This file was created by scanning the printed publication. Errors identified by the software have been corrected; however, some errors may remain. Effect of Uncertainty in Mapped Biodiversity Data on Optimal Conservation Decisions Michael J. Conroyl and Jennifer E. Crocker2 Abstract--Spatially referenced biological diversity data are increasingly being used to assist with conservation decisions. Examples include the identification, selection, and design of ecological reserves; the types, amount, and distribution of forest cutting; and the placement of corridors to connect reserves. These have in common: (1) an implicit long-term objective; (2) dynamic systems ; (3) alternative possible decisions ; (4) alternative uncertain outcomes; (5) uncertainty about both the current system state (sampling errors), and in predictions about future system states ("model uncertainty"). Decision-theoretic methods and predictive models can be used to find decisions that result in a long term optimum (e.g., a balance between species conservation and timber revenue), subject to updating through current surveys, research, and other information. We illustrate the approach for a hypothetical reserve design problem involving four species with different habitat affinities in a landscape with two habitat types, with both demographic and statistical uncertainty. In particular, we examine how uncertainty in predictive models (e.g., models relating animal distribution and abundance to habitat features) can result in suboptimal decisions, and how research and monitoring can be used adaptively to reduce uncertainty and improve decision making. INTRODUCTION Increasingly, managers are faced with making decisions about how to manage systems in which there are multiple, competing goals or objectives, and the system under management is both complex and incompletely understood. The failure of public decision makers to make a "good" decision can have severe consequences, ranging from loss of agency credibility and good will, to unnecessary losses of revenue or opportunity, to adverse impacts on the resource, including the loss of species. Concern over loss of biological diversity has led to efforts in the ' Assistant Leader, Georgia Cooperative Fish and Wildlrfe Research Unit, Athens, GA. Graduate research assistant, Institute of Ecologv, Universify of Georgia, Athens, GA 685 cartographic analyses of species distributions and landscape characteristics (e.g., Burley 1988; Scott et al. 1993) In these efforts, animal species richness is either observed directly or inferred from vegetation (habitat) maps and models relating abundance or presencelabsence to vegetation cover attributes and physical features (Scott et al. 1993). The mapped data are then used land-use decisions directed at the conservation of biological diversity, such as reserve design (Scott et al. 1993). Biodiversity mapping programs such as Gap Analysis frequently do not formally incorporate uncertainty about 1) mapping errors in habitat and animal species presence distributions, and 2) species-habitat associations. They implicitly assume that predictions can be made about the future state of biodiversity, given alterations in current habitat conditions (e.g., through management). Here we consider how statistical unreliability in spatially referenced data may affect the utility of mapped species richness patterns for conservation decisions such as reserve selection and design. We first briefly consider alternative approaches to obtaining optimal management decisions under the conditions in which the system is 1) stochastic, 2) dynamic, 3) imperfectly understood, and 4) incompletely observable. We illustrate the general problem by means of an artificial landscape containing 2 habitat types and 4 species (Conroy and Noon in press, Crocker and Conroy 1995). Before proceeding, we note that we are using one specific method (stochastic dynamic programming) for determining optimal policies. There are a variety of alternative analytical approaches to this problem, capably reviewed by Williams (1989). These include non-dynamic methods such as linear programming, simulation methods, and combinations of simulation and optimization. We recognize that no single approach is likely to perform adequately under all circumstances, but we strongly feel that any approach should adequately reflect both the dynamic nature of ecological systems and decision making, and the role of uncertainty in both observations and model structure. We also agree with Walters (1986) and Nichols et al. (1995) that a formally adaptive approach to natural resource management will in the long run prove most efficacious. METHODS Elements of the Problem We consider a hypothetical 100 hectare landscape with three cover types, which we have temed forest, open, and developed. Initially, the landscape is 75% forested, 25% open, and 0% developed. This landscape is initially occupied by four hypothetical species, two forest habitat specialists and two open habitat specialists. The management objective expresses a tradeoff between the conservation goal of maintaining the species in the landscape and the economic goal of allowing development. At five year intervals, the manager must decide what proportion of each habitat type to conserve, and a proportion of all unconserved land is permanently lost to development. The manager has a range of hypotheses regarding population response to habitat management. The population is monitored during the management period, both to guide management decisions