Species-specific antipredator capacities and prey refuges: interactions between piscivorous perch (Perca fluviatilis) and juvenile perch and roach (Rutilus rutilus)

The outcome of predator-prey interactions depends on the characteristics of predators and prey as well as the structure of the environment. In a replicated field enclosure experiment, we tested the effects of quantity and quality of different prey refuges (no structure, structure forming a partial refuge, and structure forming a complete refuge) on the interaction between piscivorous perch (Perca fluviatilis) and juvenile perch and roach (Rutilus rutilus). We quantified the behaviour of the predators and the prey and predator-induced prey mortality. The piscivores stayed in or close to the prey refuge and were more dispersed in the presence than in the absence of prey refuges. Survival of juvenile perch and roach decreased in the presence of predators and was higher for juvenile roach than for juvenile perch. In addition, juvenile perch survival increased with refuge efficiency Roach formed schools which were denser in the presence of predators, had a higher swimming speed (both in the open water and in the refuge) and used a larger area than juvenile perch. Both prey species decreased their distance to the prey refuge and increased the proportion of their time spent in the refuge in the presence of predators. The number of switches between the open-water habitat and the prey refuge was higher for juvenile roach than for juvenile perch. Juvenile perch used different parts of the prey refuge in a flexible way depending both on presence of predators and refuge type whereas juvenile roach used the different parts of the prey refuge in fixed proportions over all refuge treatments. Our results suggest that juvenile roach had a overall higher capacity to avoid predation than juvenile perch. However, in the presence of qualitatively different prey refuges juvenile perch responded to predators with more flexible refuge use than juvenile roach. The differences in antipredator capacities of juvenile perch and roach when subjected to piscivorous perch predation may depend on differences in life history patterns of the two species.     

Species-specific antipredatory behaviours: effects on prey choice in different habitats

Summary Prey species may use many different behaviours to avoid predation. In this study, the antipredator behaviours of juvenile roach (Rutilus rutilus) and juvenile perch (Perca fluviatilis) were studied in wading pools with three kinds of structural complexity: no structure, structure simulating vegetation and structure simulating bottom crevices. Predation experiments with piscivorous perch and habitat choice experiments with the prey were performed, and the foraging success and prey choice of the predators were related to the type of structure. Predator foraging success was lower in the vegetation than in the other treatments. In the absence of structure and with vegetation structure, predators preferred perch over roach, while the preference was reversed in the crevice treatment. Roach and perch differed in their antipredatory behaviours. Roach responded to the presence of predators by schooling, moving fast and remaining at the surface, and escaped from attacks by jumping out of the water. In contrast, perch moved more slowly, dispersed after attacks and tried to hide at the bottom. Perch always preferred the vegetation structure to the non-structured part of the pool, while roach showed preference for the vegetation structure only when predators were present. Roach never occurred in crevices, whereas perch used crevices when predators where present. Predator pursuit speed was lower in the vegetation structure than in the non-structured treatment, but prey escape speed was unaffected. The results suggest that both the quantity and quality of structural complexity interacting with species-specific antipredator behaviours are important for predator-prey dynamics. It is also suggested that the presence of structure can have substantial effects on the structure of North Eurasian fish communities, by affecting relative and absolute predation pressures from piscivorous perch on prey species. Correspondence to: B. Christensen     

Performance level and efficiency of two differing predator-avoidance strategies depend on nutritional state of the prey fish

