MULTISPECIES MODELING FOR ADAPTIVE MANAGEMENT OF HORSESHOE CRABS AND RED KNOTS IN THE DELAWARE BAY

Abstract Adaptive management requires that predictive models be explicit and transparent to improve decisions by comparing management actions, directing further research and monitoring, and facilitating learning. The rufa subspecies of red knots (Calidris canutus rufa), which has recently exhibited steep population declines, relies on horseshoe crab (Limulus polyphemus) eggs as their primary food source during stopover in Delaware Bay during spring migration. We present a model with two different parameterizations for use in the adaptive management of horseshoe crab harvests in the Delaware Bay that links red knot mass gain, annual survival, and fecundity to horseshoe crab dynamics. The models reflect prevailing hypotheses regarding ecological links between these two species. When reported crab harvest from 1998 to 2008 was applied, projections corresponded to the observed red knot population abundances depending on strengths of the demographic relationship between these species. We compared different simulated horseshoe crab harvest strategies to evaluate whether, given this model, horseshoe crab harvest management can affect red knot conservation and found that restricting harvest can benefit red knot populations. Our model is the first to explicitly and quantitatively link these two species and will be used within an adaptive management framework to manage the Delaware Bay system and learn more about the specific nature of the linkage between the two species.     

INDIVIDUAL‐BASED MODELS AND THE MANAGEMENT OF SHOREBIRD POPULATIONS

Abstract Individual‐based models (IBMs) predict how animal populations will be affected by changes in their environment by modeling the responses of fitness‐maximizing individuals to environmental change and by calculating how their aggregate responses change the average fitness of individuals and thus the demographic rates, and therefore size of the population. This paper describes how the need to develop a new approach to make such predictions was identified in the mid‐1970s following work done to predict the effect of building a freshwater reservoir on part of the intertidal feeding areas of the shorebirds Charadrii that overwinter on the Wash, a large embayment on the east coast of England. The paper describes how the approach was developed and tested over 20 years (1976–1995) on a population of European oystercatchers Haematopus ostralegus eating mussels Mytilus edulis on the Exe estuary in Devon, England. The paper goes on to describe how individual‐based modeling has been applied over the last 10 years to a wide range of environmental issues and to many species of shorebirds and wildfowl in a number of European countries. Although it took 20 years to develop the approach for 1 bird species on 1 estuary, ways have been found by which it can now be applied quite rapidly to a wide range of species, at spatial scales ranging from 1 estuary to the whole continent of Europe. This can now be done within the time period typically allotted to environmental impact assessments involving coastal bird populations in Europe. The models are being used routinely to predict the impact on the fitness of coastal shorebirds and wildfowl of habitat loss from (i) development, such as building a port over intertidal flats; (ii) disturbance from people, raptors, and aircraft; (iii) harvesting shellfish; and (iv) climate change and any associated rise in sea level. The model has also been used to evaluate the probable effectiveness of mitigation measures aimed at ameliorating the impact of such environmental changes on the birds. The first steps are now being taken to extend the approach to diving sea ducks and farmland birds during the nonbreeding season. The models have been successful in predicting the observed behavior and mortality rates in winter of shorebirds on a number of European estuaries, and some of the most important of these tests are described. These successful tests of model predictions raise confidence that the model can be used to advise policy makers concerned with the management of the coast and its important bird populations.     

CONSERVATION OF WILDLIFE. A BIO‐ECONOMIC MODEL OF A WILDLIFE RESERVE UNDER THE PRESSURE OF HABITAT DESTRUCTION AND HARVESTING OUTSIDE THE RESERVE

ABSTRACT. Biodiversity is today threatened by many factors of which destruction and reduction of habitats are considered most important for terrestrial species. One way to counteract these threats is to establish reserves with restrictions on land use and exploitation. However, very few reserves can be considered islands, wildlife species roam over large expanses, often via some density dependent dispersal process. As a consequence, habitat destruction, and exploitation, taking place outside will influence the species abundance inside the conservation area. The paper presents a theoretical model for analyzing this type of management problem. The model presented allows for both the common symmetric dispersal as well as what is called asymmetric dispersal between reserve and outside area. The main finding is that habitat destruction outside may not necessarily have negative impact upon the species abundance in the reserve. As a consequence, economic forces working in the direction of reducing the surrounding habitat have unclear effects on the species abundance within the protected area. We also find that harvesting outside the reserve may have quite modest effect on the species abundance in the reserve. This underlines the attractiveness of reserves from a conservation viewpoint.     