and to reduce uncertainties about population dynamics. In the development below we freely borrow fiom Nichols et. al. (1995), whom we thank for their lucid description of the problem. Management options In this simplistic, non-spatial problem, the manager must decide how much of the available wildlife forest and open habitat to conserve for the next five year period. The action chosen in period t is denoted 4. A policy A specifies a sequence of management actions { a , a,, ..., a,} as a function of the observed population and habitat conditions. Models The models specify the dynamics of the state variables through time. The set of population models specifies a range of alternative hypotheses about the population responses to management actions. The population dynamics and environmental dynamics are combined in a single state dynamics equation, where x, is a vector of state variables including both population status and habitat status, z, is a white noise process, and F,(x,,a,,zJis the net change in the state variables under model i. We considered three models describing population response to management actions. These models differ in the degree to which population dynamics is related to the amount, composition, and spatial arrangement of habitat. Model 1 was a simple non-dynamic habitat relation model, N=dH, where N=species abundance, d=a constant density, H=the amount of habitat (forest or open, depending on the species). Model 2 was an equilibrium source-sink model (Pulliam 1988). For each species, where N=species abundance, H=amount of source habitat, d=maximum density in the source habitat, k,= population growth rate in the source habitat, and A, = population growth rate in the sink habitat. Model 3 was a dynamic source-sink model (Pulliam 1988). For each species, dynamics were described by: N(2)t+1= { if a (l)N(l), < dH if a (I)N(l), 2 dH N(2), + 1(1)N(i), - dH where N(l)=population in source, k(l)=population growth rate in source, d= maximum density in source, H= amount of source habitat, N(2)=population in sink, a(2) = population growth rate in sink. Values of the species parameters are given in Table 1. Table 1.--Model parameters for population dynamics. Habitat affinity Species 1 Forest Species 2 Forest Species 3 Open Species 4 Open AI A2 - - d - 4 More complicated, spatially-explicit models can and will be constructed, but for the present Models 2 and 3 allow dispersal from favorable to unfavorable habitats, and thus are 'spatially informed', whereas for Model 1, in which movement between habitats does not occur, it is only the amount of favorable habitat that determines abundance for each species. Landscape dynamics followed a simple state transition model. Unconserved land was permanently lost to development at a rate of 0.049 every 5 year period. Of the remaining habitat, forest converted to open at a 5 year rate of 0.26, and open to forest at a 5 year rate of 0.95. Components of uncertainty There are four components of uncertainty which are important in managing biological systems. Environmental stochasticity is uncontrollable variation in the factors influencing population dynamics. We modelled extinction as a stochastic process, with probability distributions conditioned on population size. Structural uncertainty reflects imperfect knowledge about the relationship between management actions and population status. We explored the effect of structural uncertainty by using a weighted average of the three model predictions in the optimization procedure, and varying the weights assigned to each model. Partial observability refers to the imprecision in a monitoring program. Such imprecision leads to uncertainty about the appropriate action to take, and also can affect the ability to discriminate among alternative models. We plan to model the effect of partial observability by developing an optimal policy based on complete observability, and then using that policy in a simulation model with random variation around the population state variables. Partial controllability refers to the inability to implement management decisions without error. We have not considered partial controllability in this exercise, but plan to do so in future work. Monitoring Since decisions are partially based on population status, a monitoring program is needed. The monitoring program can also serve to discriminate among the competing models. The data recorded at time t are denoted y,, and are related to the population according to with the random variable E independent of z. The set of all data collected up to time t is denoted Y,. Model weights The likelihood at time t that model i is the best model relating population to habitat is denoted p,(t). The likelihoods are expected to change through time as data are accumulated and compared with model predictions. Objective function Our objective function V is a weighted sum of a species conservation objective and a resource use objective. In practice, the weights would be determined using multi-attribute utility theory (Keeney and Raiffa 1976). Our objective is to maximize where V(AI Y, pJ is the expected value of the objective for policy A given the current state of knowledge, W,=0.1875 and W, = 0.25 are weights for the two objectives, S(A) is the number of species at the end of 100 years, and C is the proportion of the available habitat conserved over all time steps. The weights were chosen so that the objective function falls between 0 and 1. Solution method All optimizations and simulations were done using Stochastic Dynamic