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.     

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.     

Ecology of Conus on eastern Indian Ocean fringing reefs: Diversity of species and resource utilization

The most diverse assemblages of the genus Conus known occur on fringing coral reefs in Thailand and Indonesia. As many as 27 congeneric species of these gastropods inhabit a single reef; in all, we examined 1,350 individuals of 48 species. Several attributes of the populations we observed conform to expectations of a model of ecological characteristics of bench and reef Conus proposed by Kohn (1971a). Number of species (S) averaged 15, and species diversity (H″) averaged 2.3 in the most heterogeneous habitat type — topographically complex, subtidal reef platforms (Type III habitat). Both species richness and evenness of distribution of individuals among species contribute strongly to H″. Fewer congeners and greater numerical dominance by single species characterize more homogeneous habitats. On subtidal reef platforms with large areas of sand substrate and less coral limestone (Type I–III habitat), mean values were S=10, and H″=1.6. In the one intensively studied, truncated reef-limestone platform (Type II–III intermediate habitat), S=13 and H″=1.4. Summed population density of all Conus species in Type III and I–III habitats is similar (0.02 to 0.05 individuals /m²) and comparable to estimates from similar habitats elsewhere in the Indo-West Pacific region. Mean density (0.7/m²) and other population attributes in Type II–III habitat more closely resemble those of Type II than Type III habitats in general. We combined analysis of species diversity and other attributes of assemblages in habitats of different environmental complexity with analysis of microhabitat and food-resource utilization, in order to demonstrate the extent to which specialization on different resources occurs in assemblages differing in diversity and habitat type. In the habitats studied, co-occurring species of Conus specialized to a greater extent on different prey species than on different microhabitat patches, but degree of microhabitat specialization was greater than in similarly complex habitats with assemblages of lower diversity elsewhere in the Indo-West Pacific region. While most Conus species preyed primarily on a different species or higher taxon of polychaetes, diets are not more specialized or dissimilar than in similar habitats elsewhere. Degree of specialization on different prey is not correlated with Conus species diversity in the different types of habitats studied. The data lead to the conclusion that differential predation is as important — and differential microhabitat utilization is more important — in permitting coexistence of potentially competing congeners, compared with conditions in habitats of comparable heterogeneity that support fewer congeners farther from the center of the Indo-West Pacific region. Pairwise comparisons of congeners indicate that many species pairs have low or no overlap in both microhabitat and food utilization. Members of species pairs with high overlap in microhabitat utilization typically eat different prey organisms, and those with similar diets typically occupy different habitats or microhabitats. This applies to molluscivorous as well as vermivorous species. Information on the diets of 11 species is reported here for the first time. Of 48 Indo-West Pacific Conus species whose food is now known, 35 prey on polychaetes, 2 on enteropneusts, 6 on gastropods, and 5 on fishes. Vermivorous Conus prey on relatively few of the polychaete species present in the environments. Species eaten represent only 12% of a total estimated polychaete population density of 27,000 individuals /m². Certain very abundant polychaetes may be protected from predation by Conus by their small size, others by their long tubes. Two new aspects of size-selective predation by Conus are reported: (1) Although comparisons of predation rate with prey standing-crop suggest that food is plentiful, selective predation on the largest prey individuals present suggests that only small proportions of prey-species populations may have large enough body size to repay foraging effort by the Conus present; (2) composition of the diet changes qualitatively with increase in body size in several vermivorous Conus species; shifting by larger individuals to larger prey species could be documented in C. ebraeus.     

