MORPHOLOGICAL EVOLUTION MEDIATED BY BEHAVIOR IN THE DAMSELFLIES OF TWO COMMUNITIES.

Behavior can play a mediating role in determining the selective pressures that influence the evolution of morphological structures. To examine this, I quantified patterns of morphological variation among larvae of Enallagma damselfly species (Odonata: Coenagrionidae) that use different behaviors to avoid the major predators found in each of two communities, lakes with and without fish. Specifically, I quantified the sizes and shapes of the abdomens and caudal lamellae (used for swimming) and legs for three species from fishless lakes and six species from lakes with fish. A preliminary cladistic analysis indicates that species within each lake type are not members of a single clade, which supports the conclusions of previous odonate taxonomists. Previous studies have shown that species in fishless lakes are very active, running and swimming frequently and at high rates of speed in the absence of predators, and they avoid their primary predators, large dragonflies, by swimming. These species have the widest abdomens, the largest caudal lamellae relative to overall body size, and the longest legs of the species studied, which should make them powerful swimmers and runners. Furthermore, species in fishless lakes are morphologically very similar to one another and differ greatly from fish-lake species, although each is more closely related to species in fish lakes. In contrast, species from lakes with fish move very slowly and infrequently in the absence of predators and do not attempt to evade attacking predators. However, despite their behavioral similarity, large interspecific variation in morphology exists among the fish-lake species, and the only morphological patterns were differences associated with membership in the two primary clades identified in the cladistic analysis. A modification of Felsenstein's (1985) method of evolutionary contrasts which allows character change to be isolated along single branches is introduced and is used to reconstruct the evolutionary histories of these characters. This analysis suggests that large increases in caudal lamella size, abdominal segment lengths and widths, and leg length accompany speciation events associated with habitat shifts from fish-lakes to fishless lakes. Following habitat shifts selection pressures exerted by dragonfly predation apparently favored swimming as an escape tactic, which mediated selection pressures onto morphologies used in swimming to increase swimming performance; morphological patterns in extant species reflect this adaptation to a new environment. Mechanisms by which behaviorally mediated selection could have accelerated evolutionary dynamics following founder events are discussed.