Microhabitat use by a post-settlement stage estuarine fish: evidence from relative abundance and predation among habitats

Seagrass beds provide food and shelter for many fish species. However, the manner in which fishes use seagrass bed habitats often varies with life stage. Juvenile fishes can be especially dependent on seagrass beds because seagrass and associated habitats (drift macroalgae) may provide an effective tradeoff between shelter from predation and availability of prey. This study addressed aspects of habitat use by post-settlement pinfish, Lagodon rhomboides (Linneaus), an abundant and trophically important species in seagrass beds in the western North Atlantic and Gulf of Mexico. Abundance of post-settlement fish in seagrass beds was positively related to volume of drift macroalgae, but not to percent cover of seagrass, indicating a possible shelter advantage of the spatially complex algae. Tethering experiments indicated higher rates of predation in seagrass without drift macroalgae than in seagrass with drift macroalgae. Aquarium experiments showed lower predation with higher habitat complexity, but differences were only significant for the most extreme cases (unvegetated bottom, highest macrophyte cover). Levels of dissolved oxygen did not differ between vegetated and unvegetated habitats, indicating no physiological advantage for any habitat. Seagrass beds with drift macroalgae provide the most advantageous tradeoff between foraging and protection from predation for post-settlement L. rhomboides. The complex three-dimensional shelter of drift macroalgae provides an effective shelter that is embedded in the foraging habitat provided by seagrass. Drift macroalgae in seagrass beds is a beneficial habitat for post-settlement L. rhomboides by reducing the risk of predation, and by providing post-settlement habitat within the mosaic (seagrass beds) of adult habitat, thus reducing risks associated with ontogenetic habitat shifts.     

Differences in predator composition alter the direction of structure‐mediated predation risk in macrophyte communities

Structural complexity strongly influences the outcome of predator–prey interactions in benthic marine communities affecting both prey concealment and predator hunting efficacy. How habitat structure interacts with species‐specific differences in predatory style and antipredatory strategies may therefore be critical in determining higher trophic functions. We examined the role of structural complexity in mediating predator–prey interactions across several macrophyte habitats along a gradient of structural complexity in three different bioregions: western Mediterranean Sea (WMS), eastern Indian Ocean (EIO) and northern Gulf of Mexico (NGM). Using sea urchins as model prey, we measured survival rates of small (juveniles) and medium (young adults) size classes in different habitat zones: within the macrophyte habitat, along the edge and in bare sandy spaces. At each site we also measured structural variables and predator abundance. Generalised linear models identified biomass and predatory fish abundance as the main determinants of predation intensity but the efficiency of predation was also influenced by urchin size class. Interestingly though, the direction of structure‐mediated effects on predation risk was markedly different between habitats and bioregions. In WMS and NGM, where predation by roving fish was relatively high, structure served as a critical prey refuge, particularly for juvenile urchins. In contrast, in EIO, where roving fish predation was low, predation was generally higher inside structurally complex environments where sea stars were responsible for much of the predation. Larger prey were generally less affected by predation in all habitats, probably due to the absence of large predators. Overall, our results indicate that, while the structural complexity of habitats is critical in mediating predator–prey interactions, the direction of this mediation is strongly influenced by differences in predator composition. Whether the regional pool of predators is dominated by visual roving species or chemotactic benthic predators may determine if structure dampens or enhances the influence of top–down control in marine macrophyte communities.     

Structural complexity and fish body size interactively affect habitat optimality

Predator–prey interactions are strongly influenced by habitat structure, particularly in coastal marine habitats such as seagrasses in which structural complexity (SC) may vary over small spatial scales. For seagrass mesopredators such as juvenile fishes, optimality models predict that fitness will be maximized at levels of SC that enhance foraging but minimize predation risk, both of which are functions of body size. We tested the hypothesis that in eelgrass (Zostera marina) habitat, optimal SC for juvenile giant kelpfish (Heterostichus rostratus), an abundant eelgrass mesopredator in southern California, changes through ontogeny. To do this, we quantified eelgrass SC effects on habitat associations, relative predation risk, and foraging efficiency for three size classes of juvenile giant kelpfish. We found that habitat selection differed with fish size: small fish selected dense eelgrass, whereas larger fish selected sparse eelgrass. Small kelpfish experienced the lowest relative predation risk in dense eelgrass but also had higher foraging efficiency in dense eelgrass, suggesting that dense eelgrass is selected by these fish because it minimizes risk and maximizes potential for growth. Surprisingly, larger kelpfish did not experience lower predation risk than small kelpfish. However, larger kelpfish experienced higher foraging efficiency in sparse eelgrass vs. dense eelgrass, suggesting that they select sparse eelgrass to maximize foraging efficiency. Our study highlights that trade-offs between predation risk and foraging can occur within a single habitat type, that studies should consider how habitat value changes through ontogeny, and that seagrass habitat value may be maximal when within-patch variability in SC is high.     

