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.     

Predation by gulls on crabs in rocky intertidal and shallow subtidal zones of the Gulf of Maine

Most organisms in intertidal areas are marine in origin; many have distributions that extend into the subtidal zone. Terrestrial predators such as mammals and birds may exploit these animals during low tide and can have considerable effects on intertidal food webs. Several studies have shown that avian predators are capable of reducing densities of sessile and slow-moving intertidal invertebrates but very few studies have considered avian predation on mobile invertebrate predators such as crabs. In this study, we investigated predation by Great Black-backed Gulls (Larus marinus Linnaeus) on three species of crabs (Cancer borealis Stimpson, Cancer irroratus Say, and Carcinus maenas Linnaeus). The study was at Appledore Island, ME (a gull breeding island) and 8 other sites throughout the Gulf of Maine, including breeding islands and mainland sites. On Appledore Island, intertidal and subtidal zones provided over one-third of prey remains found at gull nests, and crabs were a substantial proportion (∼ 30% to 40%) of the total remains. Similarly, collections of prey remains from intertidal areas indicated that crabs were by far the most common marine prey. C. borealis was eaten far more often and C. irroratus and C. maenas less often than expected at each site. Comparing numbers of carapaces to densities of crabs in low intertidal and shallow subtidal zones at each site, we estimated that gulls remove between 15% and 64% of C. borealis during diurnal low tides. The proportion of C. borealis eaten by gulls was independent of proximity to a gull colony. Approximately 97% of the outer coast of Maine is within 20 km of a breeding island. Thus, a lot of gull predation on crabs may occur throughout the Gulf of Maine during summer months. Crabs are important predators of other invertebrates; if predation by gulls reduces the number of crabs in intertidal and shallow subtidal areas, gulls may have important indirect effects on intertidal food webs.     

The Effect of Refuge on Dermestes ater (Coleoptera: Dermestidae) Predation on Musca domestica (Diptera: Muscidae): Refuge for Prey or the Predator?

It is believed that habitat heterogeneity can change the extent of predator-prey interactions. Therefore, in this study we examined the effect of habitat heterogeneity (characterized here as an addition of refuge) on D. ater predation on M. domestica. Predation of D. ater on M. domestica larvae was carried out in experimental habitats with and without refuge, and examined at different prey densities. The number of prey eaten by beetles over 24 h of predator-prey interaction was recorded, and we investigated the strength of interaction between prey and predator in both experimental habitats by determining predator functional response. The mean number of prey eaten by beetles in the presence of refuge was significantly higher than in the absence of refuge. Females had greater weight gains than males. Logistic regression analyses revealed the type II functional response for both experimental habitats, even though data did not fit well into the random predator model. Results suggest that the addition of refuge in fact enhanced predation, as prey consumption increased in the presence of refuge. Predators kept in the presence of refuge also consumed more prey at high prey densities. Thus, we concluded that the addition of refuge was an important component mediating D. ater-M. domestica population interactions. Refuge actually acted as a refuge for predators from prey, since prey behaviors detrimental to predators were reduced in this case.     

Within- and between-patch variability of predation intensity on the mussel Mytilus trossulus Gould on a rocky intertidal shore in Oregon, USA

Variability of predation intensity is an important cause of spatial differences of community structure and organization in the intertidal rocky shore. Field experiments were conducted to evaluate the within- and between-patch variability of the effects of different types of predators (small invertebrates and birds) on Mytilus trossulus Gould, which occupies an intermediate position in the competitive hierarchy among sessile organisms in disturbance patches within a California mussel (Mytilus californianus Conrad) bed community on the central Oregon Coast. Predation by birds did not significantly affect the mortality of M. trossulus. On the contrary, predation by small invertebrate whelks (Nucella spp.) had a significant effect on M. trossulus mortality. Predation by whelks also caused between- and within-patch variability of mortality of M. trossulus. Within patches, M. trossulus mortality at patch margins was significantly higher than at patch centers only when invertebrate predators were present. Wave exposure did not cause between-patch variability of predation intensity. Between-and within-patch variability of predation intensity may be caused by the variability of supply of whelks from the surrounding mussel mat. The movement of predators between patches and surrounding matrices may play an important role in the patch dynamics of these communities.     