Animal prey has developed a variety of behavioural strategies to avoid predation. Many fish species form shoals in the open water or seek refuge in structurally complex habitats. Since anti-predator strategies bear costs and are energy-demanding, we hypothesised that the nutritional state of prey should modify the performance level and efficiency of such strategies. In aquaria either containing or lacking a structured refuge habitat, well-fed or food-deprived juvenile roach (Rutilus rutilus) were exposed to an open-water predator (pikeperch, Sander lucioperca). Controls were run without predators. In the presence of the predator, roach enhanced the performance of the anti-predator strategy and increased the use of the refuge habitat whereby food-deprived roach were encountered more often in the structure than well-fed roach. Nonetheless more starved than well-fed roach were fed upon by the predator. In the treatments offering only open-water areas, roach always formed dense shoals in the presence of the predator. The shoal density, however, was lower in starved roach. Starving fish in shoals experienced the highest predation mortality across all experimental treatments. The experiment confirmed the plasticity of the anti-predator behaviour in roach and demonstrated that food deprivation diminished the efficiency of shoaling more strongly than the efficiency of hiding. The findings may be relevant to spatial distribution of prey and predator–prey interactions under natural conditions because when prey are confronted with phases of reduced resource availability, flexible anti-predator strategies may lead to dynamic habitat use patterns.     

Effects of predation and habitat structure on the abundance and population structure of the rock shrimp Rhynchocinetes typus (Caridea) on temperate rocky reefs

Human disturbances, such as overfishing, may disrupt predator–prey interactions and modify food webs. Underwater surveys were carried out at six shallow-water reef barrens in temperate waters of northern-central Chile from October to December 2010 to describe the effects of predation, habitat complexity (low, medium and high) and refuge availability on the abundance and population structure of the rock shrimp Rhynchocinetes typus (Rhynchocinetidae), an important mesoconsumer on subtidal hard substrata. Three sites were within managed (restricted access) areas for fishermen, and three were unmanaged (open-access). Field observations and tethering experiments were conducted to examine the relationship between fish and shrimp abundances, and the relative predation rates on shrimps. Direct effects of predation on R. typus body-size distribution were examined from shrimps collected in the field and fish stomachs. The presence and the abundance of R. typus increased with habitat reef complexity and refuge availability. Shrimp abundance was negatively related to fish abundance in managed areas, but not in open-access areas, where shrimp densities were the highest. Also, predation rates and body-size distribution of shrimps were unrelated, although fish consumed more large shrimps than should be expected from their distribution in the field. R. typus occurred most often in shelters with wide openings, offering limited protection against predators, but providing potential aggregation sites for shrimps. Overall, direct effects of predation on shrimp densities and population structure were weak, but indirect effects on shrimp distribution within reefs appear to have been mediated through behavioural responses. Our study highlights the need to assess both numerical and behavioural responses of prey to determine the effects of predator loss on mesoconsumer populations.     

Predation, habitat complexity, and variation in density-dependent mortality of temperate reef fishes

Density dependence in demographic rates can strongly affect the dynamics of populations. However, the mechanisms generating density dependence (e.g., predation) are also dynamic processes and may be influenced by local conditions. Understanding the manner in which local habitat features affect the occurrence and/or strength of density dependence will increase our understanding of population dynamics in heterogeneous environments. In this study I conducted two separate field experiments to investigate how local predator density and habitat complexity affect the occurrence and form of density-dependent mortality of juvenile rockfishes (Sebastes spp.). I also used yearly censuses of rockfish populations on nearshore reefs throughout central California to evaluate mortality of juvenile rockfish at large spatial scales. Manipulations of predators (juvenile bocaccio, S. paucispinus) and prey (kelp, gopher, and black-and-yellow [KGB] rockfish, Sebastes spp.) demonstrated that increasing the density of predators altered their functional response and thus altered patterns of density dependence in mortality of their prey. At low densities of predators, the number of prey consumed per predator was a decelerating function, and mortality of prey was inversely density dependent. However, at high densities of predators, the number of prey killed per predator became an accelerating response, and prey mortality was directly density dependent. Results of field experiments and large-scale surveys both indicated that the strength of density-dependent mortality may also be affected by the structural complexity of the habitat. In small-scale field experiments, increased habitat complexity increased the strength of density-dependent mortality. However, at large scales, increasing complexity resulted in a decrease in the strength of density dependence. I suggest that these differences resulted from scale-dependent changes in the predatory response that generated mortality. Whether increased habitat complexity leads to an increase or a decrease in the strength of density-dependent mortality may depend on how specific predatory responses (e.g., functional or aggregative) are altered by habitat complexity. Overall, the findings of this study suggest that rates of demographic density dependence and the resulting dynamics of local populations may largely depend upon attributes of the local habitat.     

