Invasive predators can strongly affect native populations. If alien predator pressure is strong enough, it can induce anti-predator responses, including phenotypic plasticity of exposed individuals and local adaptations of impacted populations. Furthermore, maternal investment is an additional pathway that could provide resources and improve performance in the presence of alien predators. We investigated the potential responses to an alien predator crayfish (Procambarus clarkii) in a threatened frog (Rana latastei) by combining field observations with laboratory measurements of embryo development rate, to assess the importance of parental investment, origin and exposure to the crayfish cues. We detected a strong variation in parental investment amongst frog populations, but this variation was not related to the invasion status of the site of origin, suggesting that mothers did not modulate parental investment in relation to the presence of alien predators. However, cues of the invasive crayfish elicited plastic responses in clutches and tadpoles development: embryos developed faster when exposed to the predator. Furthermore, embryos from invaded sites reached Gosner’s development stage 25 faster than those from non-invaded sites. This ontogenetic shift can be interpreted as a local adaptation to the alien predator and suggests that frogs are able to recognise the predatory risk. If these plastic responses and local adaptation are effective escape strategies against the invasive predator, they may improve the persistence of native frog populations.

During biotic invasions, native prey are abruptly exposed to novel predators and are faced with unprecedented predatory pressures. Under these circumstances, the lack of common evolutionary history may hamper predator recognition by native prey, undermining the expression of effective antipredator responses. Nonetheless, mechanisms allowing prey to overcome evolutionary naïveté exist. For instance, in naïve prey, history of coevolution with similar native predators or detection of general traits characterizing predators can favor the recognition of stimuli released by invasive predators. However, few studies have assessed how these mechanisms shape prey response at the community level. Here, we evaluated behavioral responses in naïve larvae of 13 amphibian species to chemical and visual cues associated with an invasive predator, the American red swamp crayfish (Procambarus clarkii). Moreover, we investigated how variation among species responses was related to their coexistence with similar native crayfish predators. Amphibian larvae altered their behavior in presence of visual stimuli of the alien crayfish, while chemical cues elicited feeble and contrasting behavioral shifts. Activity reduction was the most common and stronger response, whereas some species exhibited more heterogeneous strategies also involving distancing and rapid escape response. Interestingly, species sharing coevolutionary history with the native crayfish were able to finely tune their response to the invasive one, performing bursts to escape. These results suggest native prey can respond to invasive predators through recognition of generic risk cues (e.g., approaching large shapes), still the capability of modulating antipredator strategies may also depend on their coevolutionary history with similar native predators.

Microhabitat selection is a key component of amphibian breeding biology and can be modulated in response to the features of breeding sites and the presence of predators. Despite invasive alien species being among the major threats to amphibians, there is limited information on the role of invasive species in shaping amphibians' breeding microhabitat choice. The invasive red swamp crayfish (Procambarus clarkii) is a major predator of amphibians' larvae, including those of the brown frogs Rana dalmatina and Rana latastei. Although qualitative information about the spawning site preferences and breeding microhabitat choice of brown frogs is available in the literature, only a few studies performed quantitative analyses, and the relationship between microhabitat choice and the presence of alien predators has not been investigated yet. The aims of this study were: (1) to characterize the microhabitats selected for clutch deposition by R. dalmatina and R. latastei and (2) to test if the position and the aggregation of egg clutches differ in sites invaded or not invaded by P. clarkii. During spring 2017, we surveyed multiple times 15 breeding sites of both brown frogs in Northern Italy; in each site we assessed the features of the microhabitat where each egg clutch was laid, considering its position (distance from the shore, depth of the water column) and the degree of aggregation of clutches. In each site we also assessed the presence/absence of the invasive crayfish and the relative abundance in the breeding period. We detected egg clutches in all sites; the crayfish occurred in eight ponds. Our results showed substantial differences between the spawning microhabitat features of the two brown frogs: Rana latastei clutches showed a higher degree of aggregation and were associated with deeper areas of the ponds , while Rana dalmatina deposited more spaced out clutches in areas of the ponds that were less deep. For both species, spawning microhabitat features were not significantly different between sites with and without P. clarkii. Although we did not detect behavioural responses to P. clarkii in the choice of spawning microhabitat , additional studies are required to assess whether these frogs modulate other behavioural traits (e.g. during larval development) in response to the invasive predator. How to cite this article Romagnoli S, Ficetola GF, Manenti R. 2020. Invasive crayfish does not influence spawning microhabitat selection of brown frogs. PeerJ 8:e8985 http://doi.org/10.7717/peerj.8985 Romagnoli et al. (2020), PeerJ, DOI 10.7717/peerj.8985 2/16Arribas R, Díaz-Paniagua C, Gomez-Mestre I. 2014. Ecological consequences of amphibian larvae and their native and alien predators on the community structure of temporary ponds.

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