On the combined analysis of proximate and ultimate aspects in diel vertical migration (DVM) research

Although evolutionary ecologists agree that proximate and ultimate aspects are two sides of one coin, they are seldom interested in studies on physiological and behavioural mechanisms at the base of ecological phenomena. Nevertheless, these mechanisms are objects of selection and evolved to realise adaptive significances. This paper is a plea to bring both fields closer together, and, by means of an example of Diel Vertical Migration of Daphnia, some proximate and ultimate aspects are discussed. It is argued that light changes, not fish kairomone, is the primary cause for an individual to swim downwards at dawn and upwards at dusk. However, what is called a causal factor might differ when ecosystems or individuals are studied. In addition, causality in ecology is not simple, and has the character of a `set of necessary conditions'. To illustrate the importance of proximate analyses in DVM, two basic response mechanisms are discussed: Photobehaviour system 1 and 2. The physiological character of these systems leads to a fixed type of migration or to a phenotypically induced DVM, respectively. The adaptive significance of the first might be a reduction of the hazardous effects of UV radiation and of the second a lowering of mortality due to visually hunting predators.