POSSIBILITIES AND LIMITATIONS OF USING HISTORIC PROVENANCE TESTS TO INFER FOREST SPECIES GROWTH RESPONSES TO CLIMATE CHANGE

Abstract. Under projected changes in global climate, the growth and survival of existing forests will depend on their ability to adjust physiologically in response to environmental change. Quantifying their capacity to adjust and whether the response is species‐ or population‐specific is important to guide forest management strategies. New analyses of historic provenance tests data are yielding relevant insights about these responses. Yet, differences between the objectives used to design the experiments and current objectives impose limitations to what can be learned from them. Our objectives are (i) to discuss the possibilities and limitations of using such data to quantify growth responses to changes in climate and (ii) to present a modeling approach that creates a species‐ and population‐specific model. We illustrate the modeling approach for Larix occidentalis Nutt. We conclude that the reanalysis of historic provenance tests data can lead to the identification of species that have population‐specific growth responses to changes in climate, provide estimates of optimum transfer distance for populations and species, and provide estimates of growth changes under different climate change scenarios. Using mixed‐effects modeling techniques is a sound statistical approach to overcome some of the limitations of the data.     

MODELING AND THE MANAGEMENT OF MIGRATORY BIRDS

Mathematical modeling of migratory bird populations is reviewed in the context of migratory bird management. We focus on dynamic models of waterfowl, since most management-oriented migratory bird models concern waterfowl species. We describe the management context for these modeling efforts, with a focus on large-scale operational data collection programs and on processes by which waterfowl harvest is regulated and waterfowl habitats are protected and managed. Through their impacts on key population parameters such as recruitment and survival rate, these activities can influence population dynamics, thereby providing managers some measure of control over the status of populations. Recent applications of the modeling of waterfowl are described in terms of objectives, mathematical structures, and contributions to management. Finally, we discuss research needs and data limitations in migratory bird modeling, and offer suggestions to increase the value to managers of future modeling efforts.     

EFFECTS OF INDIVIDUAL HETEROGENEITY IN ESTIMATING THE PERSISTENCE OF SMALL POPULATIONS

ABSTRACT. Population viability models are commonly used to estimate the probability of persistence of small, threatened, or endangered populations. Demographic, temporal, spatial, and individual heterogeneity are important factors affecting the probability of persistence of small populations. Because stochastic process are intractable analytically (Lud-wig [1996]), computer simulation models are often used for estimating population viability via numerical techniques. Although demographic, spatial, and temporal stochasticity have been incorporated into some population viability models, individual heterogeneity has not been included. In this paper we include individual heterogeneity in a simulation model and examine probabilities of population persistence at different levels of heterogeneity and population size. Individual heterogeneity may increase the probability of persistence of small populations. The mechanism for the extension in persistence may be explained by natural selection. Genotypes persisting through a decline may be those that survive better under the conditions causing the decline. These individuals that survive and reproduce in the face of adverse conditions may extend the probability that a small population persists.     

MODELING THE EFFECTS OF PREDATOR EXCLOSURES ON A WESTERN SNOWY PLOVER POPULATION

Abstract In Humboldt County, California, nest exclosures were used between 2001 and 2006 to reduce the predation of eggs aiding in the recovery of a threatened population segment of western snowy plovers (Charadrius nivosus). Due to a sudden increase in adult mortality in 2006, field biologists abandoned the use of nest exclosures. This paper describes a discrete‐time stochastic model designed to compare two different management strategies (with and without exclosures) to predict the change in the plover population. The model uses beta distributions to model demographic parameters, and whenever possible, these distributions were fit to survey data of these populations. The model shows nest exclosures to be effective in increasing chick fledging rates. However, the model also shows that an increase in adult mortality potentially caused by the nest exclosures would counteract this increase in fledging rates. The model predicts that there will be a net negative effect on the population if these exclosures reduce adult survival to 90% of its unexclosed rate. The model also demonstrates that the population is dependent on immigration.     

THE ECONOMICS OF MARINE RESERVES

ABSTRACT. Marine reserves can be a useful supplement to other methods of fisheries management, but marine reserves alone are not likely to achieve a great deal in economic terms andperhaps not even in terms of conservation. The effects of marine reserves with open access elsewhere are analyzed, using a logistic model for a population with a patchy distribution. It is assumedthat a marine reserve is establishedfor the territory of one of two sub‐populations which interact through migrations. The total population increases while the total catch declines for the most part. A high rate of migration would, however, dilute the conservation effect. Examining a stochastic variant of the model shows that the variability (sum of squareddeviations) of catches may decrease as a result of protecting one of the sub‐populations. Even if all rents disappear by assumption, it is possible to identify this as an economic benefit, particularly when the average catch increases. The variability of the catch falls for a range of values of the population migration parameter and variability of growth, both when the stochastic disturbances are independent and when they are perfectly correlated for the two sub‐populations, andalso when the growth variability parameter differs between the sub‐populations.     

