Directional asymmetry of long-distance dispersal and colonization could mislead reconstructions of biogeography

Aim: Phylogenies are increasingly being used to attempt to answer biogeographical questions. However, a reliance on tree topology alone has emerged without consideration of earth processes or the biology of the organisms in question. Most ancestral-state optimization methods have inherent problems, including failure to take account of asymmetry, such as unequal probabilities of losses and gains, and the lack of use of independent cost estimates. Here we discuss what we perceive as shortcomings in most current tree-based biogeography interpretation methods and show that consideration of processes and their likelihoods can turn the conventional biogeographical interpretation on its head. Location: Southern hemisphere focus but applicable world-wide. Methods: The logic of existing methods is reviewed with respect to their adequacy in modelling processes such as geographical mode of speciation and likelihood of dispersal, including directional bias. Published reconstructions of dispersal of three plant taxa between Australia and New Zealand were re-analysed using standard parsimony and maximum likelihood (ML) methods with rate matrices to model expected asymmetry of dispersal. Results: Few studies to date incorporate asymmetric dispersal rate matrices or question the simplistic assumption of equal costs. Even when they do, cost matrices typically are not derived independently of tree topology. Asymmetrical dispersal between Australia and New Zealand could be reconstructed using parsimony but not with ML. Main conclusions: The inadequacy of current models has important consequences for our interpretation of southern hemisphere biogeography, particularly in relation to dispersal. For example, if repeated directional dispersals and colonization in the direction of prevailing winds have occurred, with intervening periods of speciation, then there is no need to infer dispersals against those winds. Failure to take account of directionality and other biases in reconstruction methods has implications beyond the simple misinterpretation of the biogeography of a taxonomic group, such as calibration of molecular clocks, the dating of vicariance events, and the prioritization of areas for conservation.