Experimental unit. (A) Top view of a cage, showing (a) the PVC ring of 25 cm in diameter, (b) the central compartment shown open to reveal (c) the Perumytilus purpuratus beds, and (d) the peripheral compartment with (e) specimens of the predatory snail Acanthina monodon. (B) View of a cage showing (f) the top mesh of the central compartment closed (using plastic cable ties) in the way it was maintained during the experiment. The cages were secured to the rocky intertidal substrate with (g) screws and PVC plates

Experimental unit. (A) Top view of a cage, showing (a) the PVC ring of 25 cm in diameter, (b) the central compartment shown open to reveal (c) the Perumytilus purpuratus beds, and (d) the peripheral compartment with (e) specimens of the predatory snail Acanthina monodon. (B) View of a cage showing (f) the top mesh of the central compartment closed (using plastic cable ties) in the way it was maintained during the experiment. The cages were secured to the rocky intertidal substrate with (g) screws and PVC plates

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Predators can exert nonconsumptive effects (NCEs) on prey, which often take place through prey behavioural adjustments to minimise predation risk. As NCEs are widespread in nature, interest is growing to determine whether NCEs on a prey species can indirectly influence several other species simultaneously, thus leading to changes in community struc...

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... Thanks to these properties, dense stands of foundation species often host a high diversity of associated species (Altieri & van de Koppel, 2014;Ellison, 2019;Thomsen et al., 2022). Foundation species dominate many terrestrial and aquatic communities and can be either primary producers, such as trees, shrubs, and seaweeds, or consumers, such as mussels and corals (Catalán et al., 2021(Catalán et al., , 2023Ellison et al., 2019;Stachowicz, 2001). In particular, the body structures of foundation species (branches, leaves, shells, etc.) create a tridimensional matrix where various environmental stresses are reduced and associated species find protection against large predators (Bulleri et al., 2016;He et al., 2013;Watt & Scrosati, 2013). ...
... These findings open the door to studies examining how the effects of foundation species on their associated communities might be influenced by other species. For example, predatory snails exert negative nonconsumptive effects on mussels through waterborne chemical cues, which in turn can affect the colonization of mussel stands by associated species (Catalán et al., 2021). Such effects might ultimately affect community functioning in such systems (Scrosati et al., 2022a(Scrosati et al., , 2022b. ...
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... Predation is a key biotic interaction that can control populations of prey species (Preisser and Bolnick 2008). Predators also affect prey populations through non-consumptive effects (NCEs) resulting from changes in the behavior, morphology, and life history traits of the prey (Kats and Dill 1998;Lima 1998;Peacor and Werner 2001;Trussell et al. 2003;Werner and Peacor 2003;Preisser et al. 2005;Preisser and Bolnick 2008;Holt 2009;Czarnoleski et al. 2011;Matassa and Trussell 2011;Gaynor et al. 2019;Catalán et al. 2021;Riedemann-Saldivia et al. 2022). Predator NCEs on prey (negative fitness changes in prey) can be triggered by chemical cues (e.g., exudates) from nearby predators, cues from attacked conspecific prey, or the presence of predator faeces with cues from consumed conspecific prey (Scherer and Smee 2016;Scherer et al. 2017;Dzierżyńska-Białończyk et al. 2019). ...
... A significant body of research has revealed NCEs on various prey species triggered by predator exudates. Examples are mayfly larvae of Baetis bicaudatus with predatory stoneflies or trout (Peckarsky et al. 1993;, the sea urchin Strongylocentrotus droebachiensis with Jonah crab Cancer borealis (Selden et al. 2009), whelks Nucella emarginata with the ochre sea star Pisaster ochraceus (Gosnell and Gaines 2012), the copepod Eurytemora affinis with the mysid Neomysis integer (Heuschele et al. 2014), the barnacle Semibalanus balanoides with the dogwhelk Nucella lapillus , the mussel Perumytilus purpuratus with the whelk Acanthina monodon (Catalán et al. 2021;Riedemann-Saldivia et al. 2022), and the mussel Mytilus spp. with the shore crab Carcinus maenas (Hubert et al. 2023). However, less attention has been paid to the effect of predator density or the level of predator exudates and whether prey can respond in a risk-sensitive manner to it (Loose and Dawidowicz 1994;Von Elert and Pohner 2000;Ferrari et al. 2006;Ferland-Raymond et al. 2010;Hill and Weissburg 2013). ...