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.     

Selection of food patches by sympatric herbivores in response to concealment and distance from a refuge

Small herbivores face risks of predation while foraging and are often forced to trade off food quality for safety. Life history, behaviour, and habitat of predator and prey can influence these trade‐offs. We compared how two sympatric rabbits (pygmy rabbit, Brachylagus idahoensis; mountain cottontail, Sylvilagus nuttallii) that differ in size, use of burrows, and habitat specialization in the sagebrush‐steppe of western North America respond to amount and orientation of concealment cover and proximity to burrow refuges when selecting food patches. We predicted that both rabbit species would prefer food patches that offered greater concealment and food patches that were closer to burrow refuges. However, because pygmy rabbits are small, obligate burrowers that are restricted to sagebrush habitats, we predicted that they would show stronger preferences for greater cover, orientation of concealment, and patches closer to burrow refuges. We offered two food patches to individuals of each species during three experiments that either varied in the amount of concealment cover, orientation of concealment cover, or distance from a burrow refuge. Both species preferred food patches that offered greater concealment, but pygmy rabbits generally preferred terrestrial and mountain cottontails preferred aerial concealment. Only pygmy rabbits preferred food patches closer to their burrow refuge. Different responses to concealment and proximity to burrow refuges by the two species likely reflect differences in perceived predation risks. Because terrestrial predators are able to dig for prey in burrows, animals like pygmy rabbits that rely on burrow refuges might select food patches based more on terrestrial concealment. In contrast, larger habitat generalists that do not rely on burrow refuges, like mountain cottontails, might trade off terrestrial concealment for visibility to detect approaching terrestrial predators. This study suggests that body size and evolutionary adaptations for using habitat, even in closely related species, might influence anti‐predator behaviors in prey species.     

Direct and indirect effects of predation on mosquitofish behavior and survival

Predation can have strong direct and indirect effects on the behavior of prey. We investigated whether predation by chain pickerel (Esox niger) caused adult eastern mosquitofish (Gambusia holbrooki) to alter their habitat use and whether pickerel predation influenced survival of adult and neonate mosquitofish. The number of adult mosquitofish using the riskier of three habitats was lowest when two predators occupied the risky habitat, intermediate in the treatment with one predator, and highest when no predators occurred there. More mosquitofish neonates survived high predation treatments than treatments lacking pickerel. We conclude that pickerel predation causes adult mosquitofish to shift to refuge habitats. The pattern of neonate survival suggests that adult habitat use may create a refuge from cannibalism for neonate mosquitofish, resulting in higher neonate survival in treatments with more pickerel. Hence, pickerel predation has a direct effect on adult mosquitofish behavior and a strong indirect effect on neonate survival. Both interspecific and intraspecific predation can effect prey populations and can interact to produce important indirect effects.     

A meta‐analysis of the effects of detritus on primary producers and consumers in marine,freshwater, and terrestrial ecosystems

Detritus is a central feature in marine, freshwater, and terrestrial ecosystems. Despite the ubiquity of detritus, ecologists have largely ignored its role in influencing food web structure. We used a meta‐analytic approach to ask three questions about how detritus affects food web structure in a wide variety of ecosystems. First, what is the effect strength of detritus on primary producers, detritivores, herbivores, and predators? Second, what functional role does detritus serve for consumers (energetic, habitat, or both)? Third, how does the effect of detritus on consumers vary between aquatic and terrestrial ecosystems? We found that detritus has strong positive effects on primary producers and consumers in a wide range of ecosystems types. Detritus has a positive direct effect on detritivores by providing both an energetic resource and habitat (refuge from predators). Detritus has equally strong positive effects on herbivores and predators, driven by a positive direct effect of habitat. Detritus has positive effects on consumers in both aquatic and terrestrial ecosystems with 1.7 times stronger effects in terrestrial ecosystems. These results suggest that detritus has strong effects on food‐web structure in a variety of ecosystem types. Even the portion of the food web that is linked most strongly to living plant tissue as its primary energy source is strongly positively affected.     