Programming (Lubow 1993), which uses backward iteration to find optimal policies for discrete Markov processes. Analysis Our analysis focused on the effect of structural uncertainty on the optimal policy and the expected value of the objective function. Seven combinations of weights pi were used (Table 2), reflecting a range of likelihoods ascribed to the three models. The first three cases reflected structural certainty. In the next three cases there is greater confidence in one model than the other two, and in the third there is no prior knowledge about the appropriate model. In the extreme case in which the data y, indicate with certainty which model is correct, we obtain perfect information (Lindley 1985) about the system structure. Given prior weights pi given to the models, the expected value of the objective function with perfect information is: The difference between this value and the maximum value of the objective function calculated using the weighted (by thep,) average of the three models is the expected value of perfect information (EVPI) (Lindley 1985). The EVPI was calculated for the four combinations of weights which reflected structural uncertainty. RESULTS Table 2.--The effect of varying model weights on optimal decisions and objective V PI P2 P3 V EVPI forest area conserved open area conserved In general, the population predictions for Model 2 were higher than those for Model 1. This was not surprising, since in both models density was proportional to the amount of preferred habitat, with larger constants of proportionality in Model 2. For all combinations of state variables, the expected utility for Model 2 was equal to or greater than the expected utility for Model 1. Similarly, expected utilities for Model 3 were generally equal to or greater than those for Model 2. As habitat was lost, Model 2 predicted immediate population equilibrium with the new conditions, while Model 3 predicted a time lag. The optimal decisions also varied among the models. In general, the optimal amount to conserve was larger for the model which predicted lower population size. Varying the likelihoods p, assigned to each model affected both the optimal decision and the value of the objective function for the optimal decision (Table 2). DISCUSSION The degree of confidence placed in each model affected both the optimal management decision and the expected value of the objective function. Basing management on an incorrect model without considering structural uncertainty can lead to suboptimal decisions. The EVPI refers to the expected increase in the objective function from a hypothetical study designed to eliminate structural uncertainty. For example, a gain in objective function units of 0.033 might translate into an increase in the expected number of species conserved of 0.033/W1= 0.176, or an increase in the proportion of land open for development of 0.033/W2 = 0.132. In a realistic management scenario, data will not discriminate among the models exactly. In that case, the expected gain from gathering the data depends on the model likelihoods (Lindley 1985). The expected value of partial information can be used in an analysis in the cost-benefit tradeoff of various levels of monitoring intensity. In this exercise, we chose the model weights arbitrarily. An adaptive approach to management would involve treating the model weights as state variables whose dynamics are based on Bayesian updating. In passive adaptive management (Walters and Hilborn 1978), information feedback into the decision making process is not considered in evaluating the objective function. Optimization, management, and monitoring are done iteratively, with model weights recalculated after each iteration. In active adaptive management, the optimization specifically considers the benefits of future learning. Active adaptive management presents considerable computational difficulties, because the information state Y,increases in dimension through time. A dynamic programming approach incorporating future learning must rely on a set of summary statistics which evolve in a Markovian fashion. This model exercise represents a preliminary step in developing adaptive approaches to conservation planning. Our immediate objectives include exploring the effect of partial observability on decision-making in our model landscape, and performing similar analyses using population models with spatial structure. This will require some refinements in our solution methods due to the dimensionality limitations of dynamic programming. We anticipate taking an iterative simulationoptimization (SO) approach to dealing with more complex systems. In SO, methods such as dynamic programming or linear programming is used to develop optimal policies for simplified or static systems. More detailed simulation models are then used to track the system dynamics over time using the optimal policy. If SO proves to be an effective strategy for dynamic optimization in complex systems, we will have the analytical tools available for addressing optimization that is truly adaptive, in the sense that the information state is included in the optimization procedure. ACKNOWLEDGMENTS We thank Ken Williams, Jim Nichols, Fred Johnson, and Bill Kendall for invaluable consultation and input. REFERENCES Burley, F.W. 1988. Monitoring biological diversity for setting conservation priorities. pp. 227-230 Wilson, E.O. (ed.) Biodiversity. Washington, DC :National Academy Press. Conroy, M.J. and B.R. Noon. 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