Species distribution models and ecological theory: A critical assessment and some possible new approaches

Given the importance of knowledge of species distribution for conservation and climate change management, continuous and progressive evaluation of the statistical models predicting species distributions is necessary. Current models are evaluated in terms of ecological theory used, the data model accepted and the statistical methods applied. Focus is restricted to Generalised Linear Models (GLM) and Generalised Additive Models (GAM). Certain currently unused regression methods are reviewed for their possible application to species modelling.A review of recent papers suggests that ecological theory is rarely explicitly considered. Current theory and results support species responses to environmental variables to be unimodal and often skewed though process-based theory is often lacking. Many studies fail to test for unimodal or skewed responses and straight-line relationships are often fitted without justification.Data resolution (size of sampling unit) determines the nature of the environmental niche models that can be fitted. A synthesis of differing ecophysiological ideas and the use of biophysical processes models could improve the selection of predictor variables. A better conceptual framework is needed for selecting variables.Comparison of statistical methods is difficult. Predictive success is insufficient and a test of ecological realism is also needed. Evaluation of methods needs artificial data, as there is no knowledge about the true relationships between variables for field data. However, use of artificial data is limited by lack of comprehensive theory.Three potentially new methods are reviewed. Quantile regression (QR) has potential and a strong theoretical justification in Liebig's law of the minimum. Structural equation modelling (SEM) has an appealing conceptual framework for testing causality but has problems with curvilinear relationships. Geographically weighted regression (GWR) intended to examine spatial non-stationarity of ecological processes requires further evaluation before being used.Synthesis and applications: explicit theory needs to be incorporated into species response models used in conservation. For example, testing for unimodal skewed responses should be a routine procedure. Clear statements of the ecological theory used, the nature of the data model and sufficient details of the statistical method are needed for current models to be evaluated. New statistical methods need to be evaluated for compatibility with ecological theory before use in applied ecology. Some recent work with artificial data suggests the combination of ecological knowledge and statistical skill is more important than the precise statistical method used. The potential exists for a synthesis of current species modelling approaches based on their differing ecological insights not their methodology.