Fluctuating asymmetry as an animal welfare indicator -- a review of methodology and validity.

It has been suggested that fluctuating asymmetry (FA) reflects an animal's ability to cope with the sum of challenges during its growing period and, thus, is a potential welfare indicator. In this review we investigate the evidence of associations between FA and other welfare indicators measured at the level of the individual and of effects of welfare-relevant environmental conditions on FA in populations of captive birds and mammals including humans. As the question of validity cannot be treated independently from the quality of the available data, first a checklist for the proper measurement and analysis of FA is drafted and used to evaluate the methodological quality of the various studies. We recommend this checklist to be used as a standard for future FA studies. We found 17 relevant studies on associations between FA and other welfare indicators, and 36 studies on effects of welfare-relevant factors on FA. Frequent methodological shortcomings or insufficient methodological information allow for only cautious conclusions. The proportion of significant results supporting the link between higher FA and poorer welfare is only moderately high. Independent from statistical significance, almost all studies found the relationship between FA and welfare to be prevailingly in the expected direction. FA is a promising measure of animal welfare, despite a great number of open questions, e.g. relating to the ontogeny of FA or its sensitivity to various stressors. The considerable potential of FA as a welfare indicator makes it worthwhile to pursue more intensely validation studies as well as applied studies. These studies should pay particular attention to an appropriate methodological approach.