Community-wide ramifications of an associational refuge on shallow rocky reefs

Little attention has been given to associational refuges in ecology, despite their potential for maintaining species diversity and supporting higher trophic levels. Here I show how the colonial anemone, Corynactis californica, creates a refuge for benthic macroalgae and invertebrate fish prey on intensively grazed shallow rocky reefs in the Santa Barbara Channel off southern California, U.S.A. On reefs heavily grazed by sea urchins, benthic macroalgae and invertebrate fish prey were relatively more abundant among Corynactis colonies than adjacent areas lacking the anemone. Results from field experiments showed that Corynactis facilitated the recruitment of macroalgae and tubicolous amphipods in "urchin-barren" areas subjected to intensive grazing. In areas forested by giant kelp (Macrocystis pyrifera), where grazing intensity from urchins was low, Corynactis suppressed algal recruitment but facilitated tubicolous amphipods. A manipulation of fish and sea urchins suggested that grazing by urchins, as opposed to predation from fish (primarily surfperch Embiotocidae), suppressed tubicolous amphipods, and this activity was hindered by the presence of Corynactis. In systems where human activity has intensified herbivory, associational refuges may maintain species diversity and support higher trophic levels.     

Abundance patterns of subtidal solitary ascidians in the San Juan Islands,Washington, as influenced by food preferences of the predatory snail Fusitriton oregonensis

The assemblage of solitary ascidians in rocky subtidal habitats of the San Juan Islands, Washington, USA, is dominated numerically by a few species of stolidobranchs: Pyura haustor, Halocynthia igaboja, Cnemidocarpa finmarkiensis and Boltenia villosa. Several phlebobranch species occur abundantly only on floating docks and/or muddy bottoms, where they co-occur with P. haustor and other stolidobranchs. These patterns of abundance correlate with feeding preferences and distributional patterns of the predatory snail Fusitriton oregonensis, which occurs only in rocky subtidal sites, prefers ascidians over other invertebrates as prey and prefers phlebobranchs over stolidobranchs. With the exception of B. villosa, stolidobranchs commonly co-occuring with the snail in the rocky subtidal are not eaten. Transplants of phlebobranchs from docks to rocky subtidal sites and transplants of F. oregonensis into established dock communities suggest that predation by the snail may largely explain the observed patterns of absolute and relative abundance.     

Behavioral response to predators reverses the outcome of competition between prey species

Summary In a laboratory experiment it was shown that piscivorous predators reversed the outcome of competitive interactions between two fish prey species, juveniles of roach (Rutilus rutilus) and perch (Perca fluviatilis), by behaviorally affecting their use of two available habitats, an open water habitat and a structurally complex refuge. The shift in the competitive relationship was the result of predators forcing the juvenile fishes into a prey refuge with high structural complexity. While roach was competitively superior in the unstructured habitat, perch was superior in the structurally complex prey refuge. The reversal in competitive relationship was demonstrated both with respect to foraging rate and growth rate and resulted from the high structural complexity in the prey refuge interfering with the roach's swimming performance. Because survival and growth patterns through the juvenile stages have profound effects on the population/community dynamics of size-structured populations such as those of fish, behaviorally induced changes in competitive ability should have significant implications also at the population and community levels.     

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.     

Patch use and vigilance by sympatric lemmings in predator and competitor-driven landscapes of fear

Prey living in risky environments can adopt a variety of behavioral tactics to reduce predation risk. In systems where predators regulate prey abundance, it is reasonable to assume that differential patterns of habitat use by prey species represent adaptive responses to spatial variation in predation. However, patterns of habitat use also reflect interspecific competition over habitat. Collared (Dicrostonyx groenlandicus) and brown (Lemmus trimucronatus) lemmings represent such a system and possess distinct upland tundra versus mesic meadow habitat preferences consistent with interspecific competition. Yet, we do not know whether this habitat preference might also reflect differences in predation risk or whether the two species differ in their behavioral tactics used to avoid predation. We performed experiments where we manipulated putative predation risk perceived by lemmings by increasing protective cover in upland and meadow habitats while we recorded lemming activity and behavior. Both lemming species preferentially used cover more than open patches, but Dicrostonyx was more vigilant than Lemmus. Both species also constrained their activity to protective patches in upland and meadow habitats, but during different periods of the day. Use of cover and vigilance were independent of habitat, suggesting that both species live in a fearsome but flattened landscape of fear at Walker Bay (Nunavut, Canada), and that their habitat preference is a consequence of competition rather than predation risk. Future studies aiming to map the contours of fear in multi-prey–predator systems should consider how predation and competition interact to modify prey species’ habitat preference, patch use, and vigilance.     