Functional roles of an engineer species for coastal benthic invertebrates and demersal fish

Through their tissues or activities, engineer species create, modify, or maintain habitats and alter the distribution and abundance of many plants and animals. This study investigates key ecological functions performed by an engineer species that colonizes coastal ecosystems. The gregarious tubiculous amphipod Haploops nirae is used as a biological model. According to previous studies, the habitat engineered by H. nirae (i.e., Haploops habitat) could provide food and natural shelter for several benthic species such as benthic diatoms belonging to the gender Navicula, the micrograzer Geitodoris planata, or the bivalve Polititapes virgineus. Using data from scientific surveys conducted in two bays, this study explored whether (1) the Haploops sandy‐mud community modifies invertebrate and ichthyologic community structure (diversity and biomass); (2) H. nirae creates a preferential feeding ground; and (3) this habitat serves as a refuge for juvenile fish. Available Benthic Energy Coefficients, coupled with more traditional diversity indices, indicated higher energy available in Haploops habitat than in two nearby habitats (i.e., Sternaspis scutata and Amphiura filiformis/Owenia fusiformis habitats). The use of isotopic functional indices (IFIs) indicated (1) a higher functional richness in the Haploops habitat, related to greater diversity in food sources and longer food chains; and (2) a higher functional divergence, associated with greater consumption of a secondary food source. At the invertebrate‐prey level, IFIs indicated little specialization and little trophic redundancy in the engineered habitat, as expected for homogenous habitats. Our results partly support empirical knowledge about engineered versus nonengineered habitats and also add new perspectives on habitat use by fish and invertebrate species. Our analyses validated the refuge‐area hypothesis for a few fish species. Although unique benthic prey assemblages are associated with Haploops habitat, the hypothesis that it is a preferential feeding area was not verified. However, specialist feeding behavior was observed for predators, which calls for further investigation.     

Habitat complexity and sex-dependent predation of mosquito larvae in containers

Studies in aquatic systems have shown that habitat complexity may provide refuge or reduce the number of encounters prey have with actively searching predators. For ambush predators, habitat complexity may enhance or have no effect on predation rates because it conceals predators, reduces prey detection by predators, or visually impairs both predators and prey. We investigated the effects of habitat complexity and predation by the ambush predators Toxorhynchites rutilus and Corethrella appendiculata on their mosquito prey Aedes albopictus and Ochlerotatus triseriatus in container analogs of treeholes. As in other ambush predator-prey systems, habitat complexity did not alter the effects of T. rutilus or C. appendiculata whose presence decreased prey survivorship, shortened development time, and increased adult size compared to treatments where predators were absent. Faster growth and larger size were due to predator-mediated release from competition among surviving prey. Male and female prey survivorship were similar in the absence of predators, however when predators were present, survivorship of both prey species was skewed in favor of males. We conclude that habitat complexity is relatively unimportant in shaping predator-prey interactions in this treehole community, where predation risk differs between prey sexes.     

Habitat formation prevails over predation in influencing fouling communities

Coastal human‐made structures, such as marinas and harbors, are expanding worldwide. Species assemblages described from these artificial habitats are novel relative to natural reefs, particularly in terms of the abundance of nonindigenous species (NIS). Although these fouling assemblages are clearly distinctive, the ecosystem functioning and species interactions taking place there are little understood. For instance, large predators may influence the fouling community development either directly (feeding on sessile fauna) or indirectly (feeding on small predators associated with these assemblages). In addition, by providing refuges, habitat complexity may modify the outcome of species interactions and the extent of biotic resistance (e.g., by increasing the abundance of niche‐specific competitors and predators of NIS). Using experimental settlement panels deployed in the field for 2.5 months, we tested the influence of predation (i.e., caging experiment), artificial structural complexity (i.e., mimics of turf‐forming species), and their interactions (i.e., refuge effects) on the development of sessile and mobile fauna in two marinas. In addition, we tested the role of biotic complexity—arising from the habitat‐forming species that grew on the panels during the trial—on the richness and abundance of mobile fauna. The effect of predation and artificial habitat complexity was negligible, regardless of assemblage status (i.e., native, cryptogenic, and nonindigenous). Conversely, habitat‐forming species and associated epibionts, responsible for biotic complexity, had a significant effect on mobile invertebrates (richness, abundance, and community structure). In particular, the richness and abundance of mobile NIS were positively affected by biotic complexity, with site‐dependent relationships. Altogether, our results indicate that biotic complexity prevails over artificial habitat complexity in determining the distribution of mobile species under low predation pressure. Facilitation of native and non‐native species thus seems to act upon diversity and community development: This process deserves further consideration in models of biotic resistance to invasion in urban marine habitats.     