Differences in relative abundance and size structure of the sea stars Pisaster ochraceus and Evasterias troschelii among habitat types in Puget Sound, Washington, USA

We surveyed patterns in the relative abundance and size structure of the sea stars Pisaster ochraceus and Evasterias troschelii in five habitat types of varying structural complexity and prey availability (sand/cobble, boulder, and rocky intertidal; pilings; and floating docks) in Puget Sound and the San Juan Islands, Washington. For both species, small sea stars were most abundant in the most structurally complex habitat type (boulder), where they occurred almost exclusively under boulders during low tide. Larger individuals became more abundant as structural complexity decreased, occurring more frequently in open habitat types (rocky shores, pilings, and docks) known to have greater abundances of prey resources. Gull foraging observations and experiments demonstrated that exposed small sea stars of both species were highly vulnerable to predation, suggesting that small sea stars require structural complexity (crevice microhabitat) as a predation refuge. Large sea stars, once attaining a size refuge from predation, appear to migrate to more exposed habitat types with more abundant food resources. These results suggest parallel ontogenetic habitat shifts in two co-occurring consumer species related to a shared predation risk at early life stages and demonstrate how the relative importance of top-down and bottom-up processes may differ with ontogeny.     

Predator recognition and anti-predator responses in the rainbowfish Melanotaenia eachamensis

Predator evasion behaviour patterns of three populations of rainbowfish (Melanotaenia eachamensis) were compared. The populations differed in the level of complexity of their natural habitats and the type and extent of predation. The predator recognition abilities of fish were assessed by exposing them to models differing in their degree of predator realism. The availability of vegetated cover and the location of the models with respect to cover were manipulated. Fish from Lake Tinaroo, a relatively open habitat containing numerous predators, showed strong changes in elective group size (EGS) in response to the different models but did not rely on cover as a place of refuge. In contrast, Dirran Creek fish originate from a small, fast-flowing, structurally complex stream lacking predatory fish species, and they showed little ability to distinguish between the different models and responded to threat by spending longer in vegetated areas. Members of the Lake Eacham captive stock increased their EGS in response to models representing low threat and with more threatening models increased the amount of time spent in vegetated regions of the arena. The contrasting reactions to predatory threat displayed by these populations highlights the need to use a number of different response indices when comparing the anti-predator responses of different fish populations. These data suggest that the level of habitat complexity as well as prior predator experience influence anti-predator responses of different fish popu-lations. Received: 2 October 1996 / Accepted after revision: 27 March 1997     

Do top-down or bottom-up forces determine Stephanitis pyrioides abundance in urban landscapes?

This study examined the influence of habitat structural complexity on the collective effects of top-down and bottom-up forces on herbivore abundance in urban landscapes. The persistence and varying complexity of urban landscapes set them apart from ephemeral agroecosystems and natural habitats where the majority of studies have been conducted. Using surveys and manipulative experiments. We explicitly tested the effect of natural enemies (enemies hypothesis), host plant quality, and herbivore movement on the abundance of the specialist insect herbivore, Stephanitis pyrioides, in landscapes of varying structural complexity. This herbivore was extremely abundant in simple landscapes and rare in complex ones. Natural enemies were the major force influencing abundance of S. pyrioides across habitat types. Generalist predators, particularly the spider Anyphaena celer, were more abundant in complex landscapes. Predator abundance was related to greater abundance of alternative prey in those landscapes. Stephanitis pyrioides survival was lower in complex habitats when exposed to endemic natural enemy populations. Laboratory feeding trials confirmed the more abundant predators consumed S. pyrioides. Host plant quality was not a strong force influencing patterns of S. pyrioides abundance. When predators were excluded, adult S. pyrioides survival was greater on azaleas grown in complex habitats, in opposition to the observed pattern of abundance. Similarly, complexity did not affect S. pyrioides immigration and emigration rates. The complexity of urban landscapes affects the strength of top-down forces on herbivorous insect populations by influencing alternative prey and generalist predator abundance. It is possible that habitats can be manipulated to promote the suppressive effects of generalist predators.     