Effects of <Emphasis Type="Italic">Caulerpa racemosa</Emphasis> var. <Emphasis Type="Italic">cylindracea</Emphasis> on prey availability: an experimental approach to predation of amphipods by <Emphasis Type="Italic">Thalassoma pavo</Emphasis> (Labridae)

Alien plant species, such as Caulerpa racemosa var. cylindracea, that invade Mediterranean marine vegetated habitats can affect habitat structure. In turn, changes in habitat structure may affect the associated invertebrate assemblages, either through changes in habitat selection or as a result of altered predation efficiency. In order to test for effects of changes in habitat structure resulting from colonization by C. racemosa on prey availability for predators, the importance of amphipods as a trophic resource in natural vegetated habitat was first assessed, and later experiments were undertaken to assess the effects of the alien alga on predation by Thalassoma pavo of two dominant amphipods: Elasmopus brasiliensis (Gammaridea) and Caprella dilatata (Caprellidea). Laboratory experiments were conducted in separate aquaria with five vegetation habitat types: Halopteris scoparia, Jania rubens, C. racemosa without detritus, C. racemosa with detritus, Cymodocea nodosa, together with controls. The vegetation was first defaunated, and then 30 amphipods were introduced to each aquarium and exposed to a single Thalassoma pavo individual for 1 h, after which the fish’s gut contents were examined. Consumption (per fish per hour) of caprellids (11.7 ± 1.4) was higher overall than that of gammarids (8.7 ± 1.5) and likely reflects different microhabitat use by amphipods, which affects susceptibility to predators. Consumption of amphipods also varied by habitat type. The highest predation rate was found in the C. nodosa habitat (12.7 ± 2.19) and the lowest in the C. racemosa habitats with detritus (4.1 ± 1.78) and without detritus (5.2 ± 0.55), which did not differ. The pattern of predation across habitats, however, was similar for both caprellid and gammarid amphipods, indicating a more general effect of habitat on amphipod predation. Our findings showed that invasive species such as C. racemosa can decrease feeding by predators such as T. pavo. Changes in predator–prey interactions could have consequences for food web support in the Mediterranean.     

Partitioning mechanisms of Predator Interference in different Habitats

Prey are often consumed by multiple predator species. Predation rates on shared prey species measured in isolation often do not combine additively due to interference or facilitation among the predator species. Furthermore, the strength of predator interactions and resulting prey mortality may change with habitat type. We experimentally examined predation on amphipods in rock and algal habitats by two species of intertidal crabs, Hemigrapsus sanguineus (top predators) and Carcinus maenas (intermediate predators). Algae provided a safer habitat for amphipods when they were exposed to only a single predator species. When both predator species were present, mortality of amphipods was less than additive in both habitats. However, amphipod mortality was reduced more in rock than algal habitat because intermediate predators were less protected in rock habitat and were increasingly targeted by omnivorous top predators. We found that prey mortality in general was reduced by (1) altered foraging behavior of intermediate predators in the presence of top predators, (2) top predators switching to foraging on intermediate predators rather than shared prey, and (3) density reduction of intermediate predators. The relative importance of these three mechanisms was the same in both habitats; however, the magnitude of each was greater in rock habitat. Our study demonstrates that the strength of specific mechanisms of interference between top and intermediate predators can be quantified but cautions that these results may be habitat specific. An erratum to this article can be found at     

Fish feeding guilds in shallow rocky and soft bottom areas on the Swedish west coast

