Using pattern-oriented modeling for revealing hidden information: a key for reconciling ecological theory and application

We suggest that the conscious use of information that is "hidden" in distinct structures in nature itself and in data extracted from nature ( = pattern) during the process of modeling ( = pattern-oriented modeling) can substantially improve models in ecological application and conservation. Observed patterns, such as time-series patterns and spatial patterns of presence/absence in habitat patches, contain a great deal of data on scales, site-history, parameters and processes. Use of these data provides criteria for aggregating the biological information in the model, relates the model explicitly to the relevant scales of the system, facilitates the use of helpful techniques of indirect parameter estimation with independent data, and helps detect underlying ecological processes. Additionally, pattern-oriented models produce comparative predictions that can be tested in the field. We developed a step-by-step protocol for pattern-oriented modeling and illustrate the potential of this protocol by discussing three pattern-oriented population models: (1) a population viability analysis for brown bears (Ursus arctos) in northern Spain using time-series data on females with cubs of the year to adjust unknown model parameters; (2) a savanna model for detecting underlying ecological processes from spatial patterns of tree distribution; and (3) the incidence function model of metapopulation dynamics as an example of process integration and model generalization. We conclude that using the pattern-oriented approach to its full potential will require a major paradigm shift in the strategies of modeling and data collection, and we argue that more emphasis must be placed on observing and documenting relevant patterns in addition to attempts to obtain direct estimates of model parameters.