Scale-dependent patterns of variability of a grazing parrotfish (<Emphasis Type="Italic">Leptoscarus vaigiensis</Emphasis>) in a tropical seagrass-dominated seascape

Although herbivorous fish form critical linkages between primary producers and higher trophic levels, the major factors regulating their spatial structure in seagrass systems remain poorly understood. The present study examined the parrotfish Leptoscarus vaigiensis in seagrass meadows of a tropical embayment in the western Indian Ocean. Stomach content analysis and direct field observations showed that L. vaigiensis is an efficient grazer, feeding almost exclusively on seagrass leaves. Seagrass shoot density was highly correlated to all density variables (total, juvenile and subadult) and juvenile biomass of L. vaigiensis, while subadult biomass was predicted by distance to neighbouring coral habitat. Moreover, density and biomass of predatory fish (piscivores) were predicted by seagrass canopy height and the distribution patterns of predators followed those of L. vaigiensis. Hence, factors at local (seagrass structural complexity and feeding mode) and landscape scale levels (seascape context and distribution of piscivores) likely mutually structure herbivorous fish communities. The findings underscore the importance of incorporating multiple scale-dependent factors when managing coastal seagrass ecosystems and their associated key species.     

What attracts juvenile coral reef fish to mangroves: habitat complexity or shade?

The value of mangroves for fish species is usually explained in terms of high food abundance or shelter against predators as a result of high turbidity and structural complexity. In a field experiment, artificial mangrove units (AMUs) were designed as open cages, each of which was provided with a different degree of structural complexity and shade. Fish species that were attracted to the AMUs were identified and counted and the effects of shade and structural complexity, as well as the interaction between the two factors, were tested. Diurnal fish showed a preference for the greatest structured complexity and for a moderate increase in shade. Two nocturnal species common in local mangroves as well as seagrass beds showed statistically significant effects: densities of Haemulon sciurus were positively related to both shade and structural complexity, whereas only shade had a significant positive effect on densities of Ocyurus chrysurus. The experiment indicated that the attractiveness of mangrove vegetation for H. sciurus may be influenced by the structural complexity of the habitat as well as by the degree of shade, and that both factors are equally and separately important. Individuals of O. chrysurus that are attracted to mangroves are more likely to be influenced by the presence of shade than by the degree of structural complexity. The data thus indicated that the positive relationship between fish densities in mangrove habitats and the degree of shade and structural complexity, or both, is species-specific.Communicated by O. Kinne, Oldendorf/Luhe     

Is boldness affected by group composition in young-of-the-year perch (Perca fluviatilis)?

In young-of-the-year perch (Perca fluviatilis), individuals within groups differed in the degree of boldness, estimated by habitat utilisation and feeding activity in visual contact with a potential predator. We looked at changes in individual behaviour in connection with change of group composition. During the first period, perch were randomly assigned to groups, and time spent in open habitat versus in vegetation and number of prey attacks were registered. The perch were then categorised into personality types (shy, bold, intermediate) according to their behaviour. During the second period, fish were observed when sorted into new groups, each containing only one personality type. Shy individuals showed the largest changes in behaviour, and increased both the time spent in the open and the number of prey attacks when placed into the new groups. Feeding activity in shy fish during the second period was affected by group composition during the first period. After regrouping, bold individuals decreased their time in the open, whereas intermediate individuals did not change behaviour. Time in the open habitat was, to some extent, influenced by the behaviour of the other members of the group, but number of prey attacks was not. The behaviour of fish of the different personality types we have defined in this study seemed to be based on innate traits, but also modified by the influence of other group members and by habituation to the environment.Communicated by J.Krause     

Diet of young Atlantic bluefin tuna (<Emphasis Type="Italic">Thunnus thynnus</Emphasis>) in eastern and western Atlantic foraging grounds

