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.     

GOLDEN AGES OR MAGIC MOMENTS?

ABSTRACT. The paper is an attempt to refute Quinn's ‘the Golden Age of fisheries population models has ended “hypothesis. The approach is to show it is possible to provide simple matrix based theory for use with delay difference equations applied to products of the 0th to 4th powers of length and numbers at length (Alias” proto‐moments“). It compares these to models just based upon numbers and biomass. It also shows examples (of comparatively little moment) indicating how using multiple proto‐moments could improve the ability of delay difference equations to handle the size dependent processes of maturity, selection, natural mortality rates and predation rates and thus improving the biological reality of these models.     

SUSTAINABLE YIELDS IN FISHERIES: UNCERTAINTY,RISK‐AVERSION,AND MEAN‐VARIANCE ANALYSIS

Abstract We consider a model of a fishery in which the dynamics of the unharvested fish population are given by the stochastic logistic growth equation Similar to the classical deterministic analogon, we assume that the fishery harvests the fish population following a constant effort strategy. In the first step, we derive the effort level that leads to maximum expected sustainable yield, which is understood as the expectation of the equilibrium distribution of the stochastic dynamics. This replaces the nonzero fixed point in the classical deterministic setup. In the second step, we assume that the fishery is risk averse and that there is a tradeoff between expected sustainable yield and uncertainty measured in terms of the variance of the equilibrium distribution. We derive the optimal constant effort harvesting strategy for this problem. In the final step, we consider an approach that we call the mean‐variance analysis to sustainable fisheries. Similar as in the now classical mean‐variance analysis in finance, going back to Markowitz [1952] , we study the problem of maximizing expected sustainable yields under variance constraints, and with this, minimizing the variance, e.g., risk, under guaranteed minimum expected sustainable yields. We derive explicit formulas for the optimal fishing effort in all four problems considered and study the effects of uncertainty, risk aversion, and mean reversion speed on fishing efforts.     

THE EFFECTS OF REDUCING MATING LIKELIHOOD ON POPULATION VIABILITY

Abstract The success a species may have invading a patch previously unoccupied is of considerable interest for pest managers and conservation ecologists. The purpose here is to present a mechanistic approach to analyze reproductive Allee effects appearing through the failure in the process of fertilization in a two‐sex population and observe how the survival in an invaded patch is affected. This is in contrast to the usually employed stochastic models with a deterministic skeleton that describe the presence of Allee effects. A Poisson–Ricker model, which includes stochastic demography and sex determination with females classified as successfully fertilized or not fertilized, is used. Numerical approximations to the probabilities of extinction and the mean time to extinction are presented, for fixed parameter values, suggesting how stochasticity in the mating process combined with random fluctuations in the male and female densities, at each generation, contribute to the risk of extinction of a population which started an invasion at a low density.     

PIGS,PARTIES, AND NOISE: HOW STOCHASTICITY IMPACTS THE ROBUSTNESS OF THE TSEMBAGA CULTURAL PRACTICES

Abstract We present a probabilistic perspective on sustainable resource usage. A mathematical model is introduced to describe the interplay between a population and its renewable resource base. The amount of effort a society chooses to exert in harvesting its resource is formalized in the model. Using an indigenous population of slash and burn farmers as a case study, we derive a system of stochastic differential equations from a system of ordinary differential equations introduced by another author. The cultural mechanisms that help to stabilize the population in the deterministic system actually decrease the expected survival time in the stochastic system.     