... Moreover, predator chemical cues can indirectly affect community structure through NCEs on a foundation species, as those are species that often host many small species when occurring in dense stands. As marine invertebrate foundation species are abundant worldwide, understanding the effect of predator cue intensity on these organisms could help to better understand community regulation in systems structured by such species (Catalán et al. 2021). ...
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... Subordinates and transients can still significantly influence community functioning due to high recruitment, resource consumption, and growth rates under suitable environmental conditions (Gouhier et al., 2011;Bracken and Low, 2012). An important feature of dominant species, particularly sessile, is that they function as foundation species that change the local environment and resource availability through their own physical structure and metabolic activity (Dayton, 1972;Jones et al., 1994;Catalán et al., 2021). ...
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... Samples were collected between the middle and low intertidal zones at varying wave exposures. Invertebrates over 0.5 mm in size were preserved in 70 % ethanol and identified to the lowest possible taxonomic level with a dissecting microscope using identification guides[2][3][4] . ...
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... Much evidence for the community effects of intraspecific trait variation focuses on predators directly altering prey communities (Butler, 1989;Des Roches et al., 2013;Hughes et al., 2015;Katano, 2011;Matthews et al., 2016;Palkovacs & Post, 2009;Stemp et al., 2020;Wood et al., 2019) and genetic variation in foundation species (Bangert et al., 2008;Hays et al., 2021;Hughes et al., 2008;Whitham et al., 1999). However, predators of foundation species can have indirect effects on communities by altering habitat structure, both through nonconsumptive (Catalán et al., 2021) and consumptive effects (Hughes et al., 2015;Navarrete, 1996;Paine, 1966). Yet the community effects of trait variation in predators of foundation species remain untested. ...
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... An example is given by foundation species, which are organisms that typically form extensive canopies. These organisms dominate many terrestrial and aquatic communities and can be either primary producers (e.g., trees and algae) or consumers (e.g., corals and mussels; Catal an et al., 2021;Ellison et al., 2019;Stachowicz, 2001). By limiting abiotic stress in understory environments, these canopies often increase local species richness, or alpha diversity (Altieri & van de Koppel, 2014;Ellison, 2019;Watt & Scrosati, 2013). ...
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Beta diversity measures the spatial variation in species composition. Because it influences several community attributes, studies are increasingly investigating its drivers. Spatial environmental heterogeneity is a major determinant of beta diversity, but canopy‐forming foundation species can locally modify environmental properties. We used intertidal communities dominated by the canopy‐forming alga Mazzaella laminarioides as a model system to examine how a foundation species affects spatial environmental heterogeneity and the resulting beta diversity. Since canopies were found to reduce the spatial variation of temperature and desiccation during low tides, we hypothesized that canopies would decrease understory beta diversity, which we tested through a field experiment that contrasted canopy removal with presence treatments over 32 months. The beta diversity of sessile species was always lower under canopies, but canopies never affected the beta diversity of mobile species. The observed responses for sessile species may result from their abundance being more dependent on spatial abiotic variation than for mobile species, which can occur in stressful areas while temporarily foraging or in transit to other areas. These responses may likely apply to other systems exhibiting canopy‐forming foundation species hosting sessile and mobile species assemblages.
... Instead, prey may also increase growth rate to escape gape-limited predators 39 , but this often leads to costs in terms of a decreased size at maturity 40 and a reduced ability to neutralize free radicals 41,42 . Non-consumptive predator effects can have equally or even more negative consequences for prey communities than consumptive effects 43,44 . However, it is still unclear whether and how the non-consumptive predator effects impact SMPE in prey, especially when prey represent intermediate, cannibalistic predators in a food chain, and the predators are at the top of the food chain. ...
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... On a longer time-scale, reduced feeding rates will translate into reduced growth rates, to the benefit of the predator guild that will find a greater proportion of mussels below the size refuge. Parasite-induced fear is therefore likely to act as a major structuring force in intertidal communities, similar to the non-consumptive effects of predators on mussels that act as foundation species (Catalán et al. 2021). ...
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Fear is an integral part of predator–prey interactions with cascading effects on the structure and function of ecosystems. Fear of parasitism holds a similar ecological potential but our understanding of the underlying mechanisms in host–parasite interactions is limited by lack of empirical examples. Here, we experimentally test if blue mussels Mytilus edulis respond behaviourally to the mere presence of infective transmission stages of the trematode Himasthla elongata by ceasing filtration activity, thereby avoiding infection. Our results show that blue mussels reduced clearance rates by more than 30% in presence of parasites. The reduced filtration activity resulted in lower infection rates in experimental mussels. The identified parasite‐specific avoidance behaviour can be expected to play a significant role in regulating the ecosystem engineering function of blue mussels in coastal habitats.
