Predicting mortality in novel environments: Tests and sensitivity of a behaviour-based model

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Authors: Stillman, R.A., Goss-Custard, J.D., West, A.D., Dit Durell, S.E.A.L.V., Caldow, R.W.G., McGrorty, S. and Clarke, R.T.

Journal: Journal of Applied Ecology

Volume: 37

Issue: 4

Pages: 564-588

ISSN: 0021-8901

DOI: 10.1046/j.1365-2664.2000.00506.x

1. In order to assess the future impact of a proposed development or evaluate the cost effectiveness of proposed mitigating measures, ecologists must be able to provide accurate predictions under new environmental conditions. The difficulty with predicting to new circumstances is that often there is no way of knowing whether the empirical relationships upon which models are based will hold under the new conditions, and so predictions are of uncertain accuracy. 2. We present a model, based on the optimality approach of behavioural ecology, that is designed to overcome this problem. The model's central assumption is that each individual within a population always behaves in order to maximize its fitness. The model follows the optimal decisions of each individual within a population and predicts population mortality rate from the survival consequences of these decisions. Such behaviour-based models should provide a reliable means of predicting to new circumstances because, even if conditions change greatly, the basis of predictions - fitness maximization - will not. 3. The model was parameterized and tested for a shorebird, the oystercatcher Haematopus ostralegus. Development aimed to minimize the difference between predicted and observed overwinter starvation rates of juveniles, immatures and adults during the model calibration years of 1976-80. The model was tested by comparing its predicted starvation rates with the observed rates for another sample of years during 1980-91, when the oystercatcher population was larger than in the model calibration years. It predicted the observed density-dependent increase in mortality rate in these years, outside the conditions for which it was parameterized. 4. The predicted overwinter mortality rate was based on generally realistic behaviour of oystercatchers within the model population. The two submodels that predicted the interference-free intake rates and the numbers and densities of birds on the different mussel Mytilus edulis beds at low water did so with good precision. The model also predicted reasonably well (i) the stage of the winter at which the birds starved; (ii) the relative mass of birds using different feeding methods; (iii) the number of minutes birds spent feeding on mussels at low water during both the night and day; and (iv) the dates at which birds supplemented their low tide intake of mussels by also feeding on supplementary prey in fields while mussel beds were unavailable over the high water period. 5. A sensitivity analysis showed that the model's predictive ability depended on virtually all of its parameters. However, the importance of different parameters varied considerably. In particular, variation in gross energetic parameters had a greater influence on predictions than variations in behavioural parameters. In accord with this, much of the model's predictive power was retained when a detailed foraging submodel was replaced with a simple functional response relating intake rate to mussel biomass. The behavioural parameters were not irrelevant, however, as these were the basis of predictions. 6. Although we applied the model to oystercatchers, the general principle on which it is based applies widely. We list the key parameters that need to be measured in order to apply the model to other systems, estimate the time scales involved and describe the types of environmental changes that can be modelled. For example, in the case of estuaries, the model can be used to predict the impact of habitat loss, changes in the intensity or method of shellfishing, or changes in the frequency of human disturbance. 7. We conclude that behaviour-based models provide a good basis for predicting how demographic parameters, and thus population size, would be affected by novel environments. The key reason for this is that, by being based on optimal decision rules, animals in these models are likely to respond to environmental changes in the same way as real one s would.

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