Atlantic bluefin tuna (Thunnus thynnus) are highly migratory predators whose abundance, distribution, and somatic condition have changed over the past decades. Prey community composition and abundance have also varied in several foraging grounds. To better understand underlying food webs and regional energy sources, we performed stomach content and stable isotope analyses on mainly juvenile (60–150 cm curved fork length) bluefin tuna captured in foraging grounds in the western (Mid-Atlantic Bight) and eastern (Bay of Biscay) Atlantic Ocean. In the Mid-Atlantic Bight, bluefin tuna diet was mainly sand lance (Ammodytes spp., 29% prey weight), consistent with historic findings. In the Bay of Biscay, krill (Meganyctiphanes norvegica) and anchovy (Engraulis encrasicolus) made up 39% prey weight, with relative consumption of each reflecting annual changes in prey abundance. Consumption of anchovies apparently declined after the local collapse of this prey resource. In both regions, stable isotope analysis results showed that juvenile bluefin tuna fed at a lower trophic position than indicated by stomach content analysis. In the Mid-Atlantic Bight, stable isotope analyses suggested that >30% of the diet was prey from lower trophic levels that composed <10% of the prey weights based upon traditional stomach content analyses. Trophic position was similar to juvenile fish sampled in the NW Atlantic but lower than juveniles sampled in the Mediterranean Sea in previous studies. Our findings indicate that juvenile bluefin tuna targeted a relatively small range of prey species and regional foraging patterns remained consistent over time in the Mid-Atlantic Bight but changed in relation to local prey availability in the Bay of Biscay.     

Modeling the Effects of Fishing and Implications for the Design of Marine Protected Areas: Juvenile Fish Responses to Variations in Seafloor Habitat

Abstract: A number of recent studies have linked post-settlement survivorship of Atlantic cod (  Gadus morhua ) with the complexity of the seafloor to which fish settle. Survivorship is greater in habitats of higher complexity (e.g., pebble-cobble substratum with emergent epifauna> pebble-cobble> sand), where cover provides shelter from predators. Fishing with mobile gear such as bottom trawls and dredges reduces the complexity of seafloor habitats. We used a dynamic model to (1) link patterns in habitat-mediated survivorship of post-settlement juvenile cod with spatial variations in habitat complexity, (2) simulate habitat change based on fishing activities, and (3) determine the role of marine protected areas in enhancing recruitment success. Density-dependent natural mortality was specified as three alternative functional response curves to assess the influence of different predator foraging strategies on juvenile survivorship during the first 12 months of demersal existence. We applied the model to a theoretical patch of hard-bottom substrata and to a case study based on seafloor habitat distributions at Stellwagen Bank National Marine Sanctuary (Gulf of Maine, Northwest Atlantic). Our results demonstrate that patterns in the shape of response surfaces that show the relationship between juvenile cod survivorship and density as well as movement rate were similar regardless of functional response type, that juvenile cod movement rates and post-settlement density were critical for predicting the effects of marine protected-area size on survivorship, and that habitat change caused by fishing has significant negative effects on juvenile cod survivorship and use of marine protected areas can ameliorate such effects.     

An ecological framework for contextualizing carnivore–livestock conflict

Carnivore predation on livestock is a complex management and policy challenge, yet it is also intrinsically an ecological interaction between predators and prey. Human–wildlife interactions occur in socioecological systems in which human and environmental processes are closely linked. However, underlying human–wildlife conflict and key to unpacking its complexity are concrete and identifiable ecological mechanisms that lead to predation events. To better understand how ecological theory accords with interactions between wild predators and domestic prey, we developed a framework to describe ecological drivers of predation on livestock. We based this framework on foundational ecological theory and current research on interactions between predators and domestic prey. We used this framework to examine ecological mechanisms (e.g., density-mediated effects, behaviorally mediated effects, and optimal foraging theory) through which specific management interventions operate, and we analyzed the ecological determinants of failure and success of management interventions in 3 case studies: snow leopards (Panthera uncia), wolves (Canis lupus), and cougars (Puma concolor). The varied, context-dependent successes and failures of the management interventions in these case studies demonstrated the utility of using an ecological framework to ground research and management of carnivore–livestock conflict. Mitigation of human–wildlife conflict appears to require an understanding of how fundamental ecological theories work within domestic predator–prey systems.     