Interactions among sea otters, sea stars, and suspension-feeding invertebrates in the western Aleutian archipelago

The structuring and organizing effects of apex predators on ecosystems are becoming increasingly well documented. The enhancement of kelp forests via sea otter predation on herbivorous sea urchins is among the earliest and best known examples. This study provides evidence for direct and indirect trophic interactions among sea otters, predatory sea stars, and filter-feeding mussels (Mytilus trossulus) and barnacles (Semibalanus cariosis). In western Massacre Bay at Attu Island (173°E, 53°N), subtidal transects showed sea star body size and biomass density declined markedly between 1983 and 1994 as sea otters reinhabited this area. Mussels and barnacles translocated from the rocky intertidal zone to shallow subtidal habitats to assess loss rates from sea star predation showed lower mortality rates after the arrival of sea otters. Prey mortality rates in subtidal caged controls were consistently low and similar to those of intertidal controls in both years. These findings elucidate a trophic pathway by which sea otters can influence ecosystems separate from the well-known sea otter/sea urchin/macroalgae cascade.     

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.     

Long-term trends in invertebrate–habitat relationships under protected and fished conditions

Few studies examine the long-term effects of changing predator size and abundance on the habitat associations of resident organisms despite that this knowledge is critical to understand the ecosystem effects of fishing. Marine reserves offer the opportunity to determine ecosystem-level effects of manipulated predator densities, while parallel monitoring of adjacent fished areas allows separating these effects from regional-scale change. Relationships between two measures of benthic habitat structure (reef architecture and topographic complexity) and key invertebrate species were followed over 17 years at fished and protected subtidal rocky reefs associated with two southern Australian marine reserves. Two commercially harvested species, the southern rock lobster (Jasus edwardsii) and blacklip abalone (Haliotis rubra) were initially weakly associated with habitat structure across all fished and protected sites. The strength of association with habitat for both species increased markedly at protected sites 2 years after marine reserve declaration, and then gradually weakened over subsequent years. The increasing size of rock lobster within reserves apparently reduced their dependency on reef shelters as refuges from predation. Rising predation by fish and rock lobster in the reserves corresponded with weakening invertebrate–habitat relationships for H. rubra and sea urchins (Heliocidaris erythrogramma). These results emphasise that animal–habitat relationships are not necessarily stable through time and highlight the value of marine reserves as reference sites. Our work shows that fishery closures to enhance populations of commercially important and keystone species should be in areas with a range of habitat features to accommodate shifting ecological requirements with ontogenesis.     

The effects of predator abundance and habitat structural complexity on survival of juvenile sea urchins

We studied the effect of the abundance of predatory fishes and structural complexity of algal assemblages on the survival of juveniles of the sea urchin Paracentrotus lividus on Mediterranean infralittoral rocky bottoms. Post-settlement juveniles (2–10 mm) were placed on four distinct natural substrates with increasing structural complexity (coralline barren, algal turf, erect fleshy algal assemblages and small crevices) inside and outside the Medes Islands Marine Reserve. Predation on these sea urchins increased at greater abundance of predatory fishes, and decreased with greater structural complexity. The refuge provided by structural complexity, however, decreased with increasing size of sea urchin recruits. Predation on the smallest post-settlers was carried out almost exclusively by small fishes (<20 cm), mainly the labrid Coris julis, while the dominant predator of larger juveniles was the sparid Diplodus sargus. Our results demonstrate the cascading effects caused by the prohibition of fishing in marine reserves, and highlight the potential role of small predatory fishes in the control of sea urchin populations.Communicated by O. Kinne, Oldendorf/Luhe     

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.     