Multivariate analyses of the diet relationships within fish assemblages resulted in the recognition of five major feeding guilds in shallow (0-9 m) rocky- and soft-bottom habitats on the Swedish west coast. The two habitats had similar guilds that could be broadly described as piscivores, decapod feeders, copepod feeders and amphipod and mollusc feeders. The last guild on the rocky bottom was more strictly one of amphipod feeders, whereas the last guild on soft bottoms consisted of infauna feeders (mainly polychaetes and bivalves). A comparison of the diet of the entire fish assemblages showed that amphipods and gastropods were more prevalent as prey taxa on rocky bottoms, whereas bivalves and teleosts were more common on soft bottoms. A large proportion (12 out of 27 species) of the fish species investigated was found in both habitats. These fish species generally seemed to rely on the same major prey groups in the two habitats, but also included prey taxa characteristic for the habitat in which they resided. Sea trout Salmo trutta , mackerel Scombrus scombrus , saithe Pollachius virens , whiting Merlangius merlangus and cod Gadus morhua were found to be the major piscivores in both the investigated habitats. Herring Clupea harengus and members of the family gobiidae were the most prevalent prey species for piscivores within the rocky- as well as the soft-bottom fish assemblage. A comparison of feeding modes suggested that the fish assemblage on rocky bottoms predominantly (60% of the food items) relied on food found in vegetation. On soft bottoms, the average distribution of food categories among fish species was 44% benthic, 35% pelagic and 21% vegetation-associated food items.     

Feeding ecology and growth of age 0 year Platichthys flesus(L.) in a vegetated and a bare sand habitat in a nutrient rich fjord

A study was carried out to investigate the diet and feeding strategies of age 0 year juvenile flounder Platichthys flesus in two different micro‐tidal habitats in the nutrient enriched Mariager Fjord on the Danish east coast. Juvenile flounder and benthic macrofauna were sampled monthly from June to October 1999 in a bare sand habitat and a habitat covered by filamentous and mat forming macroalgae. The presence of the 'opportunistic' macroalgae created a shift in the dominance of surface dwelling prey such as epifaunal amphipods to more infaunal groups such as oligochaetes and polychaetes. The diet of the flounder varied considerably between the two habitats mainly reflecting prey availability relative to their abundance, prey spatial distribution, habitat structure and ontogenetic prey shifts as a function of total length. Flounder in the vegetated site fed on a diverse diet of copepods, polychaetes and oligochaetes, whereas those caught in the bare sand site fed primarily on the amphipod Corophium volutator which was numerically dominant at this site. During the growth season, two diet shifts were observed: from copepods early in the season to macrofauna organisms and, later in the season, the inclusion of more hyperbenthic prey such as Mysidea spp; Idotea spp. and the common goby Pomatoschistus microps .     

Habitat complexity dampens selection on prey activity level

Conspecific prey individuals often exhibit persistent differences in behavior (i.e., animal personality) and consequently vary in their susceptibility to predation. How this form of selection varies across environmental contexts is essential to predicting ecological and evolutionary dynamics, yet remains currently unresolved. Here, we use three separate predator–prey systems (sea star–snail, wolf spider–cricket, and jumping spider–cricket) to independently examine how habitat structural complexity influences the selection that predators impose on prey behavioral types. Prior to conducting staged predator–prey interaction encounters, we ran prey individuals through multiple behavioral assays to determine their average activity level. We then allowed individual predators to interact with groups of prey in either open or structurally complex habitats and recorded the number and individual identity of prey that were eaten. Habitat complexity had no effect on overall predation rates in any of the three predator–prey systems. Despite this, we detected a pervasive interaction between habitat structure and individual prey activity level in determining individual prey survival. In open habitats, all predators imposed strong selection on prey behavioral types: sea stars preferentially consumed sedentary snails, while spiders preferentially consumed active crickets. Habitat complexity dampened selection within all three systems, equalizing the predation risk that active and sedentary prey faced. These findings suggest a general effect of habitat complexity that reduces the importance of prey activity level in determining individual predation risk. We reason this occurs because activity level (i.e., movement) is paramount in determining risk within open environments, whereas in complex habitats, other behavioral traits (e.g., escape ability to a refuge) may take precedence.     