BENEFITS AND RISKS OF INCREASED SPATIAL RESOLUTION IN THE MANAGEMENT OF FISHERY METAPOPULATIONS UNDER UNCERTAINTY

Abstract Stock assessments and harvest guidelines are typically based on the concept of a “fish stock,” which may encompass a very large area. The presence of discrete subpopulations within managed fish stocks presents risks and opportunities for fishery management. Failure to manage catch at the same scale as the true population structure can lead to extirpation of discrete subpopulations and to declines in the productivity of the larger metapopulation. However, it may be difficult and costly to assess and manage stocks at a finer spatial scale, and there is likely greater uncertainty about the size of substocks than about the aggregate stock. We use a two‐area simulation model to compare the performance of fishery management at different spatial resolutions when there is uncertainty about growth, the size of the total population, and the relative size of the subpopulations. We show that relative benefits of finer scale management, in terms of profits and risks of depleting subpopulations, depend on a number of biological, technical, and economic factors. In some cases it may be both less risky and more profitable to manage the fishery with a single total allowable catch, even when there are biologically separate fish populations in the two areas.     

CONSERVATION OF ENDANGERED ENDEMIC SEABIRDS WITHIN A MULTI‐PREDATOR CONTEXT: THE BARAU'S PETREL IN RÉUNION ISLAND

Abstract Seabirds breeding on islands are vulnerable to introduced predators, such as rats and cats, and the removal of such predators is generally viewed as a priority for seabird conservation and restoration. However, multiple invasive mammal species interacting may generate unexpected outcomes following the removal (eradication) of one species. Generally these indirect interactions are not well understood or demonstrated. We propose and study a prey (seabird)‐mesopredator (rat)‐superpredator (cat) model, taking into account the juvenile stages in the prey population, in order to direct conservation management for seabird conservation. We give a more biologically realistic differential system than those studied before (Courchamp et al. [1999] ; Fan et al. [2005] ), in particular for long‐lived seabird species. We present a theoretical study and show existence and uniqueness of a positive solution as well as a qualitative study of the equilibria that may appear. Because standard numerical methods, usually implemented in scientific softwares, can fail to give the right biological approximations ( Anguelov et al. [2009] ), we propose a reliable algorithm that preserves most of the qualitative properties of the continuous system, using the theory of nonstandard finite difference methods. Finally, we use biologically realistic parameters available for the representative Barau's petrel (Pinet et al. [2008] ), an endemic species from Réunion island, to present numerical simulations that support the theoretical study. Cats play the major role in seabird prey population dynamics. Seasonality in seabird breeding delays but does not prevent extinction. In all scenarios, cat control (or preferably eradication) is imperative to prevent extinction of vulnerable long‐lived seabirds, like the Barau's petrel.     

A FULL ANNUAL CYCLE MODELING FRAMEWORK FOR AMERICAN BLACK DUCKS

American black ducks (Anas rubripes) are a harvested, international migratory waterfowl species in eastern North America. Despite an extended period of restrictive harvest regulations, the black duck population is still below the population goal identified in the North American Waterfowl Management Plan (NAWMP). It has been hypothesized that density‐dependent factors restrict population growth in the black duck population and that habitat management (increases, improvements, etc.) may be a key component of growing black duck populations and reaching the prescribed NAWMP population goal. Using banding data from 1951 to 2011 and breeding population survey data from 1990 to 2014, we developed a full annual cycle population model for the American black duck. This model uses the seven management units as set by the Black Duck Joint Venture, allows movement into and out of each unit during each season, and models survival and fecundity for each region separately. We compare model population trajectories with observed population data and abundance estimates from the breeding season counts to show the accuracy of this full annual cycle model. With this model, we then show how to simulate the effects of habitat management on the continental black duck population.     

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.     

MANAGING HARVESTING TO MINIMIZE THE IMPACT OF EPIDEMICS ON WILD FISH STOCKS

ABSTRACT. Epidemic diseases inflict substantial damage to stocks of harvested species. Epidemic waves can be predictable away from their origin. I use a classical epidemiology model to investigate the interaction of harvesting strategy with an epidemic. The effect of reducing populations by harvesting before the epidemic depends upon the nature of the epidemic's survivors. If these have recovered following infection, then pre‐epidemic fishing optimizes the harvest, but reduces long‐term survival. However, if these survivors avoided infection, then increased pre‐epidemic fishing effort can increase post‐epidemic populations; survival is maximized by reducing the pre‐epidemic population to the threshold required to propagate infection. Post‐epidemic harvesting provides poor returns and damages stocks. Optimal stock management strategy in the face of a predicted epidemic depends upon balancing harvesting and conservation of stocks complimentary or antagonistic goals, depending on the nature of the epidemic.     