Habitat selection by western snowy plovers during the nonbreeding season

Conservation of rare populations requires managing habitat throughout the year, especially during winter when northern populations may be limited by food and predation. Consequently, we examined distribution of nonbreeding western snowy plovers (Charadrius alexandrinus), including individually marked birds that were year-round residents and others that were migrants, in coastal northern California. Over 2 years, banded plovers exhibited high site faithfulness, occupying small linear stretches of beach (752 ± 626 m). Sites occupied by plovers had more brown algae (e.g., Macrocystis, Nereocystis, Postelsia, and Fucus) and associated invertebrates (e.g., amphipods, and flies), were wider, and had less vegetation than unoccupied sites. Our findings suggest that wintering plovers select habitats with more food and where they could more easily detect predators. Maintaining habitat with attributes that support abundant food (i.e., brown algae) and reduce predation risk (i.e., wide beaches, limited obstructive cover) may be important to individual survival and maintaining the Pacific Coast population of snowy plovers. Protecting occupied sites from human disturbance, which adversely alters nonbreeding habitat (i.e., beach grooming) and directly causes mortality, may be essential for conserving the Pacific coast population of the snowy plover, and it may benefit other shorebirds. © 2011 The Wildlife Society.     

Marine biofilm bacteria evade eukaryotic predation by targeted chemical defense

Many plants and animals are defended from predation or herbivory by inhibitory secondary metabolites, which in the marine environment are very common among sessile organisms. Among bacteria, where there is the greatest metabolic potential, little is known about chemical defenses against bacterivorous consumers. An emerging hypothesis is that sessile bacterial communities organized as biofilms serve as bacterial refuge from predation. By testing growth and survival of two common bacterivorous nanoflagellates, we find evidence that chemically mediated resistance against protozoan predators is common among biofilm populations in a diverse set of marine bacteria. Using bioassay-guided chemical and genetic analysis, we identified one of the most effective antiprotozoal compounds as violacein, an alkaloid that we demonstrate is produced predominately within biofilm cells. Nanomolar concentrations of violacein inhibit protozoan feeding by inducing a conserved eukaryotic cell death program. Such biofilm-specific chemical defenses could contribute to the successful persistence of biofilm bacteria in various environments and provide the ecological and evolutionary context for a number of eukaryote-targeting bacterial metabolites.     

Host-plant specialization by a non-herbivorous amphipod: advantages for the amphipod and costs for the seaweed