Patterns of shell repair in articulate brachiopods indicate size constitutes a refuge from predation

The cost of overcoming prey defenses relative to the value of internal tissues is a key criterion in predator/prey interactions. Optimal foraging theory predicts: (1) specific sizes of prey will result in the best returns to predators, and (2) there will often be a size at which the cost/benefit balance is low enough to effectively exclude predation. Data presented here on styles of repaired shell damage and size at which injury had been sustained was collected from samples of terebratulide brachiopods from the Antarctic Peninisula (Liothyrella uva), Falkland Islands (Magellania venosa and Terebratella dorsata) and Chile (M. venosa). The predominant form of damage on shells was indicative of predators attacking the valve margins. The modal size for repaired damage was more than 10 mm smaller than the modal size for the overall size distribution in each species and there were no repaired attacks in the largest size classes of any species. These data suggest that size forms a refuge from predation, as would be predicted by optimal foraging theory. The optimal sizes that predators appeared to attack vary between species, as do the sizes that provided a refuge from predation. High levels of multiple repairs (19% of the M. venosa population from the Falkland Islands sampled had 2 or more repairs) suggest that the mortality following attack is low, suggesting that many predators abandon their attacks.     

The role of turbidity as a constraint on predator-prey interactions in aquatic environments

Many of the world's most productive aquatic ecosystems usually contain turbid water. Paradoxically, many fish species that live in these habitats are also those that often rely on vision to detect their predators and their prey. For these fish, turbidity will reduce the distance at which predator-prey interactions occur, and there should be a reduction in the opportunity for behavioural modification to control the risk of predation. Under these conditions, we predict that most antipredator behaviour will become ineffective and that predator-prey interactions in turbid water will be primarily characterised by the direct effects of predator consumption of prey, rather than behavioural modification reducing the growth rates of prey. This hypothesis was tested in two laboratory experiments. The first experiment investigated how water turbidity, risk of predation, and their interaction affect habitat choice decisions by fathead minnows (Pimephales promelas). These data demonstrate that fathead minnows reduce their use of dangerous habitats, but that this effect is diminished in turbid water. A second experiment examined mortality patterns when these fish were preyed upon by yellow perch (Perca flavescens) in clear and turbid water. Absolute mortality rates were unaffected by visibility, but patterns of mortality were random in turbid water and skewed towards smaller individuals in clear water. Combined, these results support our hypothesis and suggest that the impact of predation risk will be reduced in turbid aquatic ecosystems. Received: 23 May 1996 / Accepted after revision: 18 November 1996     

Habitat use by juvenile Atlantic cod (Gadus morhua) in the presence of an actively foraging and non-foraging predator

Experiments were conducted in the autumn and winter of 1992/93 to examine habitat use by juvenile (age 0+) Atlantic cod, Gadus morhua L., before, during and following exposure to a passive or actively foraging predator (age 3+ cod). Experiments presented groups of juvenile cod (n=5 fish/group) with one of two combinations of three substrates; (1) gravel, sand, and a patch of artificial kelp (kelp), or (2) cobble, sand, and kelp. Cobble is known to provide juvenile cod with a refuge from predation. Kelp was used to test the hypothesis that juvenile cod associate with fleshy macroalgae in nature because of the safety it provides from predators. There was little difference in habitat use by juvenile cod before, during or following exposure to a passive predator. Under these conditions, juvenile cod appeared to prefer finer grained mineral substrates and avoided the kelp. The extent of the juvenile response to a passive predator was to avoid the predator's location in the experimental tank. In contrast, juvenile cod showed a significant shift in habitat use when exposed to an actively foraging predator, hiding in cobble or, when cobble was not available, in kelp. Use of both these habitats resulted in a significant reduction in predation risk to the juvenile cod. Our results suggest that: (1) an association with kelp provides safety from predation to juvenile cod, and (2) juvenile cod are capable for assessing the risk a predator represents and adjust their response accordingly.     