Setting Realistic Recovery Targets for Two Interacting Endangered Species,Sea Otter and Northern Abalone

Failure to account for interactions between endangered species may lead to unexpected population dynamics, inefficient management strategies, waste of scarce resources, and, at worst, increased extinction risk. The importance of species interactions is undisputed, yet recovery targets generally do not account for such interactions. This shortcoming is a consequence of species‐centered legislation, but also of uncertainty surrounding the dynamics of species interactions and the complexity of modeling such interactions. The northern sea otter (Enhydra lutris kenyoni) and one of its preferred prey, northern abalone (Haliotis kamtschatkana), are endangered species for which recovery strategies have been developed without consideration of their strong predator–prey interactions. Using simulation‐based optimization procedures from artificial intelligence, namely reinforcement learning and stochastic dynamic programming, we combined sea otter and northern abalone population models with functional‐response models and examined how different management actions affect population dynamics and the likelihood of achieving recovery targets for each species through time. Recovery targets for these interacting species were difficult to achieve simultaneously in the absence of management. Although sea otters were predicted to recover, achieving abalone recovery targets failed even when threats to abalone such as predation and poaching were reduced. A management strategy entailing a 50% reduction in the poaching of northern abalone was a minimum requirement to reach short‐term recovery goals for northern abalone when sea otters were present. Removing sea otters had a marginally positive effect on the abalone population but only when we assumed a functional response with strong predation pressure. Our optimization method could be applied more generally to any interacting threatened or invasive species for which there are multiple conservation objectives. Definición de Metas de Recuperación Realistas para Dos Especies en Peligro Interactuantes, Enhydra lutris y Haliotis kamtschatkana     

Shark-inflicted injury frequencies, escape ability, and habitat use of green and loggerhead turtles

Interactions between large marine predators and their prey are difficult to observe and little is known about the risk of predation faced by sea turtles. The frequency of predator-inflicted injuries, however, has afforded insights into the predation risk faced by many taxa. We measured the frequency of shark-inflicted injuries on green (Chelonia mydas) and loggerhead (Caretta caretta) sea turtles in Shark Bay, Western Australia with a view to determining differences between species and sex-classes in the risk of predation from tiger sharks (Galeocerdo cuvier). Furthermore, we investigated how escape ability and habitat use might influence the probability of turtles being injured by sharks. Shark-inflicted injuries were more frequent on loggerhead than on green turtles, and most frequent on adult male loggerhead turtles. Species effects could not be attributed to differences in habitat use, since green turtles were found in habitats favored by tiger sharks more often than were loggerhead turtles. Green turtles, however, were faster and maneuvered better than loggerhead turtles, suggesting that escape ability is a factor in interspecific differences in injury frequency. The sex-class difference in injury frequency of loggerhead turtles suggests that males face greater predation risk than females and may take more risks. For green turtles, the lack of a sex difference in injury frequency might be due to greater escape ability lowering overall predation risk or to no differences between sexes in the benefits of risk-taking.     

Multivariate morphometric analysis of two varieties of Littorina saxatilis from the Swedish west coast

Shell variation in Littorina saxatilis Olivi (=L. rudis) is examined with different techniques of multivariate analysis (canonical variate-, multiple discriminant-, principal component-, and principal coordinate analysis). In bivariate projections of the analyses, two populations, spatially only 60 m apart, but from different habitats, showed a phenotypic separation without overlap. One habitat is a boulder shore, while the other is a rocky cliff. A sample, geographically and environmentally intermediate to the exposed and sheltered populations, covered completely the morphologic gap between the two former groups, indicating a clinal variation in morphology over the environmental gradient exposed — sheltered. From this it is concluded that the two phenotypically distinct populations belong to the same species.     

Induced defensive responses by the bivalve Mytilus edulis to predators with different attack modes

Predators such as crabs, whelks, and sea stars attack their bivalve prey in different ways, and predator-induced defenses are an important means of protection. The degree to which induced defenses are specific to different predators, however, remains largely unknown. In laboratory experiments (June to August 1998), we raised mussels (Mytilus edulis L.) in the presence of a drilling predator [the whelk Nucella lapillus (L.)] or a crushing predator [the crab Carcinus maenas (L.)] to determine whether induced changes in prey shell thickness, size, or shape occurred and whether changes were predator-specific. Over a 2 month period, juvenile mussels were exposed to waterborne cues from actively feeding crabs or whelks. Mussels produced thicker shell lips in response to both predators relative to control mussels raised in their absence, and the difference was significantly greater in response to whelks than to crabs. Mussels exposed to whelks showed significantly smaller increases in shell length and width and total wet weight than did mussels exposed to crabs. Thus, there may be a trade-off between shell thickness and linear shell growth and a potential delay in attaining a size refuge from predation. Received: 4 August 1999 / Accepted: 31 January 2000     