MODELS FOR COOPERATION AND CONSPIRACY IN FISHERIES: CHANGING THE RULES OF THE GAME

Fisheries regulation is considered necessary to counteract the effects of competitive forces which can lead to a “tragedy of the commons”. Yet management initiatives have often failed because they did not take into account competitive responses of fishing enterprises. In particular, open access fisheries provide strong incentives for the development of excessive harvesting capacity. This in turn leads to harvesting that is concentrated in space and time, with adverse effects on both the resource and markets. A coalition of fishermen, such as a fishermen's cooperative, has interests similar to those of a sole owner, and thus would be expected to produce more efficient behaviour. In practice, however, fishermen's cooperatives seldom persist. Game theory is used to explore relationships between the coalition structure of the industry, economic variables, and regulation. The models are based loosely on a purse seine fishery for herring. The results suggest that the potential to form stable coalitions is affected by changes in price and harvest. Changes in regulation also affect stability of coalitions. When interpreted in the light of historical changes in the herring fishery, these results suggest that industry may not accept regulations which do not permit formation of stable coalitions.     

FROM HARVESTING TO NONHARVESTING UTILITY: AN OPTIMAL CONTROL APPROACH TO SPECIES CONSERVATION

The purpose of this paper is to retrace the evolution of mathematical models focused on relation and interaction between economic growth, sustainable development, and natural environment conservation. First, generic defensive expenditures are introduced into a common‐property harvesting model in order to favor the species growth. Second, a transition model comprising both harvesting and nonharvesting values of wildlife biological species emerges. The latter gives rise to a group of purely nonharvesting models where anthropic activities and economic growth may have positive or negative impact on the natural evolution of wildlife species. Several scholars have proved that optimal strategies that are relatively good for harvesting purposes are not simply “transferrable” to the context of conservation of wildlife biological species with no harvesting value. In addition, the existence of optimal policies for long‐term conservation of all biological species (with or without harvesting value) cannot be guaranteed without having relatively large species populations at the initial time. Therefore, all such strategies are incapable of enhancing the scarce populations of endangered species and, therefore, cannot save these species from eventual (local) extinction. As an alternative, policymakers may soon be compelled to design and implement short‐term defensive actions aimed at recovery and enhancement of endangered wildlife species.     

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.     

Optimal and feedback strategies for managing multicohort populations

A Beverton and Holt type linear cohort dynamics model is integrated and combined with a nonlinear stock-recruitment relationship to obtain a discrete-time multicohort harvesting model. Assuming that each age class is individually controllable, it is shown, subject to certain assumptions, that the optimal harvesting strategy is to drive the population to the maximum sustainable yield solution in one time step. In most fisheries, this controllability assumption is not met and harvesting is agewise nonselective. In this case, it may be preferable to implement a harvesting policy based on suboptimal constant effort or stock level feedback strategies, rather than implement a more complicated optimal policy. This question is addressed through numerical studies on the management of an anchovy fishery.Dedicated to G. LeitmannThe author would like to thank M. Mangel, W. Reed, P. Sullivan, and G. Swartzman for commenting on a draft of this paper.     

POPULATION EXTINCTION IN DETERMINISTICAND STOCHASTIC DISCRETE‐TIME EPIDEMIC MODELS WITH PERIODIC COEFFICIENTS WITH APPLICATIONS TO AMPHIBIAN POPULATIONS

ABSTRACT. Discrete‐time deterministic and stochastic epidemic models are formulated for the spread of disease in a structured host population. The models have applications to a fungal pathogen affecting amphibian populations. The host population is structured according to two developmental stages, juveniles and adults. The juvenile stage is a post‐metamorphic, nonreproductive stage, whereas the adult stage is reproductive. Each developmental stage is further subdivided according to disease status, either susceptible or infected. There is no recovery from disease. Each year is divided into a fixed number of periods, the first period represents a time of births and the remaining time periods there are no births, only survival within a stage, transition to another stage or transmission of infection. Conditions are derived for population extinction and for local stability of the disease‐free equilibrium and the endemic equilibrium. It is shown that high transmission rates can destabilize the disease‐free equilibrium and low survival probabilities can lead to population extinction. Numerical simulations illustrate the dynamics of the deterministic and stochastic models.     

