The application of species criteria in avian taxonomy and its implications for the debate over species concepts.

The debate over species concepts has produced a huge body of literature on how species can, may or should be delimited. By contrast, very few studies have documented how species taxa are delimited in practice. The aims of the present study were to (i) quantify the use of species criteria in taxonomy, (ii) discuss its implications for the debate over species concepts and (iii) assess recent claims about the impact of different species concepts on taxonomic stability and the 'nature' of species. The application of six species criteria was examined in taxonomic studies of birds published between 1950 and 2009. Three types of taxonomic studies were included: descriptions of new species (N = 329), proposals to change the taxonomic rank of species and subspecies (N = 808) and the taxonomic recommendations of the American Ornithologists' Union Committee on Classification and Nomenclature (N = 176). In all three datasets, diagnosability was the most frequently applied criterion, followed by reproductive isolation and degree of difference. This result is inconsistent with the popular notion that the Biological Species Concept is the dominant species concept in avian taxonomy. Since the 1950s, avian species-level taxonomy has become increasingly pluralistic and eclectic. This suggests that taxonomists consider different criteria as complementary rather than as rival approaches to species delimitation. Application of diagnosability more frequently led to the elevation of subspecies to species rank than application of reproductive isolation, although the difference was small. Hypotheses based on diagnosability and reproductive isolation were equally likely to be accepted in a mainstream checklist. These findings contradict recent claims that application of the Phylogenetic Species Concept causes instability and that broader application of the Biological Species Concept can stabilise taxonomy. The criteria diagnosability and monophyly, which are commonly associated with Phylogenetic Species Concepts, were used throughout the study period. Finally, no support was found for the idea that Phylogenetic Species Concepts have caused a change in the 'nature' of species taxa. This study demonstrates that there is a discrepancy between widely held perceptions of how species are delimited and the way species are actually delimited by taxonomists. Theoretically oriented debates over species concepts thus may benefit from empirical data on taxonomic practice.