Pursuit deterrence,predation risk,and escape in the lizard <Emphasis Type="Italic">Callisaurus draconoides</Emphasis>

The frequency of pursuit-deterrent signaling should vary with predation risk for factors affecting decisions to flee and hide. Distance to refuge, temperature, and microhabitat types affect risk, and their effects may differ among defenses. Because risk is greater farther from refuge and at lower temperatures that impair escape ability in ectotherms, I predicted that when farther from refuge ectothermic prey flee sooner, enter refuge more frequently, and signal more frequently. At low temperatures, prey should flee sooner, enter refuge, and signal less frequently. Because signaling sometimes deters attack, lowered risk might allow prey to permit closer approach when signaling. In the zebra-tailed lizard Callisaurus draconoides, which signals by waving its tail, signaling and the other behaviors were affected by the same risk factors. Probability of signaling before fleeing increased with distance to refuge and temperature, but variation in temperature accounted for the effect of distance to refuge. Lizards signaled not at all at low temperatures, after starting to flee at intermediate temperatures, and before, during, and after fleeing at higher temperatures. This pattern is consistent with honest signaling of escape ability. Refuge entry was more frequent nearer refuge (lower risk) and at lower temperature (higher risk). Display was less frequent on rock than ground, possibly due to conspicuousness or delay in attaining high speed. Flight initiation distance was shorter after signaling. This novel finding suggests that signaling reduces risk and prey alter escape decisions based on lower risk as a consequence of their own signaling behavior.     

Effect of predation threat on repeatability of individual crab behavior revealed by mark-recapture

The persistence of behavioral types in situ and the drivers of persistence are central to predicting the ecological effects of intraspecific behavioral variation. We surveyed individual refuge use of mud crabs (Panopeus herbstii), a behavior related to the strength of a trait-mediated trophic cascade in oyster reefs, in the absence and presence of toadfish (Opsanus tau) predation threat. We then released these crabs into the field and using mark-recapture, measured the repeatability of this behavior in the absence and presence of threat, and how behavioral change was affected by time in the field (a month on average, up to 81 days), crab size, and sex. Because crabs exhibited some evidence of a circatidal rhythm in refuge use, we also tested how tidal height during observation influenced behavioral change. Predation threat increased refuge use, and small crabs used the refuge more than large crabs, particularly under threat. In recaptured crabs, refuge use was more repeatable under threat. Neither time in the field, crab size, crab sex, nor tidal height had any effect on behavioral change. Our results support the non-mutually exclusive hypotheses that (1) prey organisms in the presence, rather than absence, of predation threat should exhibit less behavioral variability because the fear of dying (a severe fitness consequence) should take precedence over less immediately important influences on behavior (e.g., hunger) and that (2) individual behaviors tied to fixed traits (e.g., the body size dependence of refuge use under threat in this study), rather than variable traits, should be more repeatable over time.     

Predators of juvenile sea urchins and the effect of habitat refuges

We evaluated the effects of potential predators from intertidal habitats on Strongylocentrotus purpuratus survival using laboratory experiments and assessed abundances of main predatory species along the Pacific coast of North America. The interactive effects of urchins’ and predators’ sizes in mediating predation were quantified. Habitat complexity (substrate pits, adult spine canopy) was manipulated to examine its effects on predation of most susceptible individuals (<14 mm). Pachygrapsus crassipes was identified as a major predator of urchins up to ≈30 mm. A positive effect of predator size on consumption of progressively larger urchins was detected, probably due to a mechanical limitation on crabs’ ability to consume large prey. Larger claws of males with respect to females of comparable sizes facilitated the handling of larger prey. Substrate refuges significantly reduced mortality on juvenile urchins. These results show that crab predation may be important in organizing intertidal communities, despite multiple ecological mechanisms promoting sea urchin survival.