A HABITAT BASED MODEL FOR THE DISTRIBUTION OF FOREST INTERIOR NESTING BIRDS IN A FRAGMENTED LANDSCAPE

ABSTRACT. Increased awareness of the plight of many forest dwelling species has made necessary the development of methods for projecting the spatial distribution of these populations. This is particularly important for populations that currently occupy forest fragments and that are likely to be exposed to further disruption of their natural habitat. In this paper we develop a model for predicting the distribution of a bird population that evolved as forest interior dwellers. This model uses as its basis knowledge of the relationship between demographic characteristics of the population and the qualities of the habitat where individuals reside. We make the assumption that individuals will be naturally drawn to areas where they might expect greater reproductive success and repelled from areas where there is a high degree of intraspecific competition (high density). We apply the model to the ovenbird population in a large region of the Midwest. We use the model to examine the relative extent to which the surplus production from two major source areas supports extensive sink populations. The basic diffusion model parameterized by county forest cover data projects a population distribution which compares favorably with the results from the breeding bird count.     

A MODEL OF THE ENZOOTIOLOGY OF LYME DISEASE IN THE ATLANTIC NORTHEAST OF THE UNITED STATES

A mathematical model is presented for the dynamics of the rate of infection of the Lyme disease vector tick Ixodes dammini (Acari: Ixodidae) by the spirochete Borrelia burgdorferi, in the Atlantic Northeast of the United States. According to this model, moderate reductions in the abundance of white-tailed deer Odocoileus virginianus may either decrease or increase the spirochete infection rate in ticks, provided the deer are not reservoir hosts for Lyme disease. Expressions for the basic reproductive rate of the disease are computed analytically for special cases, and it is shown that as the basic reproductive rate increases, a proportional reduction in the tick population produces a smaller proportional reduction in the infection rate, so that vector control is less effective far above the threshold. The model also shows that control of the mouse reservoir hosts Peromyscus leucopus could reduce the infection rate if the survivorship of juvenile stages of ticks were reduced as a consequence. If the survivorship of juvenile stages does not decline as the rodent population is reduced, then rodent reduction can increase the spirochete infection rate in the ticks.     

CONSIDERATION OF STOCHASTIC DEMOGRAPHY IN THE DESIGN AND MANAGEMENT OF BIOLOGICAL RESERVES

A generalized birth–and–death process serves as a simple, flexible model for computing the expected persistence time of a small population in a random world. We may reparametrize the model in ways that allow explicit incorporation of density dependence, random differences in events experienced by different individuals, and random environmental variation experienced by all individuals in concert. This model seems to capture the important features of real population dynamics for purposes of computing the mean persistence time, even though the underlying mechanisms presumed in the mathematics of the model are decidedly unrealistic. The lack of isomorphism between birth and death rates, as they feature in the model, and vital rates of real biological populations can lead to extremely misleading results, if the classic formulation, rather than the reparametrization is applied without due circumspection. Using the reparametrized model, we find that environmental variation poses a greater problem for population persistence than does individual variation. In particular, with purely individual variation, the expected persistence time increases approximately with the power of the ceiling on population size; but with purely environmental variation, the expected persistence time increases somewhat less than linearly with the size of the population ceiling. The birth–and–death process model can also be applied to calculating the persistence time of a population on an ensemble of reserves which are linked by natural migration or by deliberate reintroduction programs. Results of this model, for an idealized ensemble, show that multiple independent reserves with a sufficient recolonization rate (natural or otherwise) will confer a longer persistence time than a single reserve with the same total carrying capacity, but in the absence of recolonization, the system of smaller separate reserves confers a shorter persistence time than the single large reserve.     

THE TRANSFORMATION OF LANDSCAPE: MODELING POLICY AND SOCIAL IMPACTS ON THE AGRICULTURAL LANDSCAPE OF LESVOS

ABSTRACT. The term landscape is more and more used as an “umbrella‘ concept, covering a series of cultural, productive and ecological processes. In order to uncover mechanisms, monitor transformations and predict changes, a complicated set of interacting factors has to be taken into account. This paper presents a model for estimating social and policy impacts on agricultural landscapes, based on the assumption that agricultural landscapes are shaped at” macro “(landscape) level by” micro “interventions at farm level. The model consists of three parts: an” ecological processes “part, which deals with processes that shape the ecological and aesthetic value of a landscape, a” population dynamics “part, which examines farmer population dynamics and a” policy impact “part, which deals with direct or indirect impacts on farming systems and farmer dynamics and refers to CAP Rural Development Measures. The model is applied for the olive and graze land agricultural landscapes of Lesvos (Greece). Results, apart from revealing landscape change patterns; help to illustrate some mechanisms behind this change and indicate that Rural Development Measures are inherent with minor but important malfunctions that cannot lead to sustainable landscape management and rural development in the area.     