Vulnerability of newly settled plaice (Pleuronectes platessa L.) to predation: effects of habitat structure and predator functional response

Plaice (Pleuronectes platessa) nursery grounds on the Swedish west coast have been subject to increasing cover of annual green macroalgae during recent years, with growth of algae starting at the time of plaice settlement in April to May. A laboratory experiment was performed to investigate how the vulnerability to predation of metamorphosing plaice was affected by the presence of filamentous algae. Predation by shrimps (Crangon crangon) on settling plaice larvae was higher on sand than among algae, whereas predation by crabs (Carcinus maenas) was unaffected by habitat type, suggesting a lower overall mortality of plaice in the vegetated habitat. When predators and prey were presented with a combination of the two habitats, predation by shrimps was as high as that in the sand treatment alone, whereas predation by crabs was lower than that in the two treatments with one habitat. Based on these results, an additional experiment was performed, investigating the functional response of shrimps to six densities of juvenile plaice in a sand habitat with alternative prey present. The proportional mortality of juvenile plaice (12-16 mm total length (TL)) was density-dependent and was best described by a type III (sigmoid) functional response of the predatory shrimps. The results suggested that the combined predation pressure from shrimps and crabs was lower among algae than on sand, but settling plaice and predatory shrimps chose the sand habitat. Plaice densities in the sigmoid part of the obtained functional response curve represented normal to high field densities of plaice on the Swedish west coast, suggesting that shrimp predation could have a stabilising effect on plaice recruitment. The formation of macroalgae mats could therefore lead to a concentration of plaice juveniles in the remaining sand habitat and increased mortality through density-dependent predation by shrimps.     

Epibenthic predators control mobile macrofauna associated with a foundation species in a subarctic subtidal community

Foundation species (FS) are strong facilitators providing habitat for numerous dependent organisms. The communities shaped by FS are commonly structured by interplay of facilitation and consumer control. Predators or grazers often indirectly determine community structure eliminating either FS or their principal competitors. Alternatively, they can prey on the dependent taxa directly, which is generally buffered by FS via forming complex habitats with numerous refuges. The latter case has been never investigated at high latitudes, where consumer control is widely considered weak. We manipulated the presence of common epibenthic crustacean predators to assess their effect on mobile macrofauna of the clusters developed by a FS (barnacle Balanus crenatus and its empty tests) in the White Sea shallow subtidal (65° N). While predation pressure on the FS itself here is low, the direct effects of a spider crab Hyas araneus and a shrimp Spirontocaris phippsii on the associated assemblages were unexpectedly strong. Removing the predators did not change species diversity, but tripled total abundance and altered multivariate community structure specifically increasing the numbers of amphipods, isopods (only affected by shrimp), and bivalves. Consumer control in the communities shaped by FS may not strictly follow the latitudinal predation gradient rule.     

The effects of seafloor habitat complexity on survival of juvenile fishes: Species-specific interactions with structural refuge

Marine fishes are often associated with structurally complex microhabitats that are believed to provide a refuge from predation. However, the effects of habitat complexity on predator foraging success can be strongly modified by predator and prey behaviors. We conducted a series of laboratory experiments to evaluate the effects of sea floor habitat complexity on juvenile fish survivorship using multiple predator (striped searobin and summer flounder) and prey (winter flounder, scup, and black sea bass) species to identify potentially important species-habitat interactions. Three habitats of varying complexity (bare sand, shell, and sponge) common to coastal marine environments were simulated in large aquaria (2.4 m diameter, 2400 L volume). Prey survivorship increased significantly with greater habitat complexity for each species combination tested. However, examination of multiple prey and predator species across habitats revealed important effects of predator × habitat and prey × habitat interactions on prey survival, which appeared to be related to species-specific predator and prey behavior in complex habitats. Significant species × habitat interactions imply that the impact of reduced seafloor habitat complexity may be more severe for some species than others. Our results indicate that the general effects of seafloor habitat complexity on juvenile fish survivorship may be broadly applicable, but that the interaction of particular habitats with search tactics of predators as well as habitat affinities and avoidance responses of prey can produce differences among species that contribute to variable mortality.     

Intertidal community structure differs significantly between substrates dominated by native eelgrass (<Emphasis Type="Italic">Zostera marina</Emphasis> L.) and adjacent to the introduced oyster <Emphasis Type="Italic">Crassostrea gigas</Emphasis> (Thunberg) in British Columbia,Canada

Eelgrass beds represent important habitats for marine organisms, but are in decline in many coastal areas around the world. On Cortes Island, British Columbia, Canada, oysters coexist regionally with native eelgrass (Zostera marina L.), but eelgrass is typically absent directly seaward of oyster beds (the “below-oyster cobble zone”). We compared assemblage structure of nekton (fish and swimming macroinvertebrates) and epibenthos (macroinvertebrates and macroalgae) between eelgrass bed and below-oyster habitats. We sampled the intertidal zone on Cortes Island at low tide using two methods: quadrats to enumerate epibenthic macroinvertebrates and macroalgae, and beach seines to enumerate fish and swimming macroinvertebrates. Using multivariate analysis of similarity (ANOSIM), we found that the structure of nektonic and epibenthic assemblages associated with below-oyster cobble zones were significantly different from those in eelgrass-beds. Univariate measures showed that nektonic species richness and abundance were significantly higher in eelgrass beds than in below-oyster cobble habitat, whereas epibenthic species richness and abundance were significantly higher in below-oyster habitat. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users. Handling editor: J. Trexler     

