Predation risk by largemouth bass modulates feeding functional responses of native and non-native crayfish

Context -dependency is prevalent in nature, challenging our understanding and prediction of the potential ecological impacts of non-native species (NNS). The presence of a top predator, for example, can modify the foraging behaviour of an intermediate consumer, by means of non -consumptive effects. This raises the question of whether the fear of predation might modulate consumption rates of NNS, thus shaping the magnitude of ecological impacts. Here, we quantified the functional feeding responses of three non-native crayfish species - red swamp crayfish Procambarus clarkii, rusty crayfish Faxonius rusticus and virile crayfish Faxonius virilis - compared to the native analogue signal crayfish Pacifastacus leniusculus, considering the predation risk imposed by a top fish predator, the globally invasive largemouth bass Micropterus salmoides. We applied the comparative functional response (FR) approach using snails as prey and exposing crayfish to water containing predator and dietary chemical cues or not. All crayfish species presented a destabilising Type II FR, regardless of the presence of chemical cues. Predation risk resulted in significantly longer handling times or lower attack rates in non-native crayfish; however, no significant differences were observed in signal crayfish. We estimated per capita impacts for each species using the functional response ratio (FRR; attack rate divided by handling time). The FRR metric was lower for all crayfish species when exposed to predation risk. Rusty crayfish demonstrated the highest FRR in the absence of chemical cues, followed by signal crayfish, virile crayfish and red swamp crayfish. By contrast, the FRR of signal crayfish was nearly twice that of rusty crayfish and virile crayfish and ten times greater than red swamp crayfish when chemical cues were present. The latter result agrees with the well -recognised ecological impacts of signal crayfish throughout its globally -introduced range. This study demonstrates the importance of considering the non -consumptive effects of predators when quantifying the ecological impacts of intermediate non-native consumers on prey. The direction and magnitude of the modulating effects of predators have clear implications for our understanding of NNS impacts and the prioritisation of management actions.