Sublethal consequences of stream-dwelling predatory stoneflies on mayfly growth and fecundity
Abstract
Predators can have consequences on prey populations and communities that extend well beyond direct predator—induced mortality. Predator—prey interactions often affect prey feeding rates, growth rates, or fecundities, thereby significantly affecting reproductive success of prey adults. Thus, investigation of the sublethal fitness consequences of behavioral responses of prey to predators is essential to our understanding of the total impact of predators on prey populations and communities. Feeding (algal grazing) rates, growth rates, and fecundities of Baetis bicaudatus (Baetidae) larvae were measured in replicated circular flow—through stream channels to determine the sublethal consequences on fitness correlates of the presence of predatory stoneflies (Perlodidae). Gut fullness of 24—h starved Baetis larvae was measured using fluorometry after 24 h (short—term experiments) in four treatments: (1) ambient resource rocks and no predators, (2) low resource rocks and no predators, (3) ambient resource rocks and one female perlodid stonefly (Megarcys signata), or (4) Kogotus modestus whose mouthparts were glued to prevent lethal effects of predation. Mid—instar male and female Baetis larvae were reared for 3 wk until wing pad maturation (long—term experiments) and subjected to all but the Kogotus treatment. Predator avoidance significantly reduced levels of Baetis gut fullness, over the short term, to levels intermediate between the high and low resource treatments. Longer term residence with predatory stoneflies caused larvae of both male and female Baetis to mature at significantly smaller sizes than in the treatment with ambient food but no predators. Mayflies with predators matured at sizes similar to those held with low food levels over the long term. Interestingly, both starved larvae and those avoiding predators did not grow during the experimental period in contrast to about a 50% growth rate of larvae experiencing ambient food levels, but no predators. Similarly, egg biomass per mature female larva was significantly reduced in both starved and predator treatments. The proportion of total body mass allocated to eggs was only reduced in starved mayflies indicating that the females avoiding predators accrued less total mass, but allocated a similar proportion to eggs as did well—fed Baetis. These data convincingly demonstrate that predator avoidance can have dramatic consequences on mayfly fitness. The mechanism explaining those fitness consequences is probably a disruptive drift/swim response by Baetis to encounters with stoneflies, which reduces Baetis' feeding rates. If we accept the assumption that natural selection has shaped the foraging behavior of organisms to maximize fitness, foragers need to make decisions that maximize both survivorship (minimizing risk of predation) and fecundity. In mayflies fecundity is entirely a function of resource acquisition by the larvae, which makes them particularly vulnerable to such sublethal effects of predation. Nonetheless, we suspect that the types of sublethal costs of predator avoidance documented by this and other recent studies are nearly universal consequences or organisms foraging under predation risk.
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