Distance to neighbours influences the trade-off between hiding after disturbance and defending food patches in convict cichlids (<Emphasis Type="Italic">Archocentrus nigrofasciatus</Emphasis>)

Territories are often aggregated. Because of this, distance to neighbours should influence how territory-holders balance safety from predators with the use and defence of resources. I examined the influence of distance to a neighbour on refuge use by pairs of convict cichlids (Archocentrus nigrofasciatus) faced with a conflict between hiding and defending food patches. Neighbours could reduce the rate of intrusions by strangers as a by-product of their own resource defence. This should allow fish with near neighbours to spend more time in the refuge. Neighbours could also steal from patches that are left undefended. This should lead to a reduction in use of the refuge. When one fish was confined to its refuge (so that its patch was undefended), theft by the other increased as inter-patch distance decreased. Distance between patches did not influence the rate of intrusion by non-territorial fish. When both fish defended patches, body mass influenced the effect of inter-patch distance on refuge use. Large fish rarely used the refuge, but small territory-holders spent more time in the refuge when patches were close together, as predicted. However, when one fish was dominant at both patches, distance between patches did not influence refuge use. These results suggest that, despite increased opportunity for theft, there is no realised foraging or defensive benefit to settling near neighbours that are of similar competitive ability.Communicated by J. Krause     

Habitat complexity impacts persistence and species interactions in an intertidal whelk

Although experiments have shown that habitat structure may influence the distribution of species and species interactions, these effects are still not commonly integrated into studies of community dynamics. Since habitat structure often varies within and among communities, this may limit our understanding of how various factors influence communities. Here, we examined how mussel bed complexity (the presence and thickness of mussel layers) influenced the persistence of whelks (Nucella emarginata) and interactions with a top predator (ochre sea stars, Pisaster ochraceus) and prey (mussels, Mytilus californianus). Results from a mark?Crecapture experiment indicate that whelk recapture rates are higher in more complex habitats, and laboratory experiments demonstrate that habitat complexity affects whelk feeding, growth, and nonconsumptive interactions with a keystone predator. Habitat complexity therefore has direct effects on species and also may lead to trade-offs among feeding, refuge, and other factors, potentially influencing the distribution of whelks and the effects of both whelks and sea stars on intertidal communities. These results demonstrate that habitat structure may play an important role in intertidal communities and other habitats and should be further considered in the experimental design of future studies of community dynamics.     

Learning to recognize novel predators under weakly acidic conditions: the effects of reduced pH on acquired predator recognition by juvenile rainbow trout

Summary. Recent studies have demonstrated that under weakly acidic conditions (pH 6.0), many prey fishes, including juvenile rainbow trout (Onchorhynchus mykiss), do not exhibit overt antipredator responses to conspecific chemical alarm cues. In laboratory trials, we investigated the potential effects of reduced pH on the ability of hatchery reared, predator naïve juvenile rainbow trout to acquire the recognition of a novel predator (yellow perch, Perca flavenscens). Initially, we exposed trout to the odour of a predatory yellow perch, buffered to pH 6.0 (weakly acidic) or pH 7.0 (neutral) paired with conspecific skin extracts (also buffered to pH 6.0 or 7.0) or a distilled water control. Juvenile trout exhibited significant increase in antipredator behaviour when exposed to neutral skin extract (pH 7.0). When retested 48 hours later to perch odour alone (pH 7.0), only trout initially conditioned with neutral skin extracts (pairs with either neutral or acidic perch odour) exhibited a learned recognition of perch odour as a predator risk. Those initially exposed to weakly acidic skin extract or the distilled water control did not show a learned response to predator odour. These results demonstrate that the ability to acquire the recognition of novel predators is impaired under weakly acidic conditions, as would occur in natural waterways affected by acidic precipitation.