Habitat Heterogeneity and Fish Behavior: Units of Heterogeneity as a Resource and as a Source of Information1

摘要

A review of studies, mainly experimental, on modifications of fish behavior caused by microscale habitat heterogeneity. Elements or units of heterogeneity influence on decision making in fish either as contestable physical resources, or as informationcues or signals. Habitat heterogeneity arises from abiotic physical objects, aggregations of prey, and grouping fish. Feeding behavior of fish including food search, choice, and consumption are significantly dependent on the structure of heterogeneity of the habitat, where fish are foraging. Depending on the parameters of heterogeneity, prey characteristics and a predator foraging mode, heterogeneous habitats can either facilitate feeding behavior, or makes it more difficult. Habitat heterogeneity plays significant and, as a rule, positive role providing various refuges for fish hiding from predators. Landmarks help fish to find the shortest route to shelters. If a habitat is rather homogeneous or in a novel habitat, which appears to be homogeneous, shoaling of fish makes surroundings of each individual in the school structured providing fish with a substitute of shelters and landmarks. Recent experimental and field results convincingly demonstrate that the effects of main biotic and abiotic factorscan be significantly modified by the structure (level of spatial heterogeneity) of habitats. When a habitat is physically structured, tendencies to disperse and establish individual territories prevail. In uniform, poorly structured habitats, fish tend to gather in schools or shoals and maintain larger aggregations. Food is considered the major contestable resource, but fish often demonstrate interference competition not for food, but for heterogeneous sites in the habitat, where they vigorously fight either for a shelter or just for visually non-uniform area. Visually heterogeneous sites can be used by fish as a template of a future individual territory, where fish can find not only food but also a refuge from predators. Fish use individual territories for much longer period than food patches. Just the presence of either physical refuge or "social refuge" neutralized the inhibiting effect of kairomons and allowed fish to feed more intensively despite the potential danger. We suggest that the decision-making was influenced only by available information of possibility to use a refuge. Habitat complexity is almost always accompanied by visual and other types of heterogeneity. Adaptive significance of fish attraction to the units of heterogeneity is probably related to the fact that under natural situations vital for fish objects are often tightly coupled with heterogeneous sites. Thus, units of habitat heterogeneity can be reliable signals or information cues in uncertain, i.e. changeable and poorly predictable, habitats.