Density-dependent costs of an inducible morphological defense in Crucian carp

Theoretical models for the evolution of inducible anti-predator defenses predict that there should be fitness costs associated with the defense, otherwise it should be permanent. Thus, in the absence of predators and with limited resources, defended individuals are expected to be outcompeted by individuals lacking the costly defense. Crucian carp, Carassius carassius, increase in body depth in response to chemical stimuli from piscivorous fish, and the deeper body constitutes a morphological defense against gape-limited piscivores. Here, we have performed a field experiment investigating effects of population density on the relative competitive performance of shallow-bodied and deep-bodied crucian carp. We assigned groups of six individually marked fish (the 'focal' individuals) of either morph into enclosures with either low or high crucian carp density, and monitored changes in body mass, length, and body depth of these fish for 4 mo. In addition, we determined sex, and final liver and gonadal mass, of all focal fish. At high population density, there was a reduction in food resources (zooplankters), which resulted in reduced gonad mass, reduced gonadosomatic indices, and reduced growth in fish of both morphs. Unexpectedly, the relative allocation to livers was higher at high density. There was no difference in body mass gain between morphs at the low density, whereas at the high density shallow-bodied fish gained twice as much body mass as deep-bodied fish. Hence, when predators were absent and food resources were limited, deep-bodied individuals suffered a substantial cost when competing with shallow-bodied conspecifics. This study verifies the theoretical prediction that an inducible defense results in fitness costs in the absence of predators. Further, as we found no measurable disadvantage of the deep-bodied morphology at high food availability but a strong disadvantage under intense competition for food, our results also demonstrate the importance of relating defense costs to levels of intraspecific competition. Key words: antipredator; body size; Carassius carassius; competition; cost; density dependence; field-enclosure experiment; fish; inducible defense; phenotypic plasticity; resource allocation; Sweden.
INTRODUCTION Predation is an important selective force in most habitats. Through direct, lethal effects, predators may have profound effects on abundance and size structure of prey populations and on species [...]