MODELING THE SPATIAL DISTRIBUTION OF THE ECONOMIC COSTS AND BENEFITS OF ILLEGAL GAME MEAT HUNTING IN THE SERENGETI

ABSTRACT. Illegal game meat hunting in the Serengeti National Park, Tanzania, and adjacent game reserves provides an important source of protein and cash income to local communities. We construct a profitability model that describes the spatial distribution of the economic costs and benefits of illegal hunting in the Serengeti during the late 1980s and early 1990s. Costs included capital investment in hunting weapons, W_R, and the opportunity cost of hunting, W_O, both held to be constants; and two spatially variable components, the logistic effort of traveling to hunting areas, W_L, and the penalties incurred if arrested, W_P. Benefit was the expected income from the sale of meat from resident wildlife species. The model suggests: (1) W_R is the most important cost. (2) W_L is the second most important cost and likely to determine the spatial distribution of hunting activity if hunters seekto minimize costs. (3) W_O and W_P are of minor importance, the former because alternative sources of income provide low pay, the latter because the overall chance of being arrested is low. (4) W_P exceeds W_L only in areas close to the boundary of protected areas. (5) Although resident wildlife contributes only a minor share of illegal offtake compared to the migratory herds, hunting resident wildlife is profitable in 68% of the area. This suggests that hunting of resident and migratory wildlife is highly profitable and may explain why the utilization of the target populations has become increasingly unsustainable.     

RUMINATIONS ON THE DEVELOPMENT AND FUTURE OF POPULATION DYNAMICS MODELS IN FISHERIES

ABSTRACT. I trace the development of fisheries models (i.e., fish population dynamics models of species subject to fisheries) to the 21st century. The first real efforts occurred in the period 1900 1920 with the work of Baranov (the “Grandfather” of fisheries population dynamics) and the formation of the International Council for the Exploration of the Sea (ICES). The establishment of the science occurred between 1920 1960 with multi‐species modeling, age‐ and size‐structure dynamics, and production models. Fundamental work during this time was done by Ricker (the “Father” of fisheries population dynamics), Beverton and Holt (the “Prophets” of fisheries population dynamics), Chapman, Dickie, DeLury, Graham, Gulland, Leslie, Lotka and Volterra, Russell, Schaefer, and Thompson. During this time, most of the workwas deterministic and mathematical. Between 1960 and 1980, statistical methodology evolved greatly but was separate from mathematical advances for the most part. The development of statistical principles for the estimation of animal abundance was further enhanced by Arnason, Buckland, Burnham and Anderson and White, Cormack, Eberhardt, Jolly, Manly, Pollock, Ricker, Robson, and Seber, among others. Fisheries models evolved in a deterministic setting, with advances in age‐structured models (Gulland, Pope, Doubleday), surplus production models (Pella, Tomlin‐son, Schnute, Fletcher, Hilborn), growth models, bioeconomic models (C. Clark) and management control models (Hilborn, Walters). The period 1980 2000 was the Golden Age. The integration between mathematics and statistics occurred when likelihood and least squares techniques were formally combined with mathematical models of population change. The number of fisheries modelers grew exponentially during this time, resulting in a concomitant increase in publications. A major advance in the 1990s has been the development of Bayesian and time series methods, which have allowed explicit specification of uncertainty. Currently, theory allows realistic modeling of age‐ and size‐structured populations, migratory populations and harvesting strategies. These models routinely incorporate measurement error, process error (stochasticity) and time variation. But data needs often overwhelm the performance of models, and greater demands are being placed on models to answer complex questions. There has been poor communication between fisheries and ecological modelers, between fisheries researchers and statisticians, and among fisheries researchers in different geographic locales. Future models will need to deal better with habitat and spatial concerns, genetics, multispecies interactions, environmental factors, effects of harvesting on the ecosystem, model misspecification and so‐cioeconomic concerns. Meta‐analysis, retrospective analysis and operating models are some modern approaches for dealing with uncertainty and providing for sustainable fisheries. However, I fear that current attacks on single‐species models and management may result in rejection of these advances and an attempt to substitute a less scientific approach.