OF A THREATENED WILDLIFE SPECIES

Sensitivity analysis—determination of how prediction variables affect response variables—of individual‐based models (IBMs) are few but important to the interpretation of model output. We present sensitivity analysis of a spatially explicit IBM (HexSim) of a threatened species, the Northern Spotted Owl (NSO; Strix occidentalis caurina) in Washington, USA. We explored sensitivity to HexSim variables representing habitat quality, movement, dispersal, and model architecture; previous NSO studies have well established sensitivity of model output to vital rate variation. We developed “normative” (expected) model settings from field studies, and then varied the values of ≥ 1 input parameter at a time by ±10% and ±50% of their normative values to determine influence on response variables of population size and trend. We determined time to population equilibration and dynamics of populations above and below carrying capacity. Recovery time from small population size to carrying capacity greatly exceeded decay time from an overpopulated condition, suggesting lag time required to repopulate newly available habitat. Response variables were most sensitive to input parameters of habitat quality which are well‐studied for this species and controllable by management. HexSim thus seems useful for evaluating potential NSO population responses to landscape patterns for which good empirical information is available.     

MODELING A RIGHTS‐BASED APPROACH FOR MANAGING HABITAT IMPACTS OF FISHERIES

ABSTRACT. Limiting adverse consequences of fishing on essential fish habitat has emerged as a key fishery management objective. The conventional approach to providing habitat protection is to create MPAs or marine reserves that prohibit all or certain types of fishing in specific areas. However, there may be more cost‐effective and flexible ways to provide habitat protection. We propose an individual habitat quota (IHQ) system for habitat conservation that would utilize economic incentives to achieve habitat conservation goals cost‐effectively. Individual quotas of habitat impact units (HIU) would be distributed to fishers with an aggregate quota set to maintain a target habitat “stock.” HIU use would be based on a proxy for marginal habitat damage. We use a dynamic, explicitly spatial fishery and habitat simulation model to explore how such a system might work. We examine how outcomes are affected by spatial heterogeneity in the fishery and the scale of habitat regulation. We find that the IHQ system is a highly cost‐effective means of ensuring a given level of habitat protection, but that spatial heterogeneity and the scale of regulation can have significant effects on the distribution of habitat protection.