Effects of habitat complexity, prey type, and abundance on intraguild predation between larval odonates

Intraguild predation is an important interaction in which predators feed on a shared prey as well as on each other. It occurs frequently between larval odonates in freshwater lentic communities, and understanding the factors influencing this interaction remains an important objective. An experiment carried out in mesocosms and utilizing a factorial design investigated the strength of intraguild interactions between the dragonfly, Sympetrum vicinum, and the damselfly, Enallagma civile, under two levels each of habitat complexity (high or low), prey abundance (high or low) and prey type (amphipods or blackworms). Effects of treatments on size, mortality and emergence of larval odonates were evaluated. Shared prey abundance had little impact on intraguild interactions, affecting only the mass of the intraguild prey E. civile. Habitat complexity affected the size of E. civile damselflies, as length and wet mass were significantly greater in low complexity mesocosms. Prey type seemed to be the most important factor in the experiment, influencing all response variables measured. When shared prey consisted of larger, more active blackworms, intraguild predation decreased, and E. civile damselflies experienced lower mortality, achieved greater length and mass, and had greater emergence success. Results of this study suggest that prey type and habitat complexity can be more important than prey abundance in mediating intraguild predation.     

The effect of habitat structure on prey mortality depends on predator and prey microhabitat use

Structurally complex habitats provide cover and may hinder the movement of animals. In predator–prey relationships, habitat structure can decrease predation risk when it provides refuges for prey or hinders foraging activity of predators. However, it may also provide shelter, supporting structures and perches for sit-and-wait predators and hence increase their predation rates. We tested the effect of habitat structure on prey mortality in aquatic invertebrates in short-term laboratory predation trials that differed in the presence or absence of artificial vegetation. The effect of habitat structure on prey mortality was context dependent as it changed with predator and prey microhabitat use. Specifically, we observed an ‘anti-refuge’ effect of added vegetation: phytophilous predators that perched on the plants imposed higher predation pressure on planktonic prey, while mortality of benthic prey decreased. Predation by benthic and planktonic predators on either type of prey remained unaffected by the presence of vegetation. Our results show that the effects of habitat structure on predator–prey interactions are more complex than simply providing prey refuges or cover for predators. Such context-specific effects of habitat complexity may alter the coupling of different parts of the ecosystem, such as pelagic and benthic habitats, and ultimately affect food web stability through cascading effects on individual life histories and trophic link strengths.     

Tradeoffs involved in site selection and foraging in a wolf spider: effects of substrate structure and predation risk

Understanding how animals weigh habitat features, exposure to predators and access to resources is important to determining their life history and distribution across the landscape. For example, when predators accumulate in structurally complex habitats, they face an environment with different competitive interactions, foraging opportunities and predatory risks. The wolf spider Pardosa milvina inhabits the soil surface of highly disturbed habitats such as agricultural fields throughout eastern North America. Pardosa displays effective antipredator behavior in the presence of chemical cues produced by a larger coexisting wolf spider, Hogna helluo . We used those cues to simulate predation risk in laboratory and field experiments designed to test the effects of habitat substrate and predation risk on site selection and prey consumption of Pardosa . In general, Pardosa preferred more complex substrates over bare dirt but those preferences were eliminated or reversed when cues from Hogna were present. Feeding trials revealed that substrate alone had few effects on Pardosa prey consumption, which we measured by documenting the change in the abdomen width. Although the presence of Hogna cues reduced prey consumption overall in field feeding trials, the negative effect of predation risk on prey consumption was only observed in grass and bare dirt substrates in the laboratory. We also found that prey capture was negatively affected by habitat complexity for both spider species but that same complexity offered Pardosa protection from predation by Hogna. This study provides insight into how two predator species interact to balance site selection and feeding in order to avoid predation. Shifts in foraging and distributional patterns of predators can have profound implications for their role in the food web.     