Studies of factors affecting host plant specialization by herbivores commonly highlight the value of the plant as both food and habitat, but often cannot distinguish the relative importance of these plant traits. A different approach is to study non-herbivorous animals that specialize on particular plants but do not feed on tissue from these plants. Such animals will not be affected directly by the nutritional, chemical, or morphological traits that determine the value of the plant as a food. This study reports on a filter-feeding amphipod, Ericthoniusbrasiliensis, that lives in domiciles it constructs by curling terminal segments of the green, calcified, and chemically defended seaweed Halimedatuna. We examined the temporal (1850s–1990s) and spatial (Caribbean, Mediterranean, and Pacific regions) scale of the association, the factors that may select for specialization on H. tuna, and the effect of the amphipod on growth of its host. Sampling along 125 km of coral reefs in the Florida Keys (USA) indicated that almost all populations of H. tuna had been colonized by this amphipod. Infested plants occurred on nine of ten reefs that supported H. tuna populations, with between 8 and 75% of the plants on those reefs colonized by the amphipod. For infested plants, 2–23% of all segments on each plant had been curled by the amphipod. Common co-occurring congeners of H. tuna (H. opuntia and H. goreaui) were never used for domicile construction. A survey of 1498 Halimeda specimens collected during the last 140 years and archived in the U.S. National Museum of Natural History (Smithsonian Institution, Washington, D.C.) indicated that the association has existed for >100 years and occurs throughout the Caribbean region, never in the Indo-Pacific or Mediterranean, and only on H. tuna. Predation by fishes could select for amphipod specialization on H. tuna. Laboratory experiments demonstrated that amphipods inhabiting curled segments of H. tuna were relatively immune from fish predation while those on the exterior surface of the plant or in open water were rapidly eaten. Segments of H. tuna are large enough to provide full protection from predators, while those of the co-occurring congeners H. goreaui and H. opuntia are of a size that may provide only partial protection. Experimental addition of E. brasiliensis to H. tuna plants in the field significantly decreased segment accumulation on infested relative to uninfested control plants. Whether this negative effect was a direct or indirect consequence of amphipod occupancy is unclear. Rolling plant portions into domiciles could directly decrease host growth by increasing shading and decreasing exposure of plant surface area to water column nutrient flux. Amphipod occupancy could indirectly slow net host growth if fishes selectively feed on plant sections occupied by amphipods. Underwater video showed that herbivorous fishes did not graze infested plants more than uninfested plants, but small predatory fishes did prefer feeding from infested plants. These non-herbivorous fishes may slow host growth by damaging the terminal meristematic tissues of plants during attacks on amphipods. This study demonstrates that habitat specialists can negatively impact hosts without consuming them and that specialization on a plant can occur due to its habitat value alone (as opposed to its value as a food). Received: 24 March 1998 / Accepted: 1 November 1998     

Chemical defence against predators in a sub-Arctic fjord

The development of chemical defences in marine organisms is supposed to be driven by intense pressure of predation and competition. While benthic communities in tropical and also Antarctic regions are thought to be mainly structured by intra- and interspecific interactions, these factors are proposed to be less important in northern high latitudes. Consequently, selective pressure for chemical defence should be low in these regions. To investigate the incidence of chemical defence against predation in northern high latitudes, crude extracts of 18 abundant sessile or slow-moving invertebrate species (4 sponges, 3 actinians, 1 soft coral, 4 bryozoans, 3 ascidians and the egg mass of a gastropod) from Kongsfjord (Spitsbergen) were tested for feeding deterrent activity. Laboratory assays were performed by offering artificial food with extracts to two different predators, the amphipod Anonyx nugax which is a common species in Kongsfjord, and the starfish Asterias rubens from the North Sea. Of the 18 extracts tested, 2 (Haliclona viscosa, Hormathia nodosa) exhibited significant feeding deterrent effects in the amphipod assay. Furthermore, 6 extracts had a significantly stimulating effect on the amphipod feeding, and 10 extracts did not affect consumption. In the starfish assay, only the crude extract of H. viscosa was significantly rejected. For H. viscosa, feeding deterrence could be established for two pure compounds, and for H. nodosa for one fraction. The present data show that feeding deterrent compounds are present in sub-Arctic marine invertebrates from Kongsfjord but are less abundant than in temperate, tropical and Antarctic species.     

Shell utilization,predation pressure,and thermal stress in panamanian hermit crabs: An interoceanic comparison

Shell preference patterns of two common hermit crabs from hard bottom reef flats on the Caribbean coast of Panama are examined in relation to the predation pressures and physical stresses of their habitat. Clibanarius antillensis Stimpson lives in the high intertidal habitat and minimizes exposure to predators by seeking refuge during high tides. It prefers high-spired shells which maximize protection from thermal stress. Calcinus tibicen Herbst avoids tidal emersion and prefers low-spired shells which enhance resistance to the predators common on Caribbean reef flats.The results are compared with similar results from the tropical eastern Pacific Bay of Panama. Shell-crushing predation on Caribbean hermit crabs is suggested to differ quantitatively and qualitatively from predation on hermit crabs in the Bay of Panama. Predation on hermit crabs in the Bay of Panama is more intense and effects larger individuals than predation on Caribbean reef flat hermit crabs. In addition, shell-crushing predation on hermit crabs in the Bay of Panama is primarily from teleost fish predators (Diodon spp.), while predation on Caribbean hermit crabs is primarily by bottom-dwelling crustaceans.Differences in predation pressures and tidal regimes between the Caribbean and Pacific coasts of Panama are reflected in the shell preferences and behavior of hermit crabs from the two areas.     