... The purple mussel Perumytilus purpuratus is a dominant primary-space holder at middle and low elevations in rocky intertidal habitats on the southeastern Pacific coast (Lancellotti and Vásquez, 2000;Thiel and Ullrich, 2002). Upon detection of predators when submerged, individuals of P. purpuratus reduce the gap size between their valves and, thus, their filter-feeding activity (Vial et al., 1992;Catalán et al., 2021). However, it is still unclear how predator cues affect metabolic and physiological aspects of P. purpuratus (e.g., oxygen consumption and biodeposit production). ...
... Depending how long the NCEs last, and whether a compensatory increase in feeding occurs after prolonged predator exposure , this response could have consequences at the community level, since mussels are foundation species that provide substrate to many small species and can influence them through changes in filtration and excretion (Khalaman et al., 2021, Jahnsen-Guzmán et al., 2021. For instance, Catalán et al. (2021) found that waterborne cues from A. monodon decreased the larviphagy rate of P. purpuratus, which was ultimately related to an increase in the recruitment of other bivalves in P. purpuratus beds. Therefore, NCEs could increase the local abundance of pelagic bivalve larvae and promote a greater bivalve recruitment in coastal areas where A. monodon is abundant. ...
... This effect was attributed to negative NCEs on clearance rates of P. purpuratus that implied lower excretion levels and thus a reduction in the availability of nitrogenous products for algal growth. In our study, we found that predator cues decreased the inclusion of inorganic matter in the released biodeposits, which would limit algal development as suggested by Catalán et al. (2021). ...
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Predators can influence prey through direct consumption as well as through non-consumptive effects (NCEs). NCEs usually occur mediated by behavioral changes in the prey upon detection of predator cues. Such changes may involve reduction of feeding with a variety of physiological consequences. We evaluated NCEs from an intertidal predatory snail (Acanthina monodon) on a dominant habitat-forming mussel species (Perumytilus purpuratus) from the southeastern Pacific coast. We tested whether A. monodon exerts negative NCEs on clearance rate, oxygen consumption rate, biodeposit production, and between-valve gap size during feeding in P. purpuratus. We found that waterborne predator cues triggered a decrease in these variables except biodeposit production. However, the organic content of the biodeposits increased in the presence of predator cues. The snail’s physical contact with the mussels strengthened the negative NCEs on between-valve gap size. Since P. purpuratus is a dominant filter-feeder and foundation species in rocky intertidal habitats, predator NCEs on this species might indirectly influence ecosystem-level processes and community structure.
... Instead, prey may also increase growth rate to escape gape-limited predators 39 , but this often leads to costs in terms of a decreased size at maturity 40 and a reduced ability to neutralize free radicals 41,42 . Non-consumptive predator effects can have equally or even more negative consequences for prey communities than consumptive effects 43,44 . However, it is still unclear whether and how the non-consumptive predator effects impact SMPE in prey, especially when prey represent intermediate, cannibalistic predators in a food chain, and the predators are at the top of the food chain. ...
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Differences in hatching dates can shape intraspecific interactions through size-mediated priority effects (SMPE), a phenomenon where bigger, early hatched individuals gain advantage over smaller, late hatched ones. However, it remains unclear to what extent and how SMPE are affected by key environmental factors such as warming and predation risk imposed by top predators. We studied effects of warming (low and high temperature) and predation risk (presence and absence of predator cues of perch) on SMPE in life history and physiological traits in the cannibalistic damselfly Ischnura elegans . We induced SMPE in the laboratory by manipulating hatching dates, creating following groups: early and late hatchlings reared in separate containers, and mixed phenology groups where early and late hatchlings shared the same containers. We found strong SMPE for survival and emergence success, with the highest values in early larvae of mixed phenology groups and the lowest values in late larvae of mixed phenology groups. Neither temperature nor predator cues affected SMPE for these two traits. The other life history traits (development rate and mass at emergence) did not show SMPE, but were affected by temperature and predator cues. A tendency for SMPE was found for protein content, in the high temperature treatment. The other physiological traits (phenoloxidase activity and fat content) showed fixed expressions across treatments, indicating decoupling between physiology and life history. The results underline that SMPEs are trait-dependent, and only weakly or not affected by temperature and predation risk.