Occupation of submerged aquatic vegetation by fishes: testing the roles of food and refuge

Summary We conducted a series of field experiments to examine the roles of refuge and food availability in explaining the distribution and abundance of fish in tidal freshwater marsh creeks. Two hypotheses were tested: (1) relative predation pressure is less in SAV than in unvegetated areas and (2) fish food availability is greater in SAV than in nearby unvegetated areas. Tethering experiments using mummichogs (Fundulus heteroclitus) in vegetated and unvegetated areas revealed that relative predation pressure was significantly less in areas with SAV. Banded killifish (Fundulus diaphanus) maintained in vegetated enclosures consumed prey associated with SAV, whereas those held in unvegetated pens had empty stomachs. No differences were found in the number of prey eaten by bluespotted sunfish (Enneacanthus gloriosus) or mummichogs when confined in vegetated or unvegetated enclosures. However, larger prey were consumed by bluespotted sunfish and mummichogs maintained in vegetated enclosures. These data suggest that foraging profitability is significantly enhanced by feeding in the SAV. Submerged plant beds in tidal freshwater marsh creeks not only afford protection from predators, but also provide a rich foraging habitat. By foraging in SAV, fish consume larger prey and may have higher growth rates, lower mortality, and higher fecundity.     

Heterogeneous landscapes and the role of refuge on the population dynamics of a specialist predator and its prey

How, and where, a prey species survives predation by a specialist predator during low phases of population fluctuations or a cycle, and how the increase phase of prey population is initiated, are much-debated questions in population and theoretical ecology. The persistence of the prey species could be due mainly to habitats that act as refuges from predation and/or due to anti-predatory behaviour of individuals. We present models for the former conjecture in two (and three) habitat systems with a specialist predator and its favoured prey. The model is based on dispersal of prey between habitats with high reproductive output but high risk of predation, and less productive habitats with relatively low risk of predation. We illustrate the predictions of our model using parameters from one of the most intriguing vertebrate predator–prey systems, the multi-annual population cycles of boreal voles and their predators. We suggest that cyclic population dynamics could result from a sequence of extinction and re–colonization events. Field voles (Microtus agrestis), a key vole species in the system, can be hunted to extinction in their preferred meadow habitat, but persist in sub-optimal wet habitats where their main predator, the least weasel (Mustela nivalis nivalis) has a low hunting efficiency. Re–colonization of favourable habitats would occur after the predator population crashes. At the local scale, the model suggests that the periodicity and amplitude of population cycles can be strongly influenced by the relative availability of risky and safe habitats for the prey. Furthermore, factors like intra-guild predation may lead to reduced predation pressure on field voles in sub-optimal habitats, which would act as a refuge for voles during the low phase of their population cycles. Elasticity analysis suggested that our model is quite robust to changes in most parameters but sensitive to changes in the population dynamics of field voles in the optimal grassland habitat, and to the maximum predation rate of weasels.     

Spatial refuge from intraguild predation: implications for prey suppression and trophic cascades

The ability of predators to elicit a trophic cascade with positive impacts on primary productivity may depend on the complexity of the habitat where the players interact. In structurally-simple habitats, trophic interactions among predators, such as intraguild predation, can diminish the cascading effects of a predator community on herbivore suppression and plant biomass. However, complex habitats may provide a spatial refuge for predators from intraguild predation, enhance the collective ability of multiple predator species to limit herbivore populations, and thus increase the overall strength of a trophic cascade on plant productivity. Using the community of terrestrial arthropods inhabiting Atlantic coastal salt marshes, this study examined the impact of predation by an assemblage of predators containing Pardosa wolf spiders, Grammonota web-building spiders, and Tytthus mirid bugs on herbivore populations (Prokelisia planthoppers) and on the biomass of Spartina cordgrass in simple (thatch-free) and complex (thatch-rich) vegetation. We found that complex-structured habitats enhanced planthopper suppression by the predator assemblage because habitats with thatch provided a refuge for predators from intraguild predation including cannibalism. The ultimate result of reduced antagonistic interactions among predator species and increased prey suppression was enhanced conductance of predator effects through the food web to positively impact primary producers. Behavioral observations in the laboratory confirmed that intraguild predation occurred in the simple, thatch-free habitat, and that the encounter and capture rates of intraguild prey by intraguild predators was diminished in the presence of thatch. On the other hand, there was no effect of thatch on the encounter and capture rates of herbivores by predators. The differential impact of thatch on the susceptibility of intraguild and herbivorous prey resulted in enhanced top-down effects in the thatch-rich habitat. Therefore, changes in habitat complexity can enhance trophic cascades by predator communities and positively impact productivity by moderating negative interactions among predators.     