A PREDICTIVE MODEL TO INFORM ADAPTIVE MANAGEMENT OF DOUBLE‐CRESTED CORMORANTS AND FISHERIES IN MICHIGAN

The proliferation of double‐crested cormorants (DCCOs; Phalacrocorax auritus) in North America has raised concerns over their potential negative impacts on game, cultured and forage fishes, island and terrestrial resources, and other colonial water birds, leading to increased public demands to reduce their abundance. By combining fish surplus production and bird functional feeding response models, we developed a deterministic predictive model representing bird–fish interactions to inform an adaptive management process for the control of DCCOs in multiple colonies in Michigan. Comparisons of model predictions with observations of changes in DCCO numbers under management measures implemented from 2004 to 2012 suggested that our relatively simple model was able to accurately reconstruct past DCCO population dynamics. These comparisons helped discriminate among alternative parameterizations of demographic processes that were poorly known, especially site fidelity. Using sensitivity analysis, we also identified remaining critical uncertainties (mainly in the spatial distributions of fish vs. DCCO feeding areas) that can be used to prioritize future research and monitoring needs. Model forecasts suggested that continuation of existing control efforts would be sufficient to achieve long‐term DCCO control targets in Michigan and that DCCO control may be necessary to achieve management goals for some DCCO‐impacted fisheries in the state. Finally, our model can be extended by accounting for parametric or ecological uncertainty and including more complex assumptions on DCCO–fish interactions as part of the adaptive management process.     

An extended discrete Ricker population model with Allee effects

Based on the classical discrete Ricker population model, we incorporate Allee effects by assuming rectangular hyperbola, or Holling-II type functional form, for the birth or growth function and formulate an extended Ricker model. We explore the dynamics features of the extended Ricker model. We obtain domains of attraction for the trivial fixed point. We determine conditions for the existence and stability of positive fixed points and find regions where there exist no positive fixed points, two positive fixed points one of which is stable and two positive fixed points both of which are unstable. We demonstrate that the model exhibits period-doubling bifurcations and investigate the existence and stability of the cycles. We also confirm that Allee effects have stabilization effects, by different measures, through numerical simulations.     

Bifurcation analysis and chaos control in a discrete‐time plant quality and larch budmoth interaction model with Ricker equation

We investigate the dynamics of two‐dimensional discrete‐time model of leaf quality and larch budmoth interaction with Ricker equation. More precisely, the qualitative behavior of larch budmoth model is discussed in which the effect of food source upon the moth population is through intrinsic growth rate. We find the parametric conditions for local asymptotic stability of the unique positive fixed point. It is also proved that under certain parametric conditions, the system undergoes period‐doubling bifurcation with the help of center manifold theory. The parametric conditions for existence and direction of Neimark‐Sacker bifurcation at positive fixed point is investigated with the help of standard mathematical techniques of bifurcation theory. The chaos control in the system is discussed through implementation of hybrid control methodology. Finally, numerical simulations are provided to illustrate theoretical results. These results of numerical simulations demonstrate chaotic long‐term behavior over a broad range of parameters. The computation of the maximum Lyapunov exponents confirms the presence of chaotic behavior in the system.     

MULTISPECIES EXPLOITATION WITH EVOLUTIONARY SWITCHING OF HARVESTING STRATEGIES

In this paper, we propose a bioeconomic model which describes a fishery in which each of two noninteracting species is harvested by a given group of fishers during a defined time period. Then the Fishing Regulatory Authority allows each fisher to reconsider the harvesting decision at fixed (discrete) periods of time. The model derives from an Italian fisheries management experience in the Northern Adriatic Sea, where this kind of “self‐adjusting” fishing policy has been proposed to regulate harvesting of two shellfish species. The proposed dynamic model assumes the form of a hybrid system, as the natural growth functions of the two species (in continuous time) are coupled with a discrete time adaptive system that regulates how agents switch from one harvesting strategy to the other period by period according to an evolutionary mechanism based on profit comparison. In order to obtain some insights into the basic mechanisms of the system, some relevant benchmark cases are analyzed before tackling (mainly numerically) the complete hybrid model. Our results suggest that, for proper sets of parameters, this kind of myopic and adaptive self‐regulation may ensure a virtuous trade‐off between profit maximization and resource conservation, driven by cost externalities and market pressure.     