Predation risk and habitat complexity modify intermediate predator feeding rates and energetic efficiencies in a tri‐trophic system

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Coarse woody debris as a refuge from predation in aquatic communities

This study demonstrates experimentally that coarse woody debris (CWD) can provide refuge from predation in aquatic habitats. In the Rhode River subestuary of Chesapeake Bay, Maryland, (USA), we (1) measured the abundance of CWD, (2) examined the utilization of CWD by mobile epibenthic fish and crustaceans, and (3) tested experimentally the value of CWD as a refuge from predation. CWD was the dominant above-bottom physical structure in shallow water, ranging in size from small branches (<2 cm diameter) to fallen trees (>50 cm diameter). In response to experimental additions of CWD, densities of common epibenthic cpecies (Callinectes sapidus, Fundulus heteroclitus, Fundulus majalis, Gobiosoma bosc, Gobiesox strumosus, Palaemonetes pugio, and Rithropanopeus harrisii) increased significantly compared to control sites without CWD. In laboratory experiments, grass shrimp (P. pugio) responded to predatory fish (F. heteroclitus and Micropogonias undulatus) by utilizing shelter at CWD more frequently than in absence of fish. Access to CWD increased survivorship of grass shrimp in laboratory and field experiments. These experimental results (1) support the hypothesis, commonly proposed but untested for freshwater habitats, that CWD can provide a refuge from predation for epibenthic fish and invertebrates and (2) extend the recognized functional importance of CWD in freshwater to estuarine and marine communities. We hypothesize that CWD is an especially important refuge habitat in the many estuarine and freshwater systems for which alternative physical structure (e.g., vegetation or oyster reefs) are absent or in low abundance.     

Tube-building in two epifaunal amphipod species,Corophium insidiosum andJassa falcata

Jassa falcata andCorophium insidiosum are epifaunal tube-building marine amphipods, whose niches overlap in habitat and food requirements. Laboratory experiments were conducted to study the influence of the quality of different available particulate substrates on settlement and tube-building behaviour of these two amphipod species. Our experiments suggest thatC. insidiosum is less specialized in this respect thanJ. falcata. C. insidiosum is able to use organic materials for tube-building such as artificial (Ulva spec. powder) or mixed natural detritus as well as inorganic material (coarse sand); whereasJ. falcata utilizes organic materials, but sand only to a very limited extent.     

Feeding and Avoiding Predation Hazard: the Behavioral Response of the Prey

A forager may respond to the hazard of predation by (1) ignoring the hazard and eating maximally at all levels of food (risk-reckless); (2) avoiding the hazard and minimizing eating (risk-avoiding); (3) reducing feeding by the same amount at each level of food, making greater reductions as hazard increases (risk-adjusting); (4) reducing feeding at low levels of food, while accepting a greater hazard when food levels are high (risk-balancing). We experimentally distinguished these four nominal responses using 9 three-spined sticklebacks, Gasterosteus aculeatus in a 3 times 4 factorial, repeated measures design. The fish were tested in an aquarium containing a refuge inaccessible to predatory trout, Salmo trutta, in 12 combinations of food (low, medium, high) and predation hazard (predator density = 0, 1, 2, 3). With predators, sticklebacks ate less and used the refuge more. Amount eaten, but not time spent out of the refuge, was positively correlated with food, even in the absence of predators. Since sticklebacks as a group appeared to depress feeding by equal amounts at all levels of food, they employed a risk-adjusting response. However, individuals varied in their response to hazard.