Marine reserves indirectly affect fine‐scale habitat associations,but not overall densities,of small benthic fishes

Many large, fishery‐targeted predatory species have attained very high relative densities as a direct result of protection by no‐take marine reserves. Indirect effects, via interactions with targeted species, may also occur for species that are not themselves targeted by fishing. In some temperate rocky reef ecosystems, indirect effects have caused profound changes in community structure, notably the restoration of predator–urchin–macroalgae trophic cascades. Yet, indirect effects on small benthic reef fishes remain poorly understood, perhaps because of behavioral associations with complex, refuge‐providing habitats. Few, if any, studies have evaluated any potential effects of marine reserves on habitat associations in small benthic fishes. We surveyed densities of small benthic fishes, including some endemic species of triplefin (Tripterygiidae), along with fine‐scale habitat features in kelp forests on rocky reefs in and around multiple marine reserves in northern New Zealand over 3 years. Bayesian generalized linear mixed models were used to evaluate evidence for (1) main effects of marine reserve protection, (2) associations with habitat gradients, including complexity, and (3) differences in habitat associations inside versus outside reserves. No evidence of overall main effects of marine reserves on species richness or densities of fishes was found. Both richness and densities showed strong associations with gradients in habitat features, particularly habitat complexity. In addition, some species exhibited reserve‐by‐habitat interactions, having different associations with habitat gradients inside versus outside marine reserves. Two species (Ruanoho whero and Forsterygion flavonigrum) showed stronger positive associations with habitat complexity inside reserves. These results are consistent with the presence of a behavioral risk effect, whereby prey fishes are more strongly attracted to habitats that provide refuge from predation in areas where predators are more abundant. This work highlights the importance of habitat structure and the potential for fishing to affect behavioral interactions and the interspecific dynamic attributes of community structure beyond simple predator–prey consumption and archetypal trophic cascades.     

Predation rates of nemertean predators: the case of a rocky shore hoplonemertean feeding on amphipods

Estimates of the predation rates of benthic nemerteans are often based on observations of single individuals, and consequently they may not be representative for all members of a population of these predators. Herein we conducted controlled and repeatable laboratory experiments on the predation rate of the hoplonemertean Amphiporus nelsoni Sánchez 1973, which is common at exposed rocky shores along the central Chilean coast. During the austral fall (April, May 2000), nemerteans were observed in relatively high numbers crawling in the intertidal zone during early morning or late-afternoon low tides. When these nemerteans were offered living amphipods held by a forceps, they immediately attacked the amphipods and fed on them. In the laboratory experiments, nemerteans preferred the amphipod Hyale maroubrae Stebbing, 1899, which is also very common in the natural habitat of A. nelsoni. The nemerteans preyed to a higher extent on small males and non-ovigerous females than would have been expected from their abundance. We suggest that these (non-reproductive) stages of H. maroubrae are very mobile and therefore have a high likelihood of encounters with nemerteans. Predation rates reached maxima when nemerteans were provided prey densities of four or more of their preferred prey species, H. maroubrae, furthermore indicating that encounter rates with prey may affect predation rates. In long-term laboratory experiments, A. nelsoni consumed more amphipods during low tide conditions than during high tide conditions. Many nemerteans in the field prefer particular environmental conditions (e.g. nocturnal low tides), which restricts the time available for successful feeding. In the long-term experiment, predation rates of A. nelsoni never exceeded 0.5 amphipods nemertean^–1 d^–1. Maximum feeding events were 3 or 4 amphipods nemertean^–1 d^–1, but this only occurred during 10 out of a possible 2634 occasions. Nemerteans that had consumed 3 or 4 amphipods during 1 day, consumed substantially less prey during the following days. Towards the end of the long-term experiment, average predation rates decreased to 0.2 amphipods nemertean^–1 d^–1, corresponding to predation rates reported for other nemertean species (0.1–0.3 prey items nemertean^–1 d^–1). We suggest that predation rates from laboratory experiments represent maximum estimates that may not be directly transferable to field populations. Additionally, low predator–prey encounter rates with preferred prey in the field may further limit the predation impact of nemertean predators in natural habitats.