一类非线性随机种群动力学模型的最优收获控制

研究了一类非线性随机种群动力学模型的最优收获控制问题,得出了在外界环境对系统产生影响的条件下,最优控制所满足的必要条件及其最优性组,所得到的结论是确定性种群系统的扩展.     

A STRONG ERGODIC THEOREM FOR SOME NONLINEAR MATRIX MODELS FOR THE DYNAMICS OF STRUCTURED POPULATIONS

A general class of matrix difference equation models for the dynamics of discrete class structured populations in discrete time which possess a certain general type of nonlinearity introduced by Leslie for age-structured populations is considered. Arbitrary structuring is allowed in that transitions between any two classes are permitted. It is shown that normalized class distributions for such nonlinear models globally approach a “stable class distribution” and thus possess a strong ergodic property exactly like that of the classical linear theory of demography. However, unlike in the linear theory according to which the total population size grows or dies exponentially, the dynamics of total population size in these nonlinear models are shown to be governed by a nonlinear, nonautonomous scalar difference equation. This difference equation is asymptotically autonomous, and theorems which relate the dynamics of total population size to those of this limiting equation are proved. Examples in which the results are applied to some nonlinear age-structure models found in the literature are given.     

A STAGE‐STRUCTURED MODEL WITH TWO TIME‐STEPS FOR ANALYZING THE POPULATION DYNAMICS OF RHEA AMERICANA UNDER VARIABLE ENVIRONMENTAL CONDITIONS

ABSTRACT. The Ñandú or Rhea americana is an autochthonous species perfectly adapted to the pampas environment and only distributed in South America. The species exhibits an unusual breeding system combining polygyny, polyandry, communal nests and exclusive male parental care, which seems to contradict the idea of selfish genes. Our aim has been to construct a mathematical model based on the short term population dynamics of Rhea, living in the wild or in semi‐captivity, and taking into account environmental factors that vary from year to year. Due to the characteristics of its life cycle, it was necessary to develop a model that allows us to differentiate between the survival and fertility rates of each age group and the distinct behavior during breeding and non‐breeding seasons. Therefore, a quarterly differentiated stage‐structured discrete model was needed. Time steps of different lengths are used for modeling chicks or “charos' on the one hand, and juveniles and adults on the other. Environmental variables have been incorporated into the model because they affect the reproductive success of the species. Different scenarios are given as illustrations of the model use. Finally, the possibility of harvesting has been introduced in the model. The model is intended as a first step towards more refined models and systematic data gathering with the purpose of leading the way to a computational tool for risk assessment and decision‐making.     

USING RESERVES TO PROTECT FISH AND WILDLIFE SIMPLIFIED MODELING APPROACHES

ABSTRACT. This paper investigates theoretically to what extent a nature reserve may protect a uniformly distributed population of fish or wildlife against negative effects of harvesting. Two objectives of this protection are considered: avoidance of population extinction and maintenance of population, at or above a given precautionary population level. The pre‐reserve population is assumed to follow the logistic growth law and two models for post‐reserve population dynamics are formulated and discussed. For Model A by assumption the logistic growth law with a common carrying capacity is valid also for the post‐reserve population growth. In Model B, it is assumed that each sub‐population has its own carrying capacity proportionate to its distribution area. For both models, migration from the high‐density area to the low‐density area is proportional to the density difference. For both models there are two possible outcomes, either a unique globally stable equilibrium, or extinction. The latter may occur when the exploitation effort is above a threshold that is derived explicitly for both models. However, when the migration rate is less than the growth rate both models imply that the reserve can be chosen so that extinction cannot occur. For the opposite case, when migration is large compared to natural growth, a reserve as the only management tool cannot assure survival of the population, but the specific way